Leptynoptera sulfurea, a psyllid species that feeds on Calophyllum inophyllum trees, has been found for the first time on the Cocos (Keeling) Islands. L. sulfurea specimens were collected on five islands and were associated with C. inophyllum, which is found throughout the tropical Indian and Pacific Oceans. This expands the known range of L. sulfurea, which has been reported from numerous locations in Asia and the Pacific and feeds exclusively on C. inophyllum.

1. First record of the psyllid Leptynoptera sulfurea Crawford
(Hemiptera: Psylloidea: Triozidae) from the Cocos (Keeling) Islands
Peter J. Neville a
, Daniel Burckhardt b
, Alan L. Yen c,d,
⁎
a
Mosquito-Borne Disease Control, Environmental Health Hazards Unit, Department of Health, P.O. Box 8172 Perth Business Centre, Perth, Western Australia 6849, Australia
b
Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Switzerland
c
Biosciences Research Branch, Department of Economic Development, Jobs, Transport & Resources, La Trobe University, Bundoora, AgriBio, 5 Ring Road, Melbourne Victoria 3083, Australia
d
School of Applied Systems Biology, La Trobe University, Bundoora, Melbourne, Victoria 3086, Australia
a b s t r a c ta r t i c l e i n f o
Article history:
Received 2 November 2014
Revised 5 June 2015
Accepted 27 June 2015
Available online 3 July 2015
Keywords:
Leptynoptera sulfurea
Psyllid
Cocos (Keeling) Islands
The psyllid Leptynoptera sulfurea Crawford (Hemiptera: Psylloidea: Triozidae) is reported for the first time from
the Cocos (Keeling) Islands located in the Indian Ocean. It is associated with the tree Calophyllum inophyllum L.
(Calophyllaceae). It is the only species of psyllid currently known from these islands.
Crown Copyright © 2015 Published by Elsevier B.V. on behalf of Korean Society of Applied Entomology, Taiwan
Entomological Society and Malaysian Plant Protection Society. All rights reserved.
Introduction
Leptynoptera sulfurea Crawford is a species of psyllid associated with
Calophyllum inophyllum L., an evergreen tree widely dispersed through-
out the tropical regions of the Indian and Pacific Oceans. It was originally
described by Crawford (1919) when he erected the genus Leptynoptera
and described the species L. sulfurea from Amboina (Ambon Island) in
the Moluccas Islands (Maluku Islands), Indonesia. Burckhardt and
Ouvrard (2012) confirmed its placement in the psylloid family Triozidae.
Since its initial description, L. sulfurea has been recorded from several lo-
cations, mainly in the Asian region and on islands in the Pacific Ocean. It
is monophagous to C. inophyllum, and most of the locations are coastal.
This paper records the occurrence of L. sulfurea in the Cocos (Keeling)
Islands, a new location for this species.
Materials and methods
The Cocos (Keeling) Islands are an Australian Territory located in the
northeastern Indian Ocean, 2,400 km northwest of the North West Cape
in Australia, 1,300 km southwest of Java and 980 km west of Christmas
Island. They are located in a humid tropical zone, with predominantly
South East Trade Winds, and occasional cyclones. It is comprised of
22 vegetated islands, with the main ones (Direction, Home, Horsburgh,
South and West Islands) located in the South Keeling chain, and the
northern part only has North Keeling Island. The islands have had
human settlement for over 100 years and consequently the vegetation
is a mixture of native and introduced plants.
A general invertebrate survey of the Cocos (Keeling) atoll was con-
ducted from 1 to 24 June 2005 to inventory the fauna (Neville et al.,
2008). Due to difficulties reaching some of the islands, inclement
weather and inability to transport equipment, the survey focused on
three islands: West Island (Pulu Panjang), Direction Island (Pulu
Tikus) and Home Island (Pulu Selma), with opportunistic sampling
of additional islands as permitted: Horsburgh Island (Pulu Luar) and
North Keeling Islands.
Quantitative invertebrates sampling involved setting up transects
in the dominant vegetation types, coconut (Cocos nucifera), cabbage
bush (Scaevola taccada) and grassland habitats, on West, Home and
Direction Islands. Test tube pitfall traps (1.8 cm diameter), yellow
pan traps (12.5 × 20 × 7.5 cm), sweep netting (45 cm diameter) and
direct searches were conducted along the transects. Direct sampling
was conducted on the dominant plant species near each transect and
aimed to collect a representative sample of the invertebrate fauna
seen within 10 min. Samples were collected by scanning the upper
and lower side of foliage, bark, branches and trunks. Tree and shrub
species sampled included pisonia (Pisonia grandis R.Br.: Nyctaglinaceae),
cabbage bush (S. taccada (Gaertn.): Goodeniaceae), octopus bush
(Heliotropium foetherianum Diane & Hilger: Boraginaceae), coconut
(C. nucifera L. Arecaceae), Alexandrian laurel (Callophylum inophyllum
L.: Callophyllaceae), tea shrub (Pemphis acidula J.R. Forst. & G. Forst.:
Journal of Asia-Pacific Entomology 18 (2015) 497–499
⁎ Corresponding author at: Biosciences Research Branch, Department of Economic
Development, Jobs, Transport & Resources, AgriBio, 5 Ring Road, La Trobe University,
Bundoora, Victoria 3083, Australia. Tel.: +61 3 90327351.
E-mail addresses: Peter.neville@health.wa.gov.au (P.J. Neville),
daniel.burckhardt@bs.ch (D. Burckhardt), alan.yen@ecodev.vic.gov.au (A.L. Yen).
http://dx.doi.org/10.1016/j.aspen.2015.06.008
1226-8615/Crown Copyright © 2015 Published by Elsevier B.V. on behalf of Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection
Society. All rights reserved.
Contents lists available at ScienceDirect
Journal of Asia-Pacific Entomology
journal homepage: www.elsevier.com/locate/jape

2. Lythraceae), fish plate shrub (Guettarda speciosa L.: Rubiaceae), sea
perch (Terminalia catappa L.: Combretaceae), hopbush (Dodonaea
viscosa Jacq.: Sapindaceae), cheesefruit (Morinda citrifolia L.: Rubiaceae),
sea hibiscus (Hibiscus tilaceus L.: Malvaceae) and kerosene wood (Cordia
subcordata Lam.: Boraginaceae) (Neville et al., 2007).
Results and discussion
Specimens of L. sulfurea were collected on five of the islands in the
Cocos (Keeling) Islands during the survey: Direction Island: 1♀ found
through direct searching of C. inophyllum at the Jetty (12°05′22.5″S
96°53′02.3″E) on 4 June 2005; Home Island: 3♂ in two yellow pan
traps (run 13–17 June 2005) in grasslands (12°07′06.9″S 96°54′03.5″E),
36♂, 32♀ in three yellow pan traps (run 13–17 June 2005) at Oceania
House (12°07′14.1″S 96°53′40.0″E), 1♀ found through direct searching
of Scaevola sp. (7 June 2005) north of the Jetty (12°06′57.9″S 96°53′
37.3″E), 5 immatures found through direct searching of C. inophyllum
(7 June 2005) north of the Jetty (12°06′57.9″S 96°53′37.3″E), and 1 im-
mature found through direct searching of the litter (7 June 2005) north
of the Jetty (12°06′57.9″S 96°53′37.3″E); Horsburgh Island: 1♂1♀
found through direct searching of C. inophyllum at Possession Point
(19 June 2005); North Keeling Island: 1♂ from yellow pan trap (run
20–22 June 2005) in Pisonia forest (11°49′52.4″S 96°49′05.3″E); and
West Island: 5♂8♀ found through direct searching of C. inophyllum
and 1♂ from Scaevola sp. at the Trannies (12°08′30.7″S 96°49′03.8″E)
on 8 June 2005.
Currently, it is assumed that the psyllid immatures collected are
L. sulfurea because no other psyllid species have been recorded in the
past or were collected during this survey. Gibson-Hill documented the
non-marine invertebrates that he collected on the Cocos (Keeling)
Islands in a series of papers published in 1950. He listed 15 species of
Hemiptera but there were no psyllids in this list (Gibson-Hill, 1950).
Crawford (1919) erected the genus Leptynoptera and described the
species L. sulfurea based on one damaged specimen from Amboina
(Ambon Island) in the Moluccas Islands (Maluku Islands), Indonesia.
Uichanco (1919) published a photograph of marginal leaf galls on
C. inophyllum from Luzon (Philippines). Uichanco (1919) stated that
adult psyllids were collected at the time but they were not identified
to species. Later, Uichanco (1921) described these psyllids as a new va-
riety of L. sulfurea rubrocincta and provided illustrations of the forewing,
and the male and female genitalia. Takahashi (1927) mentions this sub-
species from Formosa (Taiwan), and Boselli (1930) lists and illustrates
specimens collected on Ficus by R. Takahashi in Formosa (Taiwan).
Laing (1922) originally described Leptynoptera didactyla from specimens
collected on C. inophyllum in Fiji. Caldwell (1942) made Leptynoptera
didactyla a synonym of L. sulfurea. In the same paper, Laing (1922) incor-
rectly spelt L. sulfurea as L. sulphurea. This mistake was corrected by
Martin and Hollis (1992).
L. sulfurea is mentioned in several papers other than the ones
already mentioned. Kuwayama (1931) lists L. sulfurea from Moluccas,
Philippines and Taiwan on C. inophyllum but from published infor-
mation (Crawford, 1919; Uichanco, 1921; Takahashi, 1927 and in
Kuwayama, 1931) and Swezey (1941) records adults on young leaves
of C. inophyllum in Samoa. Caldwell (1942) reported on specimens col-
lected in Guam with the observation by O.H. Swezey that the immatures
feed beneath the rolled over edges of C. inophyllum leaves; Caldwell
(1942) synonymised L. sulfurea rubrocincta Uichanco, 1921 with
L. sulfurea Crawford, 1919. Tuthill (1951) gives locations in the Caroline
and Palau Islands, including an observation by Esaki that this was a com-
mon gregarious feeding on young leaves of C. inophyllum. Tuthill (1964)
illustrated L. sulfurea and reported that it is widespread throughout
western Micronesia with its host plant C. inophyllum in Ambon, Fiji(?),
South Mariana Islands, Caroline Islands, Guam, and Palau. In his paper
describing Leptynoptera alagari Miyatake from Calophyllum blancoi Pl.
& Tr, in the Philippines, Miyatake (1971) also illustrates the forewing
and male genitalia of L. sulfurea. Anthony (1974, cited in Martin and
Hollis, 1992) described the gall of L. sulfurea on the leaves of Calophyllum
in Singapore.
Beardsley (1981) reported light trap capture of L. sulfurea at Honolulu
International Airport (Oahu) in 1977 and at Molokai in 1978. A survey
made in 1980 found moderate to heavy infestations of all stages of
this psyllid on young terminal growth of kamani trees (C. inophyllum)
at Honolulu International Airport, Fort Kamehameha, Salt Lake
and Kakaako. Subsequently, infestations have been seen in Manoa
(University of Hawaii Campus), Nuuanu, and in Kailua. Infestations of
immatures produce obvious crinkling and curling of the edges of
infested leaves. Yang (1984) illustrated the forewing, head, male and fe-
male genitalia, and immatures (1, 3, 4 and 5 instars) from C. inophyllum
in Taiwan. Yang (2005) gives the distribution of L. sulfurea as Taiwan,
Philippines, Ryukyu, S. Mariana Is., Caroline Is., Amboina and Fiji.
Braza and Calilung (1981) discuss more specimens collected from the
Philippines, and Hutson (1981) lists the species from C. inophyllum on
Diego Garcia Atoll in the middle of the Indian Ocean (Hutson, 1981).
Hollis (2004) cites records from C. inophyllum in the Northern Territory
and Queensland in Australia, and Li (2011) from the same host in China.
There is, finally, a series from New Caledonia in the collection of the
Muséum d’histoire naturelle, Geneva, Switzerland.
In summary, L. sulfurea has been found in Australia (mainland, Cocos
(Keeling) Islands), China (Hainan Island), Cook Islands, Diego Garcia,
Federated States of Micronesia (mainly Caroline Islands), Fiji, France
(New Caledonia), India, Indonesia (Ambon, Maluku, Sulawesi), Japan
(Ryuku Islands), Malaysia (peninsula), Mariana Islands, Palau, Papua
New Guinea, Philippines, Samoa, Singapore, Taiwan, Thailand, Tonga
and the USA (Guam and Hawaii) (Hodkinson, 1983, 1986; Martin and
Hollis, 1992; Hollis, 2004).
The host plant for L. sulfurea is C. inophyllum, and it is likely that other
host plant records are casual occurrences. C. inophyllum is a large ever-
green ornamental and timber tree that is widely dispersed throughout
the tropical regions of the Indian and Pacific Oceans. It occurs naturally
in coastal and adjacent lowland forests in East Africa (Madagascar),
India, Southeast Asia (Cambodia, Indonesia, Laos, Malaysia, Myanmar,
Philippines, Sri Lanka, Thailand, Vietnam), East Asia (Japan, Taiwan),
Australia, Papua New Guinea, and the South Pacific (Cook Islands, Fiji,
French Polynesia, Kiribati, Marshall Islands, New Caledonia, Norfolk
Island, Reunion, Samoa, Solomon Islands, Tonga, Vanuatu) (Stevens,
1980; CAB International, 2000; Friday and Okano, 2006; Orwa et al.,
2009). It is thought to be indigenous to India, Malaysia, Indonesia and
the Philippines (CAB International, 2000). C. inophyllum is often planted
within its natural distribution range because of its value as a timber tree
(Orwa et al., 2009). It has been planted in Asia (Bangladesh, China, India,
Indochina, Japan, Malaysia, Philippines, Sri Lanka, Taiwan and Thailand),
Africa (Madagascar, Nigeria, Senegal, Seychelles, Tanzania and Uganda),
Central America, Oceania (Australia, Cook Islands, Nauru, Pacific Islands,
Papua New Guinea and Solomon Islands), Hawaii and Mauritius (CAB
International, 2000). Williams (1994) records C. inophyllum from
Horsburgh and West Islands in the Cocos (Keeling) Islands, but it does
occur also on some of the other islands.
For such a widespread psyllid species, relatively little is known about
its biology. It is a gall-forming psyllid (Uichanco, 1919; Anthony, 1974;
Martin and Hollis, 1992). The main reference to the biology of
L. sulfurea is from Thailand (Tigvatnanont, 1991). The immatures were
restricted to the developing shoots and young leaves of C. inophyllum
while the adults fed on both younger and older leaves. Eggs were laid
in rows along the margin of very young leaves. The immatures excreted
copious quantities of honeydew on which sooty mould developed.
Tigvatnanont (1991) reared L. sulfurea on C. inophyllum and observed
that both males and females mated several times during their lives.
The females deposited an average of 517.95 ± 13.01 eggs; the mean
egg duration was 4.18 ± 0.08 days, and for immatures, it was 9.68 ±
0.06 days. Mean female longevity was 44.47 ± 1.77 days and for males,
it was 39.33 ± 1.30 days (Tigvatnanont, 1991). The observations by
Tigvatnanont (1991) suggest that L. sulfurea does not form an enclosed
498 P.J. Neville et al. / Journal of Asia-Pacific Entomology 18 (2015) 497–499

3. gall, and that it is more likely to distort the leaves to provide a shelter for
the immatures. Uichanco (1919, 1921) describes the species as forming
galls, while Braza and Calilung (1981) call it a gall-former that rolls up
and curls the leaf margins to enclose the immatures inside. They found
shed exuviae on the leaf surface suggesting immatures leave the galls
for adult emergence. They also stated that the immatures secrete pow-
dery wax and are associated with ants. This is the only reference to
ants associated with L. sulfurea, but is something to keep an eye on in
the Cocos (Keeling) Islands in view of the invasion of those islands by
the Yellow crazy ant, Anoplolepis gracilipes F. Smith (Neville et al., 2008).
Waterhouse (1997) lists L. sulfurea as a pest of C. inophyllum in planta-
tion forests in the southern and western Pacific (Waterhouse, 1997 p. 43)
and a pest of lesser importance in Guam (Waterhouse, 1997 p. 48).
Campbell (2003) suggests that it is an invasive alien species in Guam.
The genus Calophyllum has 187 species (Stevens, 1980) and eleven
species of Leptynoptera have been recorded as using Calophyllum as host
plants (Martin and Hollis, 1992). L. sulfurea is the most widespread spe-
cies of Leptynoptera, possibly reflecting the widespread distribution of
its host plant. Of the 10 other species of Leptynoptera, one is found in
Zanzibar, another in Sri Lanka, and the remaining eight species are dis-
tributed from Singapore to the Solomon Islands (Martin and Hollis, 1992).
One of the interesting questions about the insects of the Cocos
(Keeling) Islands is how they colonised such small islands located nearly
1,000 km from the nearest land mass, Christmas Island, to the ENE. There
are similarities between the flora and fauna of Christmas and the Cocos
(Keeling) Islands, and it may be possible that some of the insects found
on the latter originated from Christmas Island. C. inophyllum occurs on
Christmas Island (Mitchell, 1974), but L. sulfurea has yet to be found
there. The predominant winds throughout the year at the Cocos (Keeling)
Islands are the trade winds from Africa that actually arrive there from the
southeast. Winds from the Christmas Island direction (ENE) occur an
average of 4.4% over the whole year, with peaks of 7.6% (March), 7.7%
(April) and 8.5% (July) (Windfinder, 2014). These WSW winds from
Christmas Island are estimated to take 72 h to cover the 980 km journey
based on NOAA Hysplit modelling (J. Weiss pers. comm. 2014). The Cocos
(Keeling) Islands are also subjected to periodic cyclones (Bureau of
Meteorology, 2014). C. inophyllum is an oceanic seed dispersed tree and
this could explain its colonisation to these islands because there is no
evidence that C. inophyllum was planted there after human settlement.
As the centre of distribution of L. sulfurea is either Asia or Oceania, the
westerly records in the Indian Ocean, Diego Garcia and the Cocos
(Keeling) Islands, are interesting. Unless this species is found further
west in Africa, then L. sulfurea either arrived at these location by human
transport or involved a very risky transoceanic wind dispersal event.
Acknowledgments
The authors are grateful to Parks Australia North for funding this pro-
ject through DPI Victoria, and especially the staff on Cocos (Keeling)
Islands with facilitating and assisting the study. This project would not
have been possible without the encouragement and assistance of
Wendy Murray, Robert Thorn (Greenie), Ismail MacRae and Mohammad
Chongkin. Field work was conducted with the assistance of Melanie
Archer, Kylie McGregor, Dennis O’Dowd and Ben Heally. John Weiss
provided information on wind movements from Christmas Island to
the Cocos (Keeling) Islands.
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