AProceedingsdventive T hysof theanopter Hawaiiana in HEantomologicalwaii Society (2017) 49:17–28 17

Adventive Thysanoptera in the Hawaiian Islands: New Records and Putative Host Associations

Laurence A. Mound1, Janis N. Matsunaga2, Brian Bushe3, Mark S. Hoddle4, and Alice Wells1 1CSIRO Australian National Collection, Canberra, Australia Laurence.mound@ csiro.au; 2Hawaii Department of Agriculture, Pest Control Branch, 1428 S. King St., Honolulu, , 96814; 3University of Hawaii at Manoa, College of Tropical Agriculture, Agricultural Diagnostic Service Center; 4Department of Entomology and Center for Invasive Species Research, University of California

Abstract. Fifteen adventive species of Thysanoptera, comprising eight and seven , are recorded for the first time from the Hawaiian Islands. Four of these are native Australian species, seven are native to south- eastern Asia, with one species known only from Costa Rica and one known from several sites around the Caribbean. New host and locality records are given for a further 15 species, and the situation is noted that some species established on the Hawaiian Islands are pests in other parts of the world but have failed to become problematic here.

Key words: Hawaii, new state records,

The number of insect species recog- Checklist (Nishida 2002) over 330 arthro- nized as being adventive to the Hawaiian pod species adventive to Hawaii have been Islands is large and continues to increase, recorded (Howarth et al. 2013). However, although recent summary statistics about as noted by Beardsley (1962), discovery species establishment rates are not readily and reporting of many alien in available. According to Beardsley (1962), Hawaii is associated with special projects over the 25-year period 1937–1961 ap- and interests of local and visiting research- proximately 400 adventive insect species ers, and are not indicative of true dates of were reported as new to the islands, with arrival or establishment (Howarth et al. an average rate of establishment as high 2013). as 20 species per year (Beardsley 1979). Incursions of exotic resulting In a Farm Bill Report to the United States in successful establishment are ongoing, Department of Agriculture (USDA), because Hawaii is a major hub for tour- Howarth et al. (2013) indicated that the ism, trade, and military traffic (Messing nursery trade and various cut flower and et al. 2007). The result of this ongoing agricultural industries, in importing live process is that the present-day insect fauna plant material, is probably a major source of Hawaii is dominated by introduced of adventive and invasive phytophagous species, a situation that is facilitated by insects in Hawaii. These authors reported anthropogenic activities (Messing et al. an estimate of 27 to 29 insect species 2007). Native and endemic Hawaiian in- arriving and establishing each year in sect species are restricted to small areas at Hawaii. Moreover, since the last revision higher elevations, although even these are of the Hawaiian Terrestrial often altered by non-native such 18 Mound et al. as goats and pigs, and the invasion by a ogy presented at the USDA Pacific Basin diversity of weedy . The extent to Agricultural Research Center (PBARC) which adventive insect species can poten- in Hilo, samples of thrips were taken on tially colonize this native flora is a major three islands, Hawaii, Maui, and Oahu. concern. Examples are given As a result, 15 additional adventive thrips below of species that are pests in some species were discovered, and information parts of the world, but when introduced to was gathered on the putative host associa- the Hawaiian Islands have failed to cause tions of several other species. The purpose noticeable damage to crop plants or native of this work is to present new information vegetation. In this context we distinguish about these non-native thrips species. For between “adventive” and “invasive” spe- the sake of brevity, full nomenclatural cies. Adventive species are non-native, details of collected species are not given accidentally introduced, immigrants here, but are available for all thrips taxa or are species whose populations were at ThripsWiki 2017. Specimens will be intentionally introduced (e.g., biological deposited into the collections both at control agents) (Frank and McCoy 1995). the Hawaii Department of Agriculture In this context, we refer to accidentally (HDOA) and CSIRO, Australian National introduced adventive thrips as those caus- Insect Collection, Canberra (ANIC), or as ing no economic or ecological harm to noted. the ecosystem that has been infiltrated. In contrast, non-native populations that have Newly Recorded Thripidae become pestiferous after establishing are Biltothrips minutus Bhatti. This min- considered invasive species. ute white insect with black forewings was The insect Thysanoptera com- abundant in the vicinity of Hilo, Hawaii, prises just over 6000 species worldwide, breeding on the leaves of both taro (Co- with most of these species residing in locasia esculenta [Araceae]) and cassava tropical and subtropical areas. However, (Manihot esculenta [Euphorbiaceae]). despite the fact that these insects, known Feeding damage by this thrips on C. es- as thrips, are small and potentially dis- culenta was unusual (Fig. 1), comprising persive, there are only around 200 spe- small linear markings that resembled leaf- cies recorded from the Hawaiian Islands. mines of minute Lepidoptera. Similar leaf Moreover, of the 99 species of the sub- damage in association with this thrips was order currently listed from also noted on cabbage (Brassica oleracea these islands (Mound et al. 2016), no more [Brassicaceae]), avocado (Persea ameri- than seven are considered to have evolved cana [Lauraceae]), and blueberry (Vac- here, with the other 90 Terebrantia species cinium sp [Ericaceae]). This biparental having been accidentally introduced from species was described from West Bengal, other countries through human activities. India. Subsequently it was reported from At least two species are known to have Thailand, and from Malaysia (both sexes been deliberately introduced as biocon- were collected from cassava [Ng et al. trol agents: urichi against the 2014]), and also the Society Islands (Hod- highly invasive forest weed, dle et al. 2008, Ng and Mound 2015). In hirta (), and Sericothrips the key to Hawaiian Terebrantia (Mound et staphylinus against the noxious weed, al. 2016) B. minutus will run to the gorse, Ulex europaeus (Fabaceae). Dur- , because of the presence of ing July 2016, in association with a short closely spaced rows of microtrichia later- course on thrips identification and biol- ally on the tergites. However, in contrast Adventive Thysanoptera in Hawaii 19

Figures 1–5. Thrips new to Hawaii. 1. Leaf-damage on Colocasia esculenta by Bil- tothrips minutus (photo: S. Chun). 2. Head of Indusiothrips seshadrii. 3. Head and pronotum of Monilothrips kempi. 4. Head and thorax of Trichromothrips priesneri. 5. Forewing of Coremothrips pallidus. 20 Mound et al. to Scirtothrips species, Biltothrips species Indusiothrips seshadrii Priesner. Un- have the median pair of setae on the sixth til now, this species was known only from to the eighth tergites as large as, and situ- the original specimens collected in south- ated close to, the submedian setal pair, ern India. It is identified here by means instead of being small and close together of the key that was provided by Wilson in the mid-line. (1975) in describing the only other species manicatus (Haliday). The in the genus. Both of them apparently live record from the Hawaiian Islands of this on ferns, and I. nakaharai has been found common grass thrips of temperate coun- in large numbers on Polypodiaceae in tries was considered doubtful by Mound Honshu, Japan (Okajima and Urushihara et al. (2016), but an overlooked report by 1994). The two species are distinctive in Krushelnycky et al. (2007) indicated that having the head reticulate (Fig. 2), and all the species had been found at Haleakala the major setae minute on the body and National Park, Maui, during a survey forewings, except for two pairs of remark- between 2001 and 2004, and in July 2016 ably long setae with broadly expanded substantial populations were found on apices on tergite IX. Both species are the upper slopes of Haleakala within the small and white in color, but seshadrii has national park. As with other species of uniformly dark forewings and a dark area Chirothrips, this thrips breeds and pupates on the anterior half of the head, whereas within the florets of grasses, and it is thus nakaharai is almost entirely yellow. One easily distributed in commercial grass female of seshadrii was taken from an seed. Moreover, it seems to be catholic unidentified fern on the Mokuleia Trail, in the range of Poaceae on which it can Oahu (in ANIC). breed, and large populations have been Monilothrips kempi Moulton. De- studied from Australian native grass spe- scribed from northern India, with one cies in montane areas of Tasmania. synonym from South Africa, this fern- Coremothrips pallidus Hood. In living Panchaetothripine is also known December 2016 Randy Hamasaki found from California (Wilson 1975), where it larvae and adult females of this species was collected recently in large numbers on feeding on the leaves of avocado at Laupa- the fern Woodwardia fimbriata [Blechna- hoehoe, Hawaii. Also in December 2016, ceae]. On Hawaii, at Kipuka Ki, Hawaii a single female was taken from avocado Volcanoes National Park, it was found foliage in Waimanalo, Oahu. This is a breeding on the fronds of the fern Micro- very small, pale species, but the adults are lepia strigosa [Dennstaedtiaceae] in as- remarkable for having the major setae on sociation with a substantial population of the forewings (Fig. 5), head and pronotum the predatory ant-mimic, Franklinothrips exceptionally long with expanded fringed vespiformis. There is only one species apices. Similar setae also occur on the recognised in the genus Monilothrips, and dorsal surface of the larvae. This thrips kempi is a large dark brown species that was described from specimens taken from is distinctive among avocado on Trinidad, and is also reported in lacking reticulate sculpture on the from Panama, Puerto Rico, Guadeloupe fore femora and much of the head. The and St. Vincent (Mound and Marullo posterior third of the head has a strongly 1996). However, a survey of thrips associ- reticulate transverse band (Fig. 3), and the ated with avocado in several Neotropical meso- and metanota are also reticulate, countries (Hoddle et al. 2002) did not find the forewing bears two complete rows of this species. veinal setae, the sense cones on antennal Adventive Thysanoptera in Hawaii 21 segments III and IV are long and forked, sandwicensis [Oleaceae], one on Hawaii and antennal segments II–IV are almost on Mauna Loa at 2050 m from flowers of yellow. Vaccinium sp. [Ericaceae], and a third on Pseudanaphothrips achaetus (Bag- Maui at Waihee Ridge, from the flowers nall). Both sexes of this highly polypha- of an introduced Acacia sp. [Fabaceae]. gous Australian flower thrips were found Trichromothrips priesneri (Bhatti). on Maui, Haleakala National Park, in Two females of this species, previously the flowers of the native plant Geranium known only from India, were collected cuneatum [Geraniaceae] and the flowers on Maui, on the wetlands behind Waihee of an introduced Oenothera stricta [Ona- Beach amongst dodder and Cyperaceae graceae]. It is distinguished from the other (deposited in ANIC). These females were members of this predominantly Australian identified from the description given by genus by the absence of any long setae Bhatti (2000) in a key to the species of the on the pronotum. In the key to genus. These specimens agree well with genera in Mound et al. (2016) this species that description, except that they have the will run to , but in contrast basal half of antennal segments III and to members of that genus the forewing in IV more extensively yellow. T. priesneri Pseudanaphothrips species resembles that is remarkable for its color, even in the of species of in having both field. The abdomen and forewings are longitudinal veins with a complete row of dark brown, but the pterothorax is white, setae. the pronotum is white with brown lateral Thrips safrus Mound and Masumoto. margins (Fig. 4), and the head is white This species was identified from the Ha- medially but brown at the anterior and waiian Islands as Thrips imaginis Bagnall posterolaterally. The only other member (see Mound et al. 2016), based on speci- of this genus with a similar dark brown mens collected on Leptecophylla tamei- abdomen is T. elegans from Japan, but that ameiae [Ericaceae] flowers at Haleakala has a uniformly dark head and a smaller National Park, Maui. Contrary to previous pale area medially on the pronotum (Ma- information, those specimens were not re- sumoto and Okajima 2005). tained in 2003 by the Systematic Entomol- ogy Laboratory, USDA, Beltsville. They Newly Recorded Phlaeothripidae were returned to the collector, Paul Krush- Adraneothrips alajuela Mound and elnycky, through whose courtesy they Marullo. Previously known only from have now been re-examined. In contrast Costa Rica, four females were taken on Ha- to specimens of imaginis, the Australian waii in July, 2016; two from a dead branch plague thrips, the females collected from of Metrosideros polymorpha [Myrtaceae] Maui lack discal setae on the abdominal at Tom’s Trail, 30 km southwest of Hilo, pleurotergites. This condition is typical one from dead branches of Metrosideros of safrus, a polyphagous species that is polymorpha at the Volcano Research Sta- common in northern Australia (Mound tion, and one from a dead branch in a and Masumoto 2005). The two species are garden at Volcano. Generally medium otherwise closely similar in structure, and brown in color (Fig. 6), this species has both are notable for the numerous (15–25) abdominal segment II yellow with its an- discal setae on the abdominal sternites. terior margin weakly shaded, and tergites Three further females of safrus were taken VI and VII lighter brown. The compound on the Hawaiian Islands in July, 2016; one eyes are prolonged ventrally, and the pelta on Hawaii near Volcano from Nestegis has very small lateral wings and bears a 22 Mound et al.

Figures 6–9. Thrips new to Hawaii. 6. Head and pronotum of Adraneothrips alajuela. 7. Abdominal tergites 1 and 2 of A. alajuela. 8. Head and pronotum of Azaleothrips siamensis. 9. Head and antenna of Sophiothrips annulatus. pair of campaniform sensilla (Fig. 7). lacking duplicated cilia on the forewing. Adraneothrips russatus (Haga). Azaleothrips siamensis Okajima. Described from Japan, but known from Identified here from the original descrip- southern China and northeastern Australia tion and key to species (Okajima and (Dang et al. 2013), several females of this Masumoto 2014), this thrips was found on species were taken from dead branches dead branches in considerable numbers on in July 2016 on the Mokuleia Trail, Oahu Hawaii. It was collected in July 2016 from (deposited in ANIC). Strikingly bicolored, recently felled trees at USDA PBARC in with a brown head, pale pronotum, and Hilo, in lower Puna, and also in the forest banded tergites, this species is unusual in at Kipuka Ki, Hawaii Volcanoes National Adventive Thysanoptera in Hawaii 23 Park. These specimens have the color, more extensively yellow, and the antennae chaetotaxy, and sculpture character states are slightly longer (Fig. 9). The pelta of the of siamensis (Fig. 8), although the mouth apterae varies amongst the six available cone appears to be rather shorter in not specimens, from rectangular with small extending across the mesosternum. lateral wings to almost as rounded as on Cartomothrips neboissi Mound and the paratypes from Okinawa (Okajima Walker. A substantial breeding popula- 1994). tion of this Australian species was found on Hawaii, on the slopes of Mauna Loa Notes on Recent Host- just above the tree line at about 2200 m. association and Locality Records The adults and first instar larvae were Anisopilothrips venustulus (Priesner). taken from Metrosideros polymorpha The specimens of this Panchaetothripi- [Myrtaceae], where they were living nae species on which the record from within old seed capsules. The trees at this Hawaii was based (Mound et al. 2016) altitude were sparsely distributed and were were taken from leaves of stellata no more than 6 ft in height. This thrips is [] at Keeau, Hawaii, in May common in southeastern Australia, living 2010 (in Hawaii Department of Agricul- in the seed capsules of Kunzea ericoides ture Collection, Honolulu). Considerable [Myrtaceae], and is also recorded from populations of this thrips were found in New Zealand, North Island (Mound and July 2016 causing leaf damage on Alyxia Walker 2012). stellata at Kipuka Puaulu, Hawaii Volca- Plectrothrips sp. A single female of noes National Park, Hawaii. The species this genus was taken on Hawaii from dead is otherwise known from Taiwan, Japan, wood at Kipuka Ki, Hawaii Volcanoes Na- Australia, Fiji, and Florida, but with little tional Park, and one male and one female reliable host plant information. This ap- of the same species were taken on Oahu pears to be a new host association for this from dead branches on the Mokuleia Trail species. (deposited in ANIC). The species cannot Dichromothrips smithii (Zimmer- at present be identified, because all three mann). As previously reported by Hawaii specimens are de-alate. However, it ap- Department of Agriculture (2009) and pears to share all the available character Hollingsworth et al. (2012), substantial states with orientalis Okajima from Ma- populations of this thrips were found on laysia (Okajima 1981). the bamboo orchid, Arundina graminifo- Pygothrips sp. One female, possibly re- lia, that grows extensively along the road- lated to mikrommatos Moulton from Fiji, sides in lower Puna. This thripid occurs was taken on Hawaii from dead branches widely in southeast Asia, and is a pest of at USDA PBARC, Hilo (deposited in Vanilla [Orchidaceae] in India. ANIC). Dolichothrips franae Mound and Sophiothrips annulatus Okajima. Okajima. This impressive large species Four female and two male apterae, also of has previously been two female macropterae and one female known only from the islands of Oahu hemimacroptera, were taken on Maui in and Kauai (Mound and Okajima 2015), July 2016, from dead branches at Iao Val- where it was taken in good numbers ley, and on the trail to Waihee Ridge. In from the flowers of Macaranga tanarius contrast to two paratype female apterae [Euphorbiaceae], with a few specimens of annulatus that have been studied, these also recorded from Hibiscus tiliaceus specimens have the femora and tibiae [Malvaceae] (Mound and Matsunaga 24 Mound et al. 2017). However, on Hawaii, at Waikolola within the upper slopes of Mt. Haleakala, Dry Forest Reserve, a substantial breeding only two females of occidentalis were population was found amongst the apical taken, whereas on the lower slopes where leaves of Hibiscus brackenridgei [Malva- there are many non-native plants this ceae]. A similar pair of contrasting host thrips was common. associations involving these two unrelated (Trybom). plant genera has been recorded for Doli- The tomato thrips has been recorded from chothrips reuteri (Karny) in Australia, and several of the Hawaiian Islands, but it was possibly reflects some physical or chemi- found in July 2016 only in low numbers cal similarity in the apical tissues of these and in dry areas, including Waikoloa Dry unrelated plants. This is the first report of Forest Reserve, and Puu Waawaa Forest D. franae for the island of Hawaii. Reserve. Only the pale form of this species Frankliniella insularis (Franklin). was found. This Central and South American spe- Hercinothrips bicinctus (Bagnall). Al- cies is likely to be widely established on though first listed for the Hawaiian Islands the Hawaiian Islands, although currently in 1998, from Oahu (Mound et al. 2016), recorded only from Hawaii island. In this species was found commonly in July July 2016 on Hawaii, it was abundant on 2016 at various sites on Hawaii and Oahu Hibiscus flowers in lower Puna, and also (see also Kumashiro et al. 2002). Large on Jacaranda flowers at Puu Waawaa populations were present on the leaves Forest Reserve. At the later site it was also of recently planted specimens of the rare taken from the flowers of the endangered endemic plant Clermontia lindseyana endemic plant [Mal- [Campanulaceae] near Kipuka Puaulu, vaceae]. Hawaii Volcanoes National Park, and this Frankliniella occidentalis (Pergande). is the first report of this host plant associa- This ubiquitous, worldwide pest, the west- tion. In contrast, only a few specimens of ern flower thrips, was collected widely on Hercinothrips femoralis were found, and Hawaii. Large populations were found these were on the leaves of taro in as- on Mauna Kea in the flowers of the na- sociation with populations of Biltothrips tive plant species, Sophora chrysophylla minutus at Hilo (deposited in ANIC). [Fabaceae], and oahuense Pezothrips kellyanus (Bagnall). [], as well as in the flowers Known as Kelly’s citrus thrips, this spe- of the introduced plant species, Senecio cies is considered a pest of citrus crops in madagascariensis and Achillea millefo- South Australia, New Zealand, and south- lium [Asteraceae]. These infestations of ern Europe. It is not recorded from citrus native plants may have been facilitated in the Hawaiian Islands, but both sexes by the extent to which the vegetation at were collected in small numbers at various that site was degraded and dominated by sites on Hawaii, Maui, and Oahu, mainly non-native plants that are hosts for this from white flowers of species in the genera pest thrips. In contrast, within healthy Aleurites [Euphorbiaceae], Gardenia [Ru- native forest at Kipuka Puaulu, Hawaii biaceae], Myoporum [Scrophulariaceae] Volcanoes National Park, F. occidentalis and Sida [Malvaceae]. Specimens of this was found only in the flowers ofIpomoea thrips are also available in the Hawaii indica [Convolvulaceae] and was not col- Department of Agriculture collection lected from the flowers of any endemic from loquat (Eriobotrya japonica) [Ro- plant species. Moreover, on the extensive saceae], Persea americana [Lauraceae], areas of native plants that grow on Maui and Myoporum sandwicense. Adventive Thysanoptera in Hawaii 25 First recorded in 2006 on the Hawaiian bean, and is a significant pest of orna- Islands, collected at Kula, Maui, records mental and horticultural crops (Kumar et also show that this species was identified al. 2012). S. dorsalis was first detected on from the same area as far back as 2000 Oahu in 1987, then on Maui in 1988. On (Hawaii Department of Agriculture 2007). Hawaii, in July 2016, it was found in large Krushelnycky et al. (2007), reported numbers on the leaves of Mimosa pudica collections of this species at Haleakala [Fabaceae] growing on the sea cliffs in National Park, Maui on Leptocophylla lower Puna, and in 2009 was found heav- tameiameiae and Vaccinium reticulatum ily infesting Myoporum sandwicense (all of these specimens were collected [Scrophulariaceae] foliage at Waikoloa. simply by beating these plants, so may not S. dorsalis continues to be another pest of be definitive host-association data), and in the native M. sandwicense that is already pitfall traps. Although not specified, these under threat from Klambothrips myopori. collections were in 2003 (Krushelnycky Scirtothrips inermis Priesner. Previ- 2017, review comments, 22 July). ously recorded from Maui (Nishida 2002; Pseudanaphothrips araucariae Mound et al. 2016), a single female of this Mound and Palmer. Described from species was taken in July 2016 from an Australia, but also known from Hawaii unidentified plant in Hawaii Volcanoes and Oahu where it was collected from National Park, Hawaii. the male cones of the Norfolk Island Pine, Scirtothrips perseae Nakahara. The Araucaria heterophylla [Araucariaceae], avocado thrips is a pest in California, a single female of this species was taken damaging the leaves and young fruit of from flowers ofDodonaea viscosa [Sapin- Persea americana [Lauraceae], although daceae] in July 2016 on Maui on the lower it has a wide distribution on these trees slopes of Mt. Haleakala. This is the first southwards through Mexico to Guatemala report of this species from Maui. (Mound and Hoddle 2016). It appears to Sciothrips cardamomi (Ramak- have become established in Maui since rishna). The cardamom thrips was found 2006, but was first confirmed from Ha- widely on both Hawaii and Oahu. This waii in 2016 where it remains localized at thrips occurred in large numbers, breed- Kamuela. At present, there is limited in- ing within the apical rolled leaves and formation on its pest status on the islands. leaf axils of Hedychium gardnerianum According to Mach Fukada (pers. comm. [Zingiberaceae]. A few adults were also 2017), avocado thrips seems to be limited found in a similar situation on Hedychium on Maui by the environment. In lowland coronarium on Oahu, confirming a new areas populations seem to crash during island record of this species high temperatures, and severe damage to Scirtothrips citri (Moulton). This Cali- avocado trees is generally not apparent be- fornian pest species was recorded from low 2000 feet elevation. The females from collections on mango and lime foliage at the sample on Hawaii have the transverse Kihei, Maui, in November 1994. There dark markings on the tergites rather paler appear to be no more recent collections, than on specimens from California, but and it remains unclear if this polyphagous are otherwise identical in structure with species is established on the Hawaiian females from those pest populations. Islands. Thrips maculicollis Hood. As indi- Hood. This Asian cated in Mound et al. (2016) this species species is well established in Florida, as seems to be well established on Hawaii well as other localities around the Carib- island, although no specimens have been 26 Mound et al. seen from any of the other islands. It was direct feeding damage and through virus found at several sites on Hawaii in the transmission (Rosenheim et al. 1990). flowers of Fagraea berteriana [Genti- Similarly, the two widespread leaf-feeding anaceae] that are commonly used in the Chaetanaphothrips species (orchidii production of lei. and signipennis) are both damaging to various aroid plants including cultivated Discussion Anthurium, while feeding damage by C. The major purpose of the thrips sam- signipennis can lead to banana fruits being pling carried out in July 2016 was to obtain unmarketable (Hara et al. 2002) good series of the endemic species in the One exceptional adventive pest in recent genera Neurisothrips and . years has been Klambothrips myopori. Species-level in those two gen- This species originated in Tasmania, but era is currently unsatisfactory, with type is established and seriously damaging to material of several named species being in Myoporum spp. in California (Cameron very poor condition, but with some appar- and Mound 2014). On Hawaii island, this ently undescribed species present in muse- thrips has killed a large percentage of na- um collections. Because of this objective, tive Myoporum sandwicense in natural field work was primarily within areas of forests, as well as killed the prostrate natural vegetation, leading to a collection variety of this species that is used for land- of more than 50 Thysanoptera species scaping at lower elevations. Landscapers in the four weeks that field surveys were have had to replace this native species with conducted. Investigations into the range of other non-native dry tolerant species in Thysanoptera species on cultivated plants landscaping on this island. are clearly needed, and several Thripidae One problem with records of pest spe- species are known to cause problems to cies may lie in extensive dependency on horticulturalists in the Hawaiian Islands. insecticides, leading to a situation where For example, haemorrhoidalis a pest is not reported until it exhibits and rubrocinctus are both pesticide resistance. However, there are responsible for leaf damage on tree crops, several factors that could limit the nega- whereas Thrips nigropilosus can be severe tive impacts of adventive thrips in Hawaii, on hydroponic lettuce crops, and Thrips including suboptimal conditions both of parvispinus on the leaves and fruits of temperature, environmental conditions, papaya. However, the relationships of and host plants (Morse and Hoddle 2006), thrips species to cultivated plants, and to and biotic resistance in the form of com- the extensive areas of non-native plants petition from other thrips species (Fun- will require separate studies by locally derburk et al. 2016), or even population based entomologists in collaboration with suppression by resident guilds of natural the horticultural industry. enemies (Funderburk et al., 2000). One It is curious that several adventive spe- example of the latter appears to be the leaf cies recorded from these recent surveys damage to Ficus in landscapes and urban have not reached pest status in Hawaii, areas that is caused by ; even though they are problematic else- this seems to be under partial control by where (e.g., S. perseae on avocados; anthocorid bugs such as Montandoniola Pezothrips kellyanus on citrus). Notable confusa. Investigation into mechanisms exceptions are the invasive suppressing populations of adventive and F. occidentalis that are known to be thrips in Hawaii may provide useful in- pests of vegetable crops both through sights into factors influencing the invasion Adventive Thysanoptera in Hawaii 27 ecology of these tiny insects, and for the Hara, A.H., R.F.L. Mau, R. Heu, C. Jacob- development of ecologically-based man- son, and R. Niino-DuPonte. 2002. Banana agement plans for economically damaging rust thrips damage to banana and ornamen- species. tals in Hawaii. Honolulu (HI): University of Hawaii. 4 p. (Insect Pests; IP-10). Acknowledgments Hawaii Department of Agriculture. 2007. The visits to and studies on Hawaii in Hawaii Department of Agriculture Annual 2016 by three of the authors (MSH, LAM, Report for Fiscal Year 2006. Ed. Janelle Saneishi. Honolulu, HI: Hagadone Printing and AW) were facilitated by many people, Company. http://hdoa.hawaii.gov/meetings- including Rob Hollingsworth, Fran Cal- reports/annual-reports/ [accessed 1.iii.2017] vert, Karl Magnacca, Michael Sthreshley, Hoddle, M.S., S. Nakahara, and P.A. Phil- Stacey Chun, Robert Curtis, Linda Pratt, lips. 2002. Foreign exploration for Scirto- Paul Krushelnycky, Randy Hamasaki, and thrips perseae Nakahara (Thysanoptera: Dick Tsuda. Figure 1 was kindly provided Thripidae) and associated natural enemies by Stacey Chun. We are grateful to all of on avocado (Persea americana Miller). Biol. them for their input to these studies. Control 24: 251–265. Hoddle M.S., C.D. Hoddle, and L.A. Mound. 2008. An inventory of Thysanoptera col- Literature Cited lected from French Polynesia. Pac. Sci. 62: Beardsley, J.W. 1962. On Accidental Im- 509–515. migration and Establishment of Terrestrial Hollingsworth, R.G., F. Calvert, and A.H. Arthropods in Hawaii during Recent Years. Hara. 2012. Dichromothrips smithi (Zim- Proc. Hawaiian Entomol. Soc. 18: 99–109. mermann), a new thrips species infesting Beardsley, J. 1979. The current status of the bamboo orchids Arundina graminifolia names of Hawaiian aphids. Proc. Hawaii (D. Don) Hochr. and commercially grown Entomol. Soc. 13: 45–50. orchids in Hawaii. Proc. Hawaiian Entomol. Bhatti, J.S. 2000. Revision of Trichromothrips Soc. 44:1–9. and related genera (Terebrantia: Thripidae). Howarth, F.G., B.R. Kumashiro, and J.N. Oriental Ins. 34: 1–65. Matsunaga. 2013. Pathway analysis and Cameron, S.L., and L.A. Mound. 2014. dissemination of new insect and plant Trans-Bass Strait speciation and trans- disease records in Hawaii. Honolulu (HI): Pacific dispersal in the Myoporum thrips United States Department of Agriculture (Thysanoptera, Phlaeothripinae). Austral and Plant Health Inspection Ser- Entomol. 53: 36–41. vice Plant Protection and Quarantine Final Dang, L.-H., L.A. Mound, and G.-X. Qiao. Report. Hawaii Department of Agriculture 2013. Leaf-litter thrips of the genus Adra- Cooperative Agreement No. 11-8510-1485- neothrips from Asia and Australia (Thy- ca. 122 pp. sanoptera, Phlaeothripinae). Zootaxa 3716 Krushelnycky P.D., L.L. Loope, and R.G. (1): 001–021. Gillespie. 2007. Inventory of arthropods of Frank, H., and E.D. McCoy. 1995. Precinc- the west slope shrubland and alpine ecosys- tive insect species in Florida. Fla. Entomol. tems of Haleakala National Park. Honolulu 78: 21–35. (HI): Pacific Cooperative Studies Unit, Uni- Funderburk, J., J. Stavisky, and S.M. Olson. versity of Hawaii at Manoa, Department of 2000. Predation of Frankliniella occiden- Botany. PCSU Technical Report, 148. 52 pp. talis in field pepper by Orius insidiosus. Kumar, V., D.R. Seal, G. Kakkar, C.L. Environ. Entomol. 29: 376–382. McKenzie, and L.S. Osborne. 2012. New Funderburk, J., G. Frantz, C. Mellinger, tropical fruit hosts of Scirtothrips dorsalis K. Tyler-Julian, and M. Srivastava. 2016. (Thysanoptera: Thripidae) at its relative Biotic resistance limits the invasiveness abundance on them in south Florida. Fla. of the , Frankliniella Entomol. 95: 205–207. occidentalis (Thysanoptera: Thripidae), in Kumashiro, B.R., R.A. Heu, G.M. Nishida, Florida. Ins. Sci. 23: 175–182. and J.W. Beardsley. 2002. New state re- 28 Mound et al. cords of immigrant insects in the Hawaiian Thripidae) in Malaysia, with five new spe- Islands for the year 1999. Proc. Hawaiian cies and comments on Biltothrips, a related Entomol. Soc. 35: 170–184. genus. Zootaxa 3856 (2): 253–266. Masumoto, M., and S. Okajima. 2005. Ng, Y.F., and L.A. Mound. 2015. Genera of Trichromothrips Priesner (Thysanoptera, the Scirtothrips genus-group (Thysanop- Thripidae) of Japan and Taiwan, with de- tera, Thripidae) with a new species of scriptions of four new species and a review Siamothrips from Malaysia. Zootaxa 4021 of the Trichromothrips group of genera. (2): 387–394. Zootaxa 1082: 1–27. Nishida, G.M. 2002. Hawaiian terrestrial Messing, R.H., M.N. Tremblay, E.B. Mon- arthropod checklist: Fourth edition. Hawaii dor, R.G. Foottit, and K.S. Pike. 2007. Biological Survey, Bishop Museum Techni- Invasive aphids attack native Hawaiian cal Report No. 22. 313 p. Available at: http:// plants. Biol. Invas. 9: 601–607. hbs.bishopmuseum.org/pdf/tr22.pdf Morse, J.G., and M.S. Hoddle. 2006. Inva- Okajima, S. 1981. A revision of the tribe Plec- sion biology of thrips. Ann. Rev. Entomol. trothripini of fungus-feeding Thysanoptera 51: 67–89. (Phlaeothripidae: Phlaeothripinae). Syst. Mound, L.A, and M.S. Hoddle. 2016. The Entomol. 6: 291–336. Scirtothrips perseae species-group (Thy- Okajima, S. 1994. The genus Sophiothrips sanoptera), with one new species from Hood (Thysanoptera, Phlaeothripidae) from avocado, Persea americana. Zootaxa 4079 Japan. Japanese J. Entomol. 62: 29–39. (3): 388–392. Okajima, S., and M. Masumoto. 2014. Spe- Mound, L.A., and M. Masumoto. 2005. The cies-richness in the Oriental fungus-feeding genus Thrips (Thysanoptera, Thripidae) in thrips of the genus Azaleothrips (Thy- Australia, New Caledonia and New Zealand. sanoptera, Phlaeothripidae). Zootaxa 3846: Zootaxa 1020: 1–64. 301–347. Mound, L.A., and J.N. Matsunaga. 2017. The Okajima, S., and H. Urushihara. 1994. Re- species of (Thysanoptera, Ph- discovery of Indusiothrips nakaharai Wil- laeothripinae) and related genera recorded son from Japan (Thysanoptera, Thripi- from the Hawaiian Islands. ZooKeys 662: dae). Trans Shikoku Entomol. Soc. 20: 79–92. 97–101. Mound, L.A., S. Nakahara, and D.M. Tsuda. Rosenheim, J., S.C. Welter, M.W. Johnson, 2016. Thysanoptera-Terebrantia of the Ha- R.F.L. Mau, and L.R. Gusukuma-Minuto. waiian Islands: an identification manual. 1990. Direct feeding damage on cucumber ZooKeys 549: 71–126. by mixed species infestations of Thrips Mound, L.A., and S. Okajima. 2015. Taxo- palmi and Frankliniella occidentalis (Thy- nomic Studies on Dolichothrips (Thy- sanoptera: Thripidae). J. Econ. Entomol. 83: sanoptera: Phlaeothripinae), pollinators 1519–1525. of Macaranga trees in Southeast Asia ThripsWiki. 2017. ThripsWiki - providing (Euphorbiaceae). Zootaxa 3956 (1): 79–96. information on the World’s thrips. [accessed Australia-New Zealand connection re-vis- 6.iii.2017] ited, with two new species of Cartomothrips Wilson, T.H. 1975. A monograph of the sub- (Thysanoptera, Phalaeothripidae). Zootaxa Panchaetothripinae (Thysanoptera: 3487: 58–64. Thripidae). Mem. American Entomol. Inst. Ng, Y.F., L.A. Mound, and A.A. Azidah. 23: 1–354. 2014. The genus Scirtothrips (Thysanoptera: