Thrips hawaiiensis (Morgan, 1913) (Thysanoptera: Thripidae), an Asian

Blackwell Publishing Ltd

pest thrips now established in Europe

P. Reynaud1, V. Balmès2 and J. Pizzol3
1Laboratoire National de la Protection des Végétaux – 7 rue Jean Dixméras 49044 Angers cedex, France; e-mail: philippe.reynaud@agriculture.gouv.fr
2Laboratoire National de la Protection des Végétaux – unité d’entomologie 2 place Viala 34060 Montpellier cedex 01, France;
e-mail: balmes@supagro.inra.fr
3INRA – URIH 400 route des Chappes BP 167 06903 Sophia Antipolis cedex, France; e-mail: jeannine.pizzol@sophia.inra.fr

An established population of Thrips hawaiiensis (Morgan) (Thysanoptera: Thripidae), a species

originating from Asia and the Pacific, is reported for the first time in the Western Palaeartic zone
(in the South of France). Notes relating to the current geographical distribution, identification,
biology and risk of establishment of this potentially invasive species in Europe are provided.

with magnification ranging from 50 to 630×). When necessary,

Introduction
Thrips are tiny phytophagous, mycophagous or predatory
insects. They belong to the order Thysanoptera, which includes
approximately 5500 species worldwide (Mound, 2001). There
are few studies of thrips in France, even for crop pest species.
Only old publications are available (e.g. Bournier, 1983, which
lists a few species of agronomic importance), but these pre-date
the introduction of a number of new species into France:
Frankliniella occidentalis in 1986 (Bournier & Bournier,
1987), Echinothrips americanus in 1996 (Reynaud, 1998),
Neohydatothrips samayunkur in 2000 (Reynaud et al., 2001)
and Pezothrips kellyanus in 2002 (Streito & Martinez, 2005).
Material and methods
Collection
During the course of routine monitoring of the Thysanoptera
fauna inside and outside greenhouses by the Integrated
Research in Horticulture Unit (URIH) of the French National
Institute for Agricultural Research (INRA), thrips were
sampled weekly, beginning April 9, 2006, in Antibes (France:
Alpes-Maritimes, Lat.: 43°37 N – Long.: 7°02 E). They were
collected by beating various cultivated and wild plants over a
white surface, and using a fine brush to transfer them to a tube
containing 10% alcohol plus wetting agent. On each occasion,
the host plant was recorded.
Identification
Samples were identified by the entomology section of the
French National Laboratory of Plant Protection (LNPV). Often,
several species were recorded from a single tube. Initially, the
thrips were screened with a stereomicroscope (LEICA MZ12
magnification 8 to 100×), then placed in lactic acid on a slide
and examined with an optical microscope (LEICA DMLB2
permanent Canada balsam mounts were made following
traditional protocols after clearing in 10% potassium hydroxide.
Voucher specimens are deposited in the collection of the LNPV.
Species identifications were made using keys provided by
Zur-Strassen (2003).
Climatic evaluation
Potential geographical distribution was evaluated using climex
3.0 software (Hearne Scientific Software, 10 Blessington
Court, Dublin 7, Republic of Ireland). This dynamic simulation
model enables predictions to be made about the distribution of
a plant or an animal, and its relative abundance, in relation to
climate. climex 3.0 uses a climatic data file derived from a 0.5°
world grid of meteorological data calculated by interpolation
by the Climatic Research Unit (CRU), Norwich (UK) (New
et al., 1999). For each cell of the grid, a weekly and annual
Growth Index (GI) describes the potential for growth of a
population during a favourable season. Then four Stress Indices
(Cold, Hot, Wet and Dry), and in some cases their interactions,
describe the extent to which the population is reduced during an
unfavourable season. The Growth and Stress Indices are
combined into an Ecoclimatic Index (EI) that gives an overall
measure of favourableness of the location for permanent
occupation by the target species (Sutherst et al., 2007).
Results
A permanent population of Thrips hawaiiensis (Morgan, 1913)
was discovered outdoors for the first time in either Europe or
the Western Palearctic. The first two specimens were captured
on July 26, 2006, on Abelia sp. and Nerium sp. (week 31).
Afterwards, the number of captures remained low until week
43; then from week 44 a strong increase was noted, mainly on
Osmanthus fragrans (Table 1). From week 46 (November 14,
2006), captures became sporadic but some adults were still

© 2008 The Authors. Journal compilation © 2008 OEPP/EPPO, Bulletin OEPP/EPPO Bulletin 38, 155–160 155
156 P. Reynaud et al.

Table 1 List of plants on which Thrips hawaiiensis was captured in France

in 2006

Plant Number of captures

Abelia sp. 2
Acacia sp. 1
Arbutus unedo 1
Buddleja sp. 3
Nerium sp. 9
Osmanthus fragrans 99
Rosa sp. 1*
Total 116

*Specimen captured indoors. Fig. 1 Numbers of Thrips hawaiiensis captured weekly in France in 2006.

captured outdoors until the beginning of January 2007. Figure 1
gives the temporal distribution of the 116 specimens identified
between week 31 (July 2006) and week 1 (January 2007). The
sex-ratio (ratio of male to female) of this population was 0.84.
Distribution
T. hawaiiensis is a polyphagous flower-dwelling thrips, widely
distributed and common in tropical Asia and the Pacific region.
It shows undeniable invasive capacities as it is now found –
although still localised – in Africa, as well as in North and
Central America. Until now, T. hawaiiensis had never been
reported as established in Europe although a unique specimen
was reported in England by Bagnall (1927), which was
captured in the natural environment in 1919 near Horden
(County Durham) on a Compositae. However this record
remains doubtful (Collins pers. comm., 2007) and at that time,
T. hawaiiensis was not included in the British list of
Thysanoptera. Interceptions have frequently been made by
quarantine services: in the UK (2006) on Musa sp. coming from
India (http://www.defra.gov.uk/planth/interc/intercold.htm)
and by the French authorities at Roissy airport (1998 and 2000)
and Nice airport (2002) on imports from Thailand (on
Azadirachta sp. and Melia sp.) and Sri-Lanka (on Musa sp.). In
Japan (Mizobuchi et al., 1991), T. hawaiiensis is one of the
predominant thrips species found in the vicinity of 4 seaports
(Kobe, Himeji, Uno and Hirao). Figure 2 presents the geographical
distribution published by the Commonwealth Institute of
Entomology (CIE) in 1983, updated to include descriptions
published since this date (indicated by a star). The current
known geographical distribution of T. hawaiiensis is as follows:
Asia: Bangladesh (CIE, 1983), Burma (CIE, 1983), China [(CIE,
1983), Yuanjiang (Yang, 2000), Guangzhou (Zeng & Lin, 1998),
Hong-Kong (CIE, 1983)], India (CIE, 1983), Indonesia (CIE,
1983), Iran (Manzari & Golmohammadzadeh-Khiaban, 2000),
Japan (CIE, 1983), South Korea (CIE, 1983), Laos (CIE, 1983),
Malaysia (CIE, 1983), Pakistan (CIE, 1983), Philippines Islands
(CIE, 1983), Singapore (CIE, 1983), Sri Lanka (CIE, 1983), Taiwan
(CIE, 1983), Thailand (CIE, 1983), Vietnam (Nakahara, 1994).
Africa: Angola (CIE, 1983), Mozambique (CIE, 1983), Nigeria
(CIE, 1983), Sierra-Leone (CIE, 1983), Uganda (CIE, 1983),
Reunion Island (Bournier, 2000).

Fig. 2 Distribution of Thrips hawaiiensis according to Commonwealth Institute of Entomology Map no. °431 (1983); later records are indicated by a star.

© 2008 The Authors. Journal compilation © 2008 OEPP/EPPO, Bulletin OEPP/EPPO Bulletin 38, 155–160

Australasia and Pacific islands: Australia (CIE, 1983)
[Queensland, New South Wales (Mound & Masumoto, 2005)],
Fiji (CIE, 1983), Guam (Nakahara, 1994), Hawaii (CIE, 1983),
Midway Islands (Jacot-Guillarmod, 1975), Norfolk Island
(CIE, 1983), New Guinea (CIE, 1983), Samoa (CIE, 1983),
Campbell Island (Mound & Walker, 1982).
North America: USA [Florida (CIE, 1983), Georgia
(CIE, 1983), South Carolina (Sakimura, 1986), California
(Nakahara, 1994), Texas (Nakahara, 1994) and Washington DC
(Nakahara, 1994)].
Central America and the Caribbean: Jamaica (Sakimura,
1986), Mexico (Nakahara, 1994).
Taxonomy
Morgan described this species from Hawaii under the name
Euthrips hawaiiensis in 1913. Its great morphological
variability has resulted in many authors subsequently
redescribing it under various names: the nominal species has 18
recognised junior synonyms.
The genus Thrips is large with about 280 described species
worldwide (Mound, 2001). Palmer (1992) separated the Thrips
species of the Oriental and Pacific regions into five groups.
Thrips hawaiiensis was assigned to group V, characterised by
Palmer as having abdominal sternites III-VII with discal setae,
pleurotergites without discal setae and metanotum usually
striate or with some median reticulations. However, this
allocation of the species into five groups, convenient from
an identification point of view, does not necessarily reflect
phylogenetic relationships within the genus Thrips (Mound,
2005). T. hawaiiensis belongs to a complex of four species
that also includes T. florum (South Asia, Pacific, Australia,
North America), T. andrewsi (Northern India and China) and
T. exilicornis (Africa). The geographical coexistence of the two
invasive species, T. florum and T. hawaiiensis, renders their
identification particularly difficult in certain regions. Bhatti
(1999) fully discusses this subject.
Under the stereomicroscope, the specimens collected in
France corresponded to the bicoloured form with a yellow to
dark orange head and prothorax, yellow legs, and distinctly
brown abdomen. Forewings were grayish with the basal quarter
paler. The antennae were 8-segmented, brown except segment
III, the apex of segment II and the base of segment IV and V
which are yellow. According to Bhatti (1999), the nominal form
is dark brown. This form was not encountered in France.
Under the optical microscope, females of T. hawaiiensis can
be separated from the 43 other species of Thrips occurring in
France by the possession of the following characters: several
discal setae on sternite VII, pleurotergites without discal setae,
metanotum with a pair of campaniform sensilla and with S1
setae long (more than half the length of the sclerite) and close
to the anterior margin. The T. hawaiiensis collected in France
have 8-segmented antennae but a form with 7 segments exists
in certain countries. Thrips angusticeps Uzel, 1895, is the most
similar common and polyphagous species found in France but
this is distinguished by the presence of 5 – 6 distal setae on the
© 2008 The Authors. Journal compilation © 2008 OEPP/EPPO, Bulletin OEPP/EPPO Bulletin 38, 155–160
Thrips hawaiiensis now established in Europe 157
main vein of the forewing and by the irregular teeth on the
posteromarginal comb on tergite VIII.
Economic importance
T. hawaiiensis, is a strict flower-dwelling species which can
attack various crops such as tobacco (Kurozawa et al., 1964), rose
(Woo & Paik, 1971; Wang, 1982), gladiolus (Chen & Lo, 1987),
Brassica oleracea (Chandra & Lal, 1973), coffee (Ketavan, 1978),
tea (Chen, 1979), mango (Lee & Wen, 1982), citrus (Srivastava &
Bhullar, 1980; Chiu et al., 1991), apple and pear (Palmer & Wetton,
1987) and banana (Tsai et al., 1992; Jhala et al., 2004). Chen
& Chang (1987) regards T. hawaiiensis as one of the main thrips
devastating vegetables in Taiwan. Adults are very polyphagous
and can feed on numerous host plants. Takahashi (1936) and
Wang (2002) give further information about host plants.
T. hawaiiensis is also considered to be a useful pollinating
agent on oil palm (Elaeis sp.) (Syed, 1979; Tay, 1981), Cosmos
bipinnatus (Varatharajan et al., 1982) and Lantana camara
(Mathur & Ram, 1986).
However, lists of host plants should be treated with caution
because T. hawaiiensis has often been confused with T. florum.
These 2 species are morphologically very similar and at
one time (between 1966 and 1985) they were synonymized.
T. hawaiiensis causes direct damage by puncturing flowers and
fruits, inducing spot lesions, scarring, necrosis or malforma-
tions depending on the intensity of the attack. It also feeds
on pollen and host plant fertility problems can be noted.
T. hawaiiensis is not a virus vector.
Biology
There are few publications about the life history of T. hawaiiensis,
mainly from Asian authors. Murai (2001), however, has
explored in detail the relationship between temperature, and
thrips development and reproduction. Under laboratory
conditions, with thrips reared on pollen of Camelia sinensis at
constant temperature, the developmental period from egg hatch
to adult emergence was 38.9 days at 10°C (but then the adults
failed to oviposit), 19.4 days at 15°C, 16 days at 20°C, 10.3
days at 25°C and 8 days at 30°C.
According to Murai (2001), the lower threshold temperature
of development from egg to adult oviposition, calculated by
linear regression, was 10.4°C and the thermal constant was
153.8 degree-days. T. hawaiiensis is a plurivoltine species.
In Western Japan, the number of generations completing
development varied annually between 11 and 18. Longevity
and fecundity were functions of the temperature. Developmental
time decreased with temperature (18 days at 25°C but 92 days
at 15°C). Maximum longevity was reached at 15°C (121 days)
and maximum fecundity at 20°C (536 eggs). Daily egg produc-
tion was fairly constant during the lifetime of the female with,
for example, 6–8 eggs per day at 20°C. The sex-ratio was 0.74.
T. hawaiiensis appears to tolerate low temperatures and
to show a greater potential voltinism than other pest thrips
(Frankliniella intonsa, Thrips palmi, Thrips tabaci, Frankliniella
158 P. Reynaud et al.

Fig. 3 Potential European distribution of Thrips

hawaiiensis predicted by climex. The grey
shades indicate the areas with suitable climatic
conditions.

Table 2 Biological parameters used by climex, partly deduced from Murai of T. hawaiiensis in Europe. The increasing number of new
(2001) and adjusted to take into account the currently known distribution of outbreaks being detected, in particular in North America,
Thrips hawaiiensis supports this hypothesis. The discovery of a population in the
South of France confirms the existence of such a pathway, even
Growing Index (GI) Stress Index
if it has not been precisely identified.
Temperature Index Cold Stress
Lower temperature: 10.4°C Temperature threshold: 1°C Probability of establishment
Lower optimum temperature: 15°C Temperature rate: −0.005 For a successful establishment, two criteria have to be met: the
Upper optimum temperature: 25°C
presence of at least one host-plant and a suitable climate to
Upper temperature threshold: 34°C Dry Stress
Degree-days for total development: 153.8 Dry stress threshold: 0.2
allow the species to complete at least one generation. The
Dry stress rate: −0.005 polyphagy of T. hawaiiensis guarantees it an easy access to
Wet Stress food resources in France and Europe, both outdoors and under
Wet stress threshold: 2.5 glass. But Europe might be a marginal climatic zone because
Wet stress rate: 0.002 this thrips is known to be of tropical origin, so climatic
parameters need to be studied more precisely. The potential
geographical distribution of T. hawaiiensis was assessed
occidentalis, Scirtothrips dorsalis, Thrips imaginis). Murai with the climex software based on biological parameters
(2001) concluded that there was considerable potential for (Table 2) partly deduced from Murai (2001) and adjusted to
T. hawaiiensis to become a major pest. Nevertheless, T. take into account the currently known distribution of this insect.
hawaiiensis has not become a major pest in Japan, possibly due The results (Figure 3) suggested that conditions suitable for
to its overwhelming preference for flowers rather than other establishment of T. hawaiiensis exist across the whole
parts of plants and an apparent lack of insecticide resistance. Mediterranean area (from Israel to Spain, and North Africa) and
It is also noted that there may be interspecific variation with along the Atlantic coast of both France and Spain. In Northern
respect to temperature tolerance. Europe, conditions are less favourable but establishment cannot
be completely excluded. The number of generations that could
complete their development annually will vary from 10 to 15
Phytosanitary risk
(up to 20 on a local basis) according to the different regional
A pest risk assessment classically includes 2 steps: firstly, conditions.
an assessment of the risk of introduction (i.e. entry and
establishment) and, secondly, an assessment of the economic
Evaluation of the potential economic and environmental
and environmental damage.
damage

T. hawaiiensis is found on many economically important plants

Risk of introduction
Probability of entry
The international trade of flowers and fruits from contaminated
regions is probably the main pathway for the entry and spread
worldwide. Adults and larvae usually feed on pollen and empty
cells of growing flowers. Feeding produces direct damage such
as necrosis, bud malformation or reduced fruit set (Childers &
Nakahara, 2006), but as previously mentioned, it is not a virus

© 2008 The Authors. Journal compilation © 2008 OEPP/EPPO, Bulletin OEPP/EPPO Bulletin 38, 155–160

Table 3 Economic indicators for crops potentially threatened by Thrips
hawaiiensis in France
Production Surfaces Number of farms Production value
Ornamentals 21 000 ha 11 000 $1.0 billion
Vegetable crops 277 400 ha 36 100 $3.1 billion
Tobacco 8 153 ha 3 662 $98 million
Orchards 188 759 ha 20 691 $2.5 billion
Sources: Agreste, Viniflhor, CTIFL and Eurostat.
vector. In France, for example, vegetable crops, ornamentals,
fruit crops and tobacco could all be at risk. As shown in Table 3,
these crops are of major economic importance.
Accurate data quantifying the economic losses (yields or
quality) caused by T. hawaiiensis is scarce in the literature, and
usually restricted to Asia or Oceania. There are few pest reports
from North America, where T. hawaiiensis was first found in
the 1960s. Therefore, the potential risk presented by this species
for France and the rest of Europe remains very uncertain.
Conclusion on the risk
T. hawaiiensis is polyphagous and because its host range
includes many plants which are widely traded, its potential for
entry into Europe appears to be rather high. Like other
Thysanoptera species, its small size makes it difficult to detect
during import controls. Although European climatic conditions
appear to be potentially suitable, T. hawaiiensis is not a virus
vector and, so far, it has not proven to be very harmful in the
non-native zones where it is now established. Therefore, based
on current knowledge, the overall risk seems low. However,
caution is recommended when considering past experiences
with similar species. Echinothrips americanus which was
thought to be a low risk when it was first discovered in Europe,
now appears to have been under-estimated as a pest (Pijnakker,
2003).
Acknowledgements
We would like to thank Dr. Dom Collins, Central Science
Laboratory, York, England for critically evaluating this work
and for helpful and extremely valuable comments. Authors are
grateful to Dr. R. Zur Strassen, Forschungsinstitut Senckenberg,
Frankfurt am Main, Germany for confirmation of Thrips
hawaiiensis identity.
We also thank D. Nammour, C. Bresch, P. Hervouet, S. Voisin
and J.P. Ziegler for their assistance in this work.
Thrips hawaiiensis (Morgan, 1913)
(Thysanoptera: Thripidae), un ravageur
originaire d’Asie désormais établi en Europe
Une population établie de Thrips hawaiiensis (Morgan)
(Thysanoptera: Thripidae), un ravageur originaire d’Asie et du
Pacifique, est signalée pour la première fois dans l’Ouest
© 2008 The Authors. Journal compilation © 2008 OEPP/EPPO, Bulletin OEPP/EPPO Bulletin 38, 155–160
Thrips hawaiiensis now established in Europe 159
paléarctique (sud de la France). Des éléments portant sur la
répartition géographique actuelle, l’identification, la biologie
et le risque d’établissement en Europe de cette espèce
potentiellement invasive sont fournis.
References
Anonymous (1983) CIE (Commonwealth Institute of Entomology) Thrips
hawaiiensis (Morg). Distribution Maps of Pests, Series A (Agricultural)
No. 431 (revised) (London, UK: Commonwealth Agricultural Bureaux),
1–2.
Bagnall RS (1927) Contributions towards a knowledge of the European
Thysanoptera III. Annals and Magazine of Natural History 20(9), 561–
585.
Bhatti JS (1999) New characters for identification of the pest species Thrips
hawaiiensis and florum (Terebrantia: Thripidae). Thrips 1, 53.
Bournier A (1983) Les thrips, Biologie, importance agronomique. Ed.
Institut National de la Recherche Agronomique Paris, France, 1–128.
Bournier A & Bournier JP (1987) L’introduction en France d’un nouveau
ravageur: Frankliniella occidentalis. Phytoma 388, 14–17.
Bournier JP (2000) Les Thysanoptères de l’île de la Réunion: Terebrantia.
Bulletin de la Société Entomologique de France 105, 65–108.
Chandra J & Lal OP (1973) Record of thrips on some vegetables and
ornamental plants from Kulu Valley. Himachal Pradesh Indian Journal
of Entomology 35, 164–166.
Chen JS & Lo PKC (1987) Differential preference of the flower-dwelling
thrips, Thrips hawaiiensis (Morgan) (Thysanoptera: Thripidae), to some
gladiolus cultivars. Journal of Agricultural Research of China 36, 317–326.
Chen LS (1979) [Thrips infesting tea in Taiwan]. Plant Protection Bulletin,
Taiwan 21, 377–382 (in Chinese).
Chen WS & Chang FI (1987) [The infestation of thrips on vegetables and
their control]. Chinese Journal of Entomology, Special Publication 1, 45–
53 (in Chinese).
Childers CC & Nakahara S (2006) Thysanoptera (thrips) within citrus
orchards in Florida: Species distribution, relative and seasonal abundance
within trees, and species on vines and ground cover plants. Journal of
Insect Science 6, 1–19.
Chiu HT, Shen SM & Wu MY (1991) [Occurrence and damage of thrips in
Citrus orchards in southern Taiwan]. Chinese Journal of Entomology 11,
310–316 (in Chinese).
Jacot-Guillarmod, CF (1975) Catalogue of the Thysanoptera of the World.
Annals of Cape Province Museum (Natural History) 7(4), 977–1255.
Jhala RC, Borad PK & Bharpoda TM (2004) Incidence of Thrips hawaiiensis
(Morgan) on banana in Gujarat. Insect Environment 10, 55.
Ketavan C (1978) Thrips of economic importance and their control in
Thailand. Report, Kasetsart University, 90.
Kurozawa M, Takaoka I & Naito T (1964) Thrips infesting the tobacco
plants in Japan (Thysanoptera). Japanese Journal of Entomology 32, 402.
160 P. Reynaud et al.
Lee HS & Wen HC (1982) [Seasonal occurrence of and injury caused by the
thrips and their control on mangoes]. Plant Protection Bulletin (Taichung)
24, 179–187 (in Chinese).
Manzari S & Golmohammadzadeh-Khiaban N (2000) Thrips hawaiiensis
(Morgan, 1913), a new species for the thrips fauna of Iran. Journal of
Entomological Society of Iran 19, 107–108.
Mathur G & Ram HYM (1986) Floral biology and pollination of Lantana
camara. Phytomorphology 36, 79–100.
Mizobuchi M, Fujiwara Y, Kobayashi K, Ikawa Y, Maruyama S & Ogino T
(1991) Notes on thrips (Thysanoptera) collected in and around ports of
Kobe, Himeji, Uno and Hirao. Research Bulletin of the Plant Protection
Service Japan 27, 115–126.
Mound LA (2001) So many thrips-so few tospoviruses. In Thrips, Plants,
Tospoviruses: the Millennial Review Proceedings of the 7th International
Symposium on Thysanoptera 15–18.
Mound LA (2005) The Thrips orientalis group from South-east Asia and
Australia: some species identities and relationships (Thysanoptera,
Thripidae). Australian Journal of Entomology 44, 420–424.
Mound LA & Masumoto M (2005) The genus Thrips (Thysanoptera,
Thripidae) in Australia, New Caledonia and New Zealand. Zootaxa 1020,
1–64.
Mound LA & Walker AK (1982) Terebrantia (Insecta: Thysanoptera).
Fauna of New Zealand 1, 1–115.
Murai T (2001) Development and reproductive capacity of Thrips hawaiiensis
(Thysanoptera: Thripidae) and its potential as a major pest. Bulletin of
Entomological Research 91, 193–198.
Nakahara S (1994) The genus Thrips Linnaeus (Thysanoptera: Thripidae)
of the New World. Technical Bulletin. United States Department of
Agriculture 1822, 1–183.
New M, Hulme M & Jones PD (1999) Representing twentieth century
space-time climate variability Part 1: development of a 1961–90 mean
monthly terrestrial climatology. Journal of Climate 12, 829–856.
Palmer JM (1992) Thrips (Thysanoptera) from Pakistan to the Pacific: a
review. Bulletin of the British Museum (Natural History), Entomology
Series 61, 1–76.
Palmer JM & Wetton MN (1987) A morphometric analysis of the Thrips
hawaiiensis (Morgan) species-group (Thysanoptera: Thripidae). Bulletin
of Entomological Research 77, 397–406.
Pijnakker J (2003) Echinothrips americanus, un ravageur sous-estimé.
PHM-Revue-Horticole 449, 25–27.
Reynaud P (1998) Les Thysanoptères des cultures légumières et ornementales:
2 années d’inventaire et un invité surprise! In First transnational work-
© 2008 The Authors. Journal compilation © 2008 OEPP/EPPO, Bulletin OEPP/EPPO Bulletin 38, 155–160
shop on biological, integrated and rational control: status and perspectives
with regard to regional and European experiences, Lille, France, 21–23
January 1998, 17–18.
Reynaud P, Bertaux F & Martinez M (2001) Premier signalement en Europe
de Neohydatothrips samayunkur (Kudo) (Thysanoptera, Thripidae).
Nouvelle Revue d’Entomologie 18, 91–93.
Sakimura, K (1986) Thrips in and around the coconut plantations in Jamaica,
with a few taxonomic notes (Thysanoptera). Florida Entomologist 69,
348–363.
Srivastava S & Bhullar JS (1980) Thrips – a new record as a pest of citrus
blossoms in Himachal Pradesh. Current Science 49, 747.
Streito JC & Martinez M (2005) Nouveaux ravageurs, 41 espèces depuis
2000. Phytoma – La défense des Végétaux 586, 16–20.
Sutherst RW, Maywald GF & Kriticos D (2007) CLIMEX version 3 User’s
Guide. CSIRO Publishing, Melbourne, 1–131.
Syed RA (1979) Studies on oil palm pollination by insects. Bulletin of
Entomological Research 69, 213–224.
Takahashi R (1936) Thysanoptera of Formosa. The Philippine Journal of
Science 60(4), 427–459.
Tay KC (1981) Observations on insects visiting oil palm inflorescences. The
Planter 57, 82–91.
Tsai YP, Hwang MT & Chen HP (1992) [Occurrence and damage of Thrips
hawaiiensis in banana orchards]. Chinese Journal of Entomology 12,
231–237 (in Chinese).
Varatharajan R, Gopinathan K & Ananthakrishnan TN (1982) Comparative
efficiency of thrips in relation to other foraging insects in the pollina-
tion of Cosmos bipinnatus Cav (Compositae). Proceedings of the
Indian National Science Academy, B (Biological Sciences) 48(6), 735–
739.
Wang C (2002) [Thrips of Taiwan: Biology and Taxonomy]. Taiwan
Agricultural Research Institute 99, 1–328 (in Chinese).
Wang CL (1982) [Insect pests and their injury on rose]. Journal of Agricultural
Research of China 31, 97–101 (in Chinese).
Woo KS & Paik WH (1971) [Studies on the thrips (Thysanoptera) unrecorded
in Korea (I)]. Korean Journal of Plant Protection 10, 69–73 (in Korean).
Yang Y (2000) [Harm Characteristics and the Control of Thrips in Mangifera
indica in Yuanjiang]. Yunnan Forestry Science and Technology 3, 57–60
(in Chinese).
Zeng XN & Lin JT (1998) [Damage of Thrips hawaiiensis (Morgan) to
banana and its control]. Plant Protection 24(6), 15–17 (in Chinese).
Zur-Strassen R (2003) Die terebranten Thysanopteren Europas und des
Mittelmeer-Gebietes. Die Tierwelt Deutschlands 74, 1–277.