Invasive Species Compendium

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Datasheet

Sophonia orientalis
(two-spotted leafhopper)

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Datasheet

Sophonia orientalis (two-spotted leafhopper)

Summary

  • Last modified
  • 15 July 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Sophonia orientalis
  • Preferred Common Name
  • two-spotted leafhopper
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • S. orientalis proved to be highly invasive on the Hawaiian Islands, although its populations decreased eventually, probably because of the impact of several natural enemies (
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  • Distribution map More information

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Identity

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Preferred Scientific Name

  • Sophonia orientalis (Matsumura)

Preferred Common Name

  • two-spotted leafhopper

Other Scientific Names

  • Nirvana orientalis Matsumura, 1912
  • Pseudonirvana rufofascia Kuoh and Kuoh, 1983
  • Sophonia rufofascia (Kuoh and Kuoh)

International Common Names

  • English: leafhopper, two-spotted

Summary of Invasiveness

Top of page S. orientalis proved to be highly invasive on the Hawaiian Islands, although its populations decreased eventually, probably because of the impact of several natural enemies (Alyokhin et al., 2001). Nevertheless, it still remains a significant problem. Outside of Hawaii, a resident, breeding population of S. orientalis was found in Los Angeles County in California, USA (Garrison, 1996). However, its density remains fairly low and this species has not (yet?) become invasive in continental USA. Similarly, S. orientalis was collected in 1999 in Tahiti (Polhemus, 2001), but there were no reports on its invasiveness on that island. There are no records of it being invasive in its native regions.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Hemiptera
  •                         Suborder: Auchenorrhyncha
  •                             Unknown: Cicadelloidea
  •                                 Family: Cicadellidae
  •                                     Genus: Sophonia
  •                                         Species: Sophonia orientalis

Notes on Taxonomy and Nomenclature

Top of page Sophonia orientalis appears to have been described at least four times in various areas of South and South-East Asia as Nirvana orientalis (Matsumura, 1912), Quercinirvana longicephala (Ahmed and Mahmood, 1970), Pseudonirvana rufofascia (Kuoh and Kuoh, 1983) and Sophonia rufofascia (Huang, 1994). Until recently, the species was called S. rufofascia Kuoh and Kuoh following revisions by Viraktamath and Wesley (1988) and Viraktamath (1992). Most literature referred to the species as S. rufofascia (Kuoh and Kuoh), but recently it has been shown that previous records of this species correspond to S. orientalis (Matsumura). Additional investigations are needed to determine if what is presently known as S. orientalis is indeed a single species, or a complex of several closely related species that are similar in appearance.

Description

Top of page The adult is approximately 5 mm long, primarily light-green or pale-yellow, and has a bold black longitudinal stripe down the centre of its body. A black spot occurs at the tip of each of the forewings. The nymphs are smaller, uniformly green or light-yellow, and have a pair of small black spots at the hind end of the abdomen. The eggs are hyaline to milky-white, elongate and slightly curved, with a whitish operculum, approximately 1.41 mm long and 0.30 mm at the greatest width (Culliney, 1998).

Distribution

Top of page S. orientalis is native to, and widely distributed in, South and South-East Asia. Following inadvertent introductions, it is currently present on all major Hawaiian Islands, on Tahiti (French Polynesia) and in southern California, USA, and has recently been recorded on Madeira and the Canary Islands.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ChinaPresentPresent based on regional distribution.
-GuizhouPresentNative Not invasive Kuoh and Kuoh, 1983
-ZhejiangPresentNative Not invasive Kuoh and Kuoh, 1983
IndiaPresentNative Not invasive Ahmed and Mahmood, 1969
JapanPresentNative Not invasive Matsumura, 1912
PakistanPresentNative Not invasive Ahmed and Mahmood, 1969
SingaporePresentNative Not invasive Huang, 1994
TaiwanPresentNative Not invasive Huang, 1994

Africa

Spain
-Canary IslandsPresentIntroducedAguin-Pombo et al., 2007

North America

USAPresentPresent based on regional distribution.
-CaliforniaPresentIntroduced1996 Not invasive Garrison, 1996
-HawaiiPresentIntroduced1987 Invasive Heu and Kumashiro, 1989

Europe

PortugalPresentPresent based on regional distribution.
-MadeiraPresentIntroducedAguin-Pombo et al., 2007
SpainPresentPresent based on regional distribution.

Oceania

French PolynesiaPresentIntroduced1999 Not invasive Polhemus, 2001

History of Introduction and Spread

Top of page S. orientalis was first detected in Hawaii, North America on Oahu Island in June 1987 (Heu and Kumashiro, 1989) and subsequently spread to the other major islands over the next 5 years. After that, it was recorded in 1996 in Los Angeles County in California, USA (Garrison, 1996) and in 1999 on Tahiti (Polhemus, 2001). It was recorded from the northeastern Atlantic archipelagos of Madeira and Canaries (Aguin Pombo et al., 2007) in 2000.  

Risk of Introduction

Top of page The small size and concealed nature of S. orientalis eggs make their detection on imported plant material very difficult. The risk of introduction is therefore fairly high.

Habitat

Top of page S. orientalis has been recorded from a wide variety of habitats (Fukada, 1996). Nevertheless, in a survey by Alyokhin et al. (2001), the leafhoppers appeared to have a preference for wetter, closed habitats rather than drier, open ones. Wild plants had a higher leafhopper infestation than cultivated plants. More eggs were found on the foliage from closed natural habitats than from open natural habitats. There was no difference in egg density between dry and mesic open natural habitats, but wet closed habitats harboured more eggs than mesic closed habitats. The type of mesic closed habitat also appeared to be an important factor, with habitats that had a shrub understorey harbouring more leafhopper eggs than habitats that had a fern understorey. The presence of preferred host plants may increase leafhopper density compared to otherwise very similar habitats (Lenz and Taylor, 2001).

Hosts/Species Affected

Top of page S. orientalis is extremely polyphagous. Its nymphs develop on over 300 species of host plants in 83 families (Fukada, 1996). However, not all these hosts are equally preferred by the leafhoppers. For example, in a survey by Alyokhin et al. (2001), egg density varied from 227.83 ± 82.22 eggs per m² on the most preferred host, Hawaiian tree fern (Cibotium chamissoi) to 2.93 ± 1.10 on the least preferred host, grapefruit (Citrus paradisi). Similarly, Lenz and Taylor (2001) captured significantly more leafhoppers on the nitrogen-fixing Myrica faya [Morella faya] that had glabrous foliage compared to the non-nitrogen-fixing 'ohi'a lehua tree (Metrosideros polymorpha) that had hirsute foliage. On the Hawaiian Islands, where most work has been carried out, the impact of the leafhoppers on some host species has received more attention than on other species because of their economic, ecological or conservation significance. This might create a potentially misleading impression that those species are most affected by S. orientalis. In fact, little is known about the host preference of this insect.

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Acacia koa (koa)FabaceaeWild host
Aleurites moluccanus (candlenut tree)EuphorbiaceaeWild host
Annona muricata (soursop)AnnonaceaeOther
Artocarpus altilis (breadfruit)MoraceaeOther
Averrhoa carambola (carambola)OxalidaceaeOther
Bidens pilosa (blackjack)AsteraceaeWild host
Capsicum annuum (bell pepper)SolanaceaeOther
Cibotium chamissoiDicksoniaceaeWild host
Cinnamomum verum (cinnamon)LauraceaeOther
Citharexylum spinosumVerbenaceaeOther
Citrus aurantiifolia (lime)RutaceaeOther
Citrus limon (lemon)RutaceaeOther
Citrus sinensis (navel orange)RutaceaeOther
Coffea arabica (arabica coffee)RubiaceaeOther
Colocasia esculenta (taro)AraceaeOther
Cordyline fruticosa (ti plant)AgavaceaeWild host
CrotalariaFabaceaeWild host
Desmodium (tick clovers)FabaceaeWild host
Dicranopteris linearisGleicheniaceaeWild host
Dodonaea eriocarpaSapindaceaeWild host
Erythrina sandwicensisFabaceaeOther
Euphorbia pulcherrima (poinsettia)EuphorbiaceaeOther
FicusMoraceaeWild host
Fragaria vesca (wild strawberry)RosaceaeOther
Garcinia mangostana (mangosteen)ClusiaceaeOther
Hibiscus rosa-sinensis (China-rose)MalvaceaeOther
Ipomoea batatas (sweet potato)ConvolvulaceaeOther
Ipomoea batatas (sweet potato)ConvolvulaceaeOther
Macadamia integrifolia (macadamia nut)ProteaceaeOther
Malus sylvestris (crab-apple tree)RosaceaeOther
Mangifera indica (mango)AnacardiaceaeOther
Manihot esculenta (cassava)EuphorbiaceaeOther
Metrosideros collinaMyrtaceaeWild host
Metrosideros polymorphaMyrtaceaeWild host
Morella faya (firetree)MyricaceaeWild host
Morus (mulberrytree)MoraceaeWild host
Murraya paniculata (orange jessamine)RutaceaeOther
Musa (banana)MusaceaeOther
Persea americana (avocado)LauraceaeOther
PittosporumPittosporaceaeOther
Plumeria rubra (red frangipani)ApocynaceaeWild host
Psidium cattleianum (strawberry guava)MyrtaceaeWild host
Psidium guajava (guava)MyrtaceaeOther
Quercus floribunda (green oak)FagaceaeWild host
Rhododendron (Azalea)EricaceaeWild host
Ricinus communis (castor bean)EuphorbiaceaeOther
Schefflera actinophylla (umbrella tree)AraliaceaeWild host
Spathodea campanulata (African tulip tree)BignoniaceaeOther
Styphelia tameiameiae (hawaiian hether)Wild host
Zea mays (maize)PoaceaeOther

Growth Stages

Top of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

Symptoms

Top of page The feeding of S. orientalis produces interveinal chlorosis and vein browning that can eventually result in the necrosis of affected leaves. It also decreases stem length and diameter, the number of nodes, leaf area and the net carbon dioxide assimilation (Jones et al., 2000; Lenz, 2000). Leafhopper oviposition can also cause the death of distal leaf tissue by interrupting the flow of water and nutrients (Culliney, 1998; Jones et al., 2000). The feeding damage is localized to the feeding sites, suggesting that the factor causing the damage (i.e. the toxins present in the leafhopper saliva) is not translocated in plant tissues (Jones et al., 2000). Microscopic examination of sections through the damaged areas of several host plants showed vascular bundle abnormalities (Jones et al., 2000). Also, leafhopper oviposition into leaf mid-veins blocks up to one-third of the vascular bundle and disrupts both the phloem and xylem (Jones et al., 2000). Not surprisingly, affected trees suffer a higher level of moisture stress (Lenz, 2000).

List of Symptoms/Signs

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SignLife StagesType
Leaves / abnormal colours
Leaves / necrotic areas
Whole plant / discoloration
Whole plant / dwarfing
Whole plant / external feeding
Whole plant / plant dead; dieback

Biology and Ecology

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Physiology and Phenology

All the information on this subject was obtained on the Hawaiian Islands, North America. Therefore, it is possible that the leafhopper phenology will be different in the areas with more pronounced seasonal changes. Under conditions typical to the Hawaiian winter (18.0-23.5ºC and 81.3-97.7% RH), it took S. orientalis between 3 and 4 months to complete a single generation (Duan and Messing, 2000). Because of the long oviposition and egg-hatching periods (approximately 10 and 4 weeks, respectively), it is likely that S. orientalis has overlapping generations (Duan and Messing, 2000). The adult leafhoppers were captured by yellow sticky cards throughout the year in a number of habitats on the islands of Kauai and Hawaii. However, their densities were usually highest during the summer months (Lenz and Taylor, 2001; Yang et al., 2002).

Reproductive Biology

S. orientalis is similar to many other cicadellid leafhoppers in that it reproduces sexually and is oviparous. In the laboratory at 23 ± 1.5ºC, adult S. orientalis did not mate until 7-9 days after eclosion. The proportion of mating pairs increased from 10 to 100% as age increased from 9 to 15 days. A complete mating process in S. orientalis consisted of mounting, back-to-back copulation, and postmating rest phases, which lasted for a mean ± SE period of 0.7 ± 0.04, 64.7 ± 1.20 and 3.5 ± 0.39 minutes, respectively (Duan and Messing, 2000). A single female produces approximately 30-35 eggs throughout her life span (Duan and Messing, 2000). Females insert their eggs on the underside (abaxial surface) of the foliage, usually at the juncture of the midrib and lamina, sometimes directly in the midrib and often near the intersection of the midrib and a cross-vein (Culliney, 1998; Yang et al., 2000). The eggs are usually located in the pith of the midrib with their long axis aligned with the axis of the vein. They may be positioned either with the operculum protruding slightly from the oviposition scar, or entirely concealed just below the scar surface (Culliney, 1998). On plants with thicker leaves, some of the eggs may be deposited away from the midrib (Yang et al., 2000). A variation in egg position relative to the basal, middle and apical third of the leaf also exists, but usually the middle third of the leaf is the most common oviposition site (Yang et al., 2000). When the adults were caged onto individual shoots of host plants [Cordyline terminalis (ti) and Psidium guajava (guava)] growing outdoors at 18.0-23.5ºC and 81.3-97.7% RH, they did not lay eggs until the second week after emergence. The oviposition period lasted 8-9 weeks, and the weekly oviposition rate peaked 4-5 weeks after emergence (9.8 ± 2.9 eggs per female on ti and 8.1 ± 0.9 eggs per female on guava). There was no significant difference in the total number of eggs laid per female on guava versus ti plants. Under outdoor ambient conditions, the eggs of S. rufofascia did not begin to hatch until 4 weeks after oviposition, and the peak rate of hatch occurred 6 weeks after oviposition. The newly hatched nymphs took a mean of 56.9 ± 2.8 days to complete development to the adult stage on guava and 47.2 ± 1.7 days on ti plants (Duan and Messing, 2000).

Environmental Requirements

S. orientalis is widely distributed in areas with a warm climate, ranging from Pakistan to California, USA. In Hawaii it has been collected from the sea level up to 1460 m (Fukada, 1996). It is not known to occur in areas with colder climates. Therefore, it appears that this species can survive under a variety of environmental conditions, as long as it does not get too cold. However, its exact environmental tolerances and preferences are unknown.

Associations

Jones et al. (1998) demonstrated that S. orientalis did not vector phytoplasmas in guava plantations in Hawaii. It is also not known to form any other interspecific associations.

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Chaetomymar sophoniae Parasite Eggs
Polynema Parasite Eggs
Schizophragma bicolor Parasite Eggs

Notes on Natural Enemies

Top of page A series of surveys conducted in Hawaii (Yang et al., 2000, 2002; Alyokhin et al., 2001; Johnson et al., 2001) revealed a fairly robust complex of generalist egg parasitoids (Hymenoptera: Mymaridae). In the survey by Alyokhin et al. (2001), approximately 40% of S. orientalis eggs, averaged over all host species and locations surveyed, were parasitized by Chaetomymar sophoniae, Schizophragma bicolor and Polynema sp. The percentage of parasitism varied widely among different plant species and habitats. Ch. sophoniae was the most abundant and widely distributed species, but the parasitoid guild varied depending on plant and habitat. Ch. sophoniae is a new species described by Huber (2003). It was previously called Chaetomymar sp. nr. bagichi (Alyokhin et al., 2001) or Chaetomymar sp. (Johnson et al., 2001). It is a new, adventive species in Hawaii that probably arrived from Asia together with S. orientalis. S. bicolor is native to North America and was first collected in Hawaii in 1963 (Huber, 1987; Beardsley and Huber, 2000). Polynema sp. is native to Hawaii, where it is known to attack the eggs of a number of native leafhopper species (Beardsley and Huber, 2000). Therefore, the parasitism of two-spotted leafhopper eggs represents a host range expansion for both S. bicolor and Polynema sp. (Johnson et al., 2001).

Means of Movement and Dispersal

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Natural Dispersal

The adult leafhoppers have fully developed wings and readily fly. The extent of their natural dispersal has never been documented, but it is likely that they can be blown by the wind over considerable distances.

Movement in Trade

All life stages can be readily moved on vegetative plant materials. A number of recent interceptions on imported plants in California, USA (Pylman and Saunders, 2000; Sokulsky and Chen, 2001) confirm the probability of such a mode of introduction.

Plant Trade

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Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Growing medium accompanying plants
Leaves
Roots
True seeds (inc. grain)
Wood

Wood Packaging

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Wood Packaging not known to carry the pest in trade/transport
Loose wood packing material
Non-wood
Processed or treated wood
Solid wood packing material with bark
Solid wood packing material without bark

Impact Summary

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CategoryImpact
Animal/plant collections None
Animal/plant products None
Biodiversity (generally) Negative
Crop production Negative
Environment (generally) Negative
Fisheries / aquaculture None
Forestry production Negative
Human health None
Livestock production None
Native fauna None
Native flora Negative
Rare/protected species Negative
Tourism None
Trade/international relations None
Transport/travel None

Impact

Top of page S. orientalis damage can result in significant economic losses. In a study by Jones et al. (1998), feeding and oviposition by this species caused 'guava yellowing', which reduced yields by 23% compared to pesticide-protected trees. Also, S. orientalis has been implicated in killing large patches of uluhe (Dicranopteris linearis), a native Hawaiian fern that is an important ground cover of agricultural watersheds (Palmer, 1993). This left bare ground subject to erosion, which in some cases was so severe that it affected agricultural irrigation systems that relied on watershed rivers (Duan and Messing, 2001). However, while proving that S. orientalis feeding is indeed detrimental to this fern, Jones et al. (2000) failed to find a significant correlation between leafhopper presence and the wide-scale death of uluhe in natural habitats.

Environmental Impact

Top of page Leafhopper feeding may have a detrimental effect on multiple species of wild plants. Among over 300 plants identified as S. orientalis hosts in Hawaii by Fukada (1996), 22% were endemic to the Hawaiian Islands (including 14 rare and endangered species). Also, leafhopper feeding may represent a selection force that favours certain plant species or phenotypes (e.g. because of foliar pubescence), causing corresponding changes in the community structure.

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Doryopteris takeuchii (Takeuch's lipfern)NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiHerbivory/grazing/browsingUS Fish and Wildlife Service, 2013b
Dracaena forbesii (hala pepe)USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiHerbivory/grazing/browsingUS Fish and Wildlife Service, 2013a
Loxioides bailleui (palila)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resourcesUS Fish and Wildlife Service, 2006
Pritchardia lanigeraEN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiHerbivory/grazing/browsingUS Fish and Wildlife Service, 2013a
Santalum freycinetianum var. lanaienseNo DetailsHawaiiHerbivory/grazing/browsingUS Fish and Wildlife Service, 2011

Risk and Impact Factors

Top of page Impact mechanisms
  • Herbivory/grazing/browsing

Detection and Inspection

Top of page Both the nymphs and adults can be easily detected by visual inspections of the plant foliage. The adult populations have also been successfully monitored using yellow sticky aphid/whitefly cards (Lenz and Taylor, 2001; Yang et al., 2002). Oviposition scars may be visible to the naked eye on some, but not all, host plants (Culliney, 1998). Yang et al. (2000) developed a more reliable method for detecting S. orientalis eggs using backlighting of excised leaves by a fibreoptic microscope light source. Also, excised foliage containing S. orientalis eggs can be successfully incubated in plastic bags in the laboratory until nymphal emergence (Alyokhin et al., 2001).

Similarities to Other Species/Conditions

Top of page The symptoms of damage caused by S. orientalis are fairly general (hopperburn) and are very similar to those caused by other leafhopper species.

Prevention and Control

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Phytosanitary Measures

Quarantine inspections may help to prevent S. orientalis introductions into new areas.

Host-Plant Resistance

Foliar pubescence appears to make plants significantly less suitable as S. orientalis hosts (Fukada, 1996; Jones et al., 2000; Lenz and Taylor, 2001).

Biological Control

A complex of two inadvertently introduced (Chaetomymar sophoniae and Schizophragma bicolor) and one or more native (Polynema sp.) egg parasitoids parasitized approximately 40% of S. orientalis eggs in Hawaii, North America. Percent parasitism increased dramatically between 1995 and 2000, and was accompanied by a substantial reduction in leafhopper densities (Yang et al., 2000, 2002; Alyokhin et al., 2001; Johnson et al., 2001). Since its first appearance in Hawaii, Ch. sophoniae rapidly expanded its range and began to dominate the parasitoid complex attacking S. orientalis. It is not known if the parasitoids were solely responsible for the observed suppression of S. orientalis, but it is likely that their contribution was substantial. Unfortunately, all the species have a fairly wide host range. Therefore, their possible future introductions into new areas for S. orientalis control should be carefully weighed against possible non-target effects on native leafhopper species.

Chemical Control

Chemical control appears to be highly efficient in controlling S. orientalis, with substantial reductions in both leafhopper populations and damage observed following the applications of a rotenone-azadiractin tank mix (Jones et al., 1998), orthene (Lenz, 2000) and imidacloprid (Follett et al., 2000).

IPM Programmes

No IPM programmes have been developed to specifically target S. orientalis.

References

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Aguin-Pombo D, Aguiar AMF, Kuznetsova VG, 2007. Bionomics and taxonomy of leafhopper Sophonia orientalis (Homoptera: Cicadellidae), a pacific pest species in the Macaronesian Archipelagos. Annals of the Entomological Society of America, 100(1):19-26. http://esa.publisher.ingentaconnect.com/content/esa/aesa/2007/00000100/00000001/art00004

Ahmed M, Mahmood SH, 1970. A new genus and two new species of Nirvaninae (Cicadellidae:Homoptera) from Pakistan. Pakistan Journal of Scientific and Industrial Research, 12:260-263.

Alyokhin AV, Yang PJ, Messing RH, 2001. Distribution and parasitism of Sophonia rufofascia (Homoptera: Cicadellidae) eggs in Hawaii. Annals of the Entomological Society of America, 94(5):664-669; 23 ref.

Alyokhin AV, Yang PJ, Messing RH, 2004. Oviposition of the invasive two-spotted leafhopper on an endemic tree: effects of an alien weed, foliar pubescence, and habitat humidity. Journal of Insect Science (Tucson), 4:13. http://www.insectscience.org/4.13/

Beardsley JW, Huber JT, 2000. Key to genera of Mymaridae in the Hawaiian islands, with notes on some species (Hymenoptera: Chalcidoidea). Proceedings of the Hawaiian Entomological Society, 34:1-22.

Culliney TW, 1998. Site of oviposition and description of eggs of Sophonia rufofascia (Homoptera: Cicadellidae: Nirvaninae), a polyphagous pest in Hawaii. Proceedings of the Hawaiian Entomological Society, 33:67-73.

Duan JJ, Messing RH, 2000. Mating, oviposition, and development of Sophonia rufofascia (Homoptera: Cicadellidae) in Hawaii. Annals of the Entomological Society of America, 93(3):554-558; 15 ref.

Follett PA, Empy-Campora C, Jones VP, 2000. Imidachloprid as a protectant for endangered plants attacked by Sophonia rufofascia. Proceedings of the Hawaiian Entomological Society, 34:199-201.

Fukada M, 1996. Distribution, host range, and seasonal abundance of the two spotted leafhopper, Sophonia rufofascia (Kuoh and Kuoh) in Hawaii. MSc thesis. Honolulu: University of Hawaii at Manoa.

Garrison RW, 1996. New agricultural pest for Southern California: two-spotted leafhopper (Sophonia rufofascia). California Pest and Disease Report, 15:6-7.

Heu R, Kumashiro B, 1989. Notes and exhibition. Proceedings of the Hawaiian Entomological Society, 29:16-17.

Huang KW, 1994. Supplement of Nirvanini of Taiwan (Homoptera: Cicadellidae: Nirvaninae). Chinese Journal of Entomology, 14:83-88.

Huber JT, 1987. Review of Schizophragma Ogloblin and the non-Australian species of Stethynium Enock (Hymenoptera: Mymaridae). Canadian Entomologist, 119(9):823-855

Huber JT, 2003. Review of Chaetomymar Ogloblin, with description of a new species in the Hawaiian Islands (Hymenoptera: Mymaridae). Journal of Hymenoptera Research, 12(1):77-101; 36 ref.

Johnson MT, Yang P, Huber JT, Jones VP, 2001. Egg parasitoids of Sophonia rufofascia (Homoptera: Cicadellidae) in Hawaii Volcanoes National Park. Biological Control, 22(1):9-15; 27 ref.

Jones VP, Anderson-Wong P, Follett PA, Yang PingJun, Westcot DM, Hu JS, Ullman DE, 2000. Feeding damage of the introduced leafhopper Sophonia rufofascia (Homoptera: Cicadellidae) to plants in forests and watersheds of the Hawaiian Islands. Environmental Entomology, 29(2):171-180; 20 ref.

Jones VP, Follett PA, Messing RH, Borth WB, Hu JS, Ullman DE, 1998. Effect of Sophonia rufofascia (Homoptera: Cicadellidae) on guava production in Hawaii. Journal of Economic Entomology, 91(3):693-698; 20 ref.

Kuoh CL, Kuoh JL, 1983. New species of Pseudonirvana (Homoptera: Nirvanidae). Acta Entomologica Sinica, 26(3):316-325

Lenz L, Taylor JA, 2001. The influence of an invasive tree species (Myrica faya) on the abundance of an alien insect (Sophonia rufofascia) in Hawai'i Volcanoes National Park. Biological Conservation, 102:301-307.

Lenz, LS, 2000. The dieback of an invasive tree in Hawaii: interactions between the two-spotted leafhopper (Sophonia rufofascia) and faya tree (Myrica faya). MSc Thesis, Honolulu: University of Hawaii at Manoa.

Matsumura S, 1912. Die Acocephalinen und Bythoscopinen Japans. Journal of Sapporo College of Agriculture, 4:279-325.

Palmer DD, 1993. Hawaiian ferns under attack. Fiddlehead, 20:37.

Polhemus D, 2001. The first record of Sophonia rufofascia (Homoptera: Cicadellidae) in Tahiti. Proceedings of the Hawaiian Entomological Society, 35:16-17.

Pylman J, Saunders R, 2000. Sacramento County 2000 Crop & Livestock Report. Sacramento County: Agricultural Commissioner's Office.

Sokulsky R, Chen J, 2001. 2001 Crop and Livestock Report. Los Angeles, USA: County of Los Angeles Agricultural Commissioner / Weights and Measures Department.

US Fish and Wildlife Service, 2006. In: Revised Recovery Plan for Hawaiian Forest Birds. US Fish and Wildlife Service, 622 pp..

US Fish and Wildlife Service, 2011. In: Santalum freycinetianum var. lanaiense, Lanai sandalwood ('iliahi). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 19 pp..

US Fish and Wildlife Service, 2013. In: Endangered and Threatened Wildlife and Plants; Determination of Endangered Species Status for 15 Species on Hawaii Island; Final Rule. 78(209) US Fish and Wildlife Service, 64638-64690. https://www.gpo.gov/fdsys/pkg/FR-2013-10-29/pdf/2013-24103.pdf

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