Final report: 07-IG-11272177-051

Preliminary exploration for natural enemies of
Rubus ellipticus in China
By
Jianqing Ding, Kai Wu & Jialiang Zhang
Invasion Biology and Biocontrol Lab
Wuhan Botanical Institute
Chinese Academy of Sciences, Wuhan, Hubei Province, 430074 China
Email: ding@wbgcas.cn
In collaboration withTracy Johnson
Institute of Pacific Islands Forestry
USDA Forest Service, Pacific Southwest Research Station
P.O. Box 236, Volcano, Hawaii 96785
Ph: 808-967-7122
Fax: 808-967-7158
E-mail: tracyjohnson@fs.fed.us

Abstract
The Yellow Himalayan Raspberry, Rubus ellipticus, is an invasive plant in Hawaii. As
chemical, physical and manual controls are expensive and difficult to implement against this plant,
biological control is being considered. The collaboration between China and the U.S. for finding
potential biological control agents in the native range of R. ellipticus was recently reinitiated in2006. Here, we report 60 arthropod species in 30 families that were directly collected on Rubus
ellipticus in field surveys in 2006-2008. We also provide a review of the potential agents,
including 49 species of arthropods in 16 families and 65 species of fungi in 3 phyla and 19
families, from literature or online data. Among these species, the warty beetles Chlamisus setosus(Bowditch) and Chlamisus spp., the flea beetles Chaetoenema, an unidentified stem borer, theleaf-rolling moth Epinotia spp. and an unidentified sawfly were the most promising potential
agents. Preliminary lab tests indicated that the warty beetles Chlamisus spp., the flea beetles
Chaetoenema, may have narrow host range. We recommend further screening of these organisms
to investigate their impacts on the target plant, host specificity, and the risk of undesired effects in
Hawaiian ecosystems.

Introduction
Rubus ellipticus, yellow Himalayan raspberry, is a serious invasive plant in Hawaii, posing a great
threat to local ecosystems. It was listed as one of the 100 World's worst invasive alien species by
IUCN. Mechanical and herbicidal control has been performed on a limited scale, but large scale
eradication is not feasible, because of the extensiveness and rugged terrain of the infested area.
The plant is native to the Himalayan region in Asia, including China, Bhutan, India, Laos,
Myanmar, Nepal, Pakistan, Philippines, Sikkim, Sri Lanka, Thailand and Vietnam.
Rubus is one of the largest genera in Rosaceae, consisting about 700 species worldwide.
According to Gu et al (1996), 201 species grow in China. They are distributed in 27 Provinces, butaround 70% of these species occur in southwestern China. Rubus ellipticus is distributed in
Guangxi, Guizhou, Sichuan, Tibet, and Yunnan. It is a small deciduous shrubby plant about 1-2 m
in height. The plant is perennial, reproducing by both vegetative means and seeds. It blooms fromMarch to April and fruits from April through May. The plant grows along hillsides, roadsides or
thickets in valleys, sparse forest, and broad-leaf forest at altitudes of 300-2000 m.. Twigs andleaves are used medicinally, and fruits are eaten.
Previous biological control efforts by Dr. Don Gardner (USGS PIERC, Honolulu) focused on
pathogens for R. ellipticus, but he retired and no promising pathogenic agent was screened. Dr.
Gardner suggested that continued efforts should be put on insects associated with the plant in its
native areas.
With funding from the state of Hawaii and USDA Forest Service, efforts for survey of and study
on natural enemies were conducted in China in since spring 2006. Specifically, the objectives ofthis project are to: (1) identify prospective biocontrol agents and source areas from Chinese
literature, (2) to locate source populations of Rubus ellipticus and associated arthropods insouthwestern China; (3) evaluate life histories and biocontrol potential of 1-2 selected insect
species; and (4) identify collaborators and plan in-depth studies at field sites in China.

Figure 1. Distribution of Rubus ellipticus in China (in grey)

Potential biocontrol agents and source areas identified from Chinese literatureand field surveys
We conducted a thorough literature search for information on Rubus ellipticus and its associatednatural enemies in China. We collected information from international databases, such as BIOSIS,
CAB, and major Chinese databases (in Chinese), and Chinese books on forest and agricultural
fauna, as well as internet information. A total of 49 arthropod species in 16 families have beenrecorded in literature as associated with Rubus in its Asian range (Table 1).
Table 1. Literature review: species of phytophagous insects on the genus Rubus in East Asia
Order : FamilySpecies

Feeds onLocationa

Host Range

References

LeavesYN, SC (China); JapanM(Lei and Zhou 1998)

Coleoptera
Buprestidae

Coraebus quadriundulatus Motschulsky

Chrysomelidae

Aphthona howenchuni (Chen)

LeavesChina

O(Yu et al. 1996)

Batophila impressa Wang

LeavesChina

O(Yu et al. 1996)

Chaetocnema simplicifrons (Baly)

LeavesChina

O(Yu et al. 1996)

Phaedon fulvescens Weise

LeavesChina; VietnamO(Yu et al. 1996)

Curculionidae

Enaptorrhinus convexiusculus Heller

Leaves

HeN, YN, ShX (China)P(Zhao and Chen
1980)

Eumolpidae

Basilepta leechi (Jacoby)

LeavesJS, ZJ, HB, JX, FJ, GD, GX, SC, GZ, YN(China); Vietnam

P(Tan et al. 1980)

Basilepta ruficolle (Jacoby)

LeavesZJ, HB, FJ, GX, SC, GZ, YN (China)

P(Tan et al. 1980)

Chlamisus indicus Jacoby

LeavesGX, SC, YN (China); India

O(Tan et al. 1980)

Chlamisus latiusculus Chûjô

LeavesFJ, TW, HaiN, GX (China)

M(R. amphidasys)(Tan et al. 1980)

Order : Family

HispidaeFeeds onLocationa

Host Range

References

Chlamisus ruficeps (Chen)

LeavesFJ, GX (China)

P(Tan et al. 1980)

Chlamisus semirufus (Chen)

LeavesJX, FJ, GD, GX (China); Vietnam

P(Tan et al. 1980)

Chlamisus setosus (Bowditch)

LeavesYN(China)

M(R. ellipticus)(Tan et al. 1980)

Alledoya vespertina (Boheman)

LeavesHB, BJ, TW (China); Korea; JapanP(HIRANO 1998; Lei
and Zhou 1998)

JL (China),P(Zhang 1985)

HB, ZJ, GS, TW (China)P(Zhang 1985)

P(Zhang 1985)

M(R. caesius)O(R. caesius,
R. fruticosus)(Aslan and Uygun
2005; Database 2007)

SpeciesHemiptera
CoreidaeDerepteryx fuliginosa (Uhler)
Derepteryx lunata (Distant)Pentatomidae

Leaves,stems
Leaves,stems

Amyntor obscurus (Dallas)

LeavesChina

Amphorophora rubi (Kaltenbach, 1843)

Unknown

Turkey; UK; Finland; Belgium, US; Canada

Aphis ruborum (Borner, 1932)

Leaves,flowers,
shootsTurkey; Europe

Acyrthosiphon rubiformosanum(Takahashi)

LeavesChina

Dysstroma cinereata (Moore)

LeavesDysstroma citrata (L.)

LeavesMesoleuca albicillata (L.)

LeavesPhotoscotosia miniosata (Walker)

LeavesPlagodis dolabraria (L.)

LeavesAbraximorpha davidii (Mabille)

LeavesSinthusa chandrana (Moore)

Leaves

Homoptera
Aphididae(Aslan and Uygun
2005)

P

(Zhang and Zhong
1983)

M

(Xue and Zhu 1999)

P(Xue and Zhu 1999)

Lepidoptera
Geometridae

Hesperiidae

Lycaenidae
Noctuidae

Nymphalidae

HN, JX, TW, SC, YN (China); Japan;
Korea; India; Burma; Sikkim; Bhutan
EU; Urals; Caucasus; Transcaucasia;
N.Kazakhstan; Mongolia; South Siberia;
Amur; Sakhalin; Kurils; China; Korea;
Japan; Canada; North US
North China; Korea; Austria; Czech
Republic; Denmark; Finland; France;
Germany; Ireland; Italy; Japan; Latvia;
Luxembourg; Moldova; Norway; Romania;
Sweden; United Kingdom
HeN, GS, HN, TW, SC, GZ, YN, TB(China); India; Sikkim; Pakistan;Philippines
GS, SC, HB, HN ZJ, JS (China); Austria;
Czech Republic; Denmark; Finland;
France; Germany; Ireland ; Italy; Japan;
Latvia; Luxembourg; Norway; Portugal ;Romania; Slovenia; South Korea; Sweden;
United Kingdom
ShX, HeN, HB, HN, JX, ZJ, SC, GD,
HaiN, YN, TW (China); Burma; Vietnam;
Indonesia
JX, ZJ, TW (China); Sikkim; Burma;
Thailand; Vietnam; Singapore; Indonesia

P

M(R. ellipticus)(Xue and Zhu 1999)

O(Sun et al. 1998)

M(Zhou 1994)

M

LeavesSC, HB, GZ, XJ (China); Korea; Japan;
Turkey; Russia; Syria; EuropeP

Anaplectoides prasina (Denis &Schiffermüller)

Leaves

HLJ, NM, XJ, HeN (China); Japan; EuropeP

Anomis mesogona (Walker)

Leaves,fruitsGrammodes geometrica (Fabricius)

Leaves,fruitsGrammodes stolida (Fabricius)

LeavesSypnoides picta Butler

Leaves

HeB, HLJ, HB, ZJ, (China); India;
Malaysia; Japan; Korea
ZJ, TW, GD, SC, HB (China ); Thailand;
India; Burma; Singapore; Sri Lanka;
Europe; Africa; Australia
YN (China); India; Burma; Iran; South
Europe; Africa - Oriental Region;
Australia.
HLJ, LN, ZJ, HN, YN (China); North
Korea; Japan

Argynnis paphia (L.)

LeavesAcronicta rumicis (L.)

Brenthis daphne (Denis &Schiffermüller)

LeavesBrenthis ino (Rottemburg)

LeavesSaturniidae

Loepa damaritis Jordan

Unknown

Tortricidae

Adoxophyes orana Fischer von
RöslerstammBuds,
leavesBuds,
leavesBuds,
flowers,
fruitAncylis comptana (Frölich)
Archips xylosteana (L.)(Sun et al. 1998; Xi etal. 2002; Xue andZhu 1999)

O(Zhou 1994)(Chen 1983; Wu et al.
1999; Zhu and Chen
1962)
(Chen 1983; Wu et al.
1999)(Chen 1983; Zhu andChen 1962)

P(Chen 1985; Zhu andChen 1962)

P(Zhu and Chen 1962)

P(Chen 1983; Zhu etal. 1964)

P(Zhou 1994)

P(Zhou 1994)

P(Zhou 1994)

China

P

(Zhu and Wang 1996)

China; Japan; Korea; Europe; Africa
China; Japan; Korea; Turkey; Greece;
South Europe
HLJ; XJ; ZJ (China); Korea; Japan;
Russian; Turkey; Spanish; Kazakhstan

East China; Europe; India; JapanP(Liu and Bai 1977)

Northeast China; Europe; Minor Asia;
North America

P(Liu and Bai 1977;
Liu et al. 1983)

East and north China; Europe; Siberia;
Korea; JapanP(Liu and Bai 1977)

Epiblema tetragonana (Stephens)

LeavesNortheast China; Europe

P

Epinotia ustulana Hübner

Leaves,buds

Northeast China; Europe

O(Liu and Bai 1977;
Liu et al. 1983)(Liu and Bai 1977;
Liu et al. 1983)

Order : FamilySpecies
Orthotaenia undulana (Denis &Schiffermüller)

NotodontidaeaFeeds onLeavesPandemis corylana (Fabricius)

Buds

syndemis musculana （Hübner）

LeavesCapua favillaceana （Hübner）

LeavesArgyroploce lacunana (Denis &Schiffermüller)

LeavesAncylis unculana Haworth

LeavesStauropus basalis Moore

LeavesLocationaNortheast China; Europe; Minor Asia
Northeast and Central China; Europe;
Siberia; Japan
Northeast China; Europe; Siberia; North
America; Japan
Northeast China; Europe; Minor Asia
Northeast China; Europe; Siberia; Minor
Asia
Northeast China; Europe; Minor Asia;
Korea
HLJ, HeB, ZJ, SD, JS, JX, TW, HB, YN,
SC, SH (China); Korea; Vietnam; Russia;
Japan

Host Range

References

P(Liu and Bai 1977;
Liu et al. 1983)

P(Liu and Bai 1977)

P(Liu and Bai 1977)

P(Liu and Bai 1977)

P(Liu and Bai 1977)

P(Liu et al. 1983)

P(Cai 1979)

Abbreviated names of Chinese Provinces, autonomous regions and municipalities: AH, Anhui; BJ, Beijing; FJ, Fujian; GD, Guangdong;GS, Gansu; GX, Guangxi; GZ, Guizhou; HaiN, Hainan; HeB, Hebei; HB, Hubei; HeN, Henan; HLJ, Heilongjiang; HN, Hunan; JL, Jilin;
JS, Jiangsu; JX, Jiangxi; LN, Liaoning; NM, Inner Mongolia; SC, Sichuan, SD, Shandong; SH, Shanghai; ShaX, Shaanxi; ShX, Shanxi;
TB, Tibet; TW, Taiwan; XJ, Xinjiang; YN, Yunnan; ZJ, Zhejiang.
b

P=polyphagous species known to feed on Rubus and other families; O=oligophagous species occurring mainly on Rubus;

M=monophagous on one Rubus species.

We also conducted extensive field surveys for potential biological control agents in Guangxi,
Guizhou, Sichuan and Yunnan Provinces. We have developed collaboration with Professor Yang
Chunhua, an ecologist at Sichuan Agricultural University. Professor Yang and her team member
conducted frequent visits (at intervals of 2-4 weeks) to field sites in Sichuan. A total of 60 species
in 30 families were directly collected on Rubus ellipticus (Table 2).

Fig. 1 Jianqing Ding in a field trip to Sichuan Province

Fig. 2 Jialiang Zhang worked with students

Table 2. Phytophagous insects collected on Rubus ellipticus in China during 2006-2007
Order/FamilySpeciesFrequency

Stages Plant partfoundaUnidentified (5 morphotypes)

commonA

Host
Economic
RangebPest

Coleoptera
Buprestidae

LeavesU

Melolonthidae

Serica orientalis Motschulsky

rare

A

LeavesP

Rutelidae

Anomala cypriogastra Ohaus

occasional

A

P

rare

A

Leaves, flowers,
buds
Leavesoccasional

A

Popillia quadrigutat Fabricius

rare

Dicranocephalus wallichi
bowringi Passos
Oxycetonia bealiae (Gory etPassoe)Cerogria chinensis Fairmsire

commonCerogria anisocena (Wied.)occasional
rare

Hispiinae

Taiawania (s.str)guadriramosa(Gresstt)
Dactylispa sp.

Cerambyciidae

Linda (Linda)atricornis Pis♀

Curculionidae

Sympiezomias menlongensis
Chao
Apoderus minimus Roelofs

rare

Anomala aulax wiede
Anomala rirens Linnaeus

Cetoniidae

Lagriidae
Cassidinae

A

P

*

rare

A

Leaves

P

*

rare

A

LeavesP*

A

LeavesU

A

LeavesU

A

LeavesUoccasionalA

LeavesUoccasional

A

P

A

Leaves, stems(borer as larva)
LeavesU

rare

A

LeavesU

Apoderus nigroapicatus Jekel

occasionalA

LeavesP

Drepanoderes leucofasciatus
Voss
Henicolabus hypomelas Faust

rare

A

LeavesP

rare

A

Leaves

U

A

Buds

O

Lixus mandaranus
fukieienensis Voss
Macrocorynus sp.

rare

A

LeavesO

rare

A

LeavesU

Phymatapoderus latipennis
Jekel
Phytoscaphus dentirostris Voss

rare

A

LeavesP

rare

A

Leaves, stems

PoccasionalA

LeavesU

Chaetoenema kingpinesis ChenoccasionalA

LeavesO

Chaetoenema sp.

occasionalA

LeavesU

Nisotra gemella (Erichson)

rare

A

LeavesU

Hemipyxis tonkinensis (Chen)

rare

A

LeavesPoccasionalA

LeavesP

Basilepta leechi (Jacoby)
Basilepta sp.

rare

A

LeavesUoccasionalA

LeavesP

Oides tarsatus (Baly)

rare

A

LeavesP

Paleosepharia liquidambara
Gressitt et Kimoto
Exosoma flavoventris(Motschulsk
Paridea sinensis Laboissiere

rare

A

LeavesP

rare

A

LeavesU

rare

A

LeavesP

rare

A

LeavesU

rare

A

LeavesU

rare

A

LeavesU

Cleoporus varriabilis (Baly)
GalerucinaePoccasional

Ptochus sp.

Eumolpinae

P

Leaves, flowers
buds
LeavesInvolvulus sp.

Alticinae*

Morphosphaera viridipennis
Laboissiere
Stenoluperus parvus Gressitt etKimotoLinaeidea placida (Chen)*Order/FamilySpeciesFrequency

Stages Plant partfoundaoccasional A, L, P Leaves

Host
Economic
RangebPest
O/M

Chlamisinae

Chlamisus sp.(2 morphoypes )Cryptocephalinae

Cryptocephalus sp.

rare

A

LeavesU

Platypleura kaempferi(Fabricius)
Unidentified

rare

A

P

rare

A

Leaves, roots,
stems
Leaves, Stems

U

Clovia quadrangularis Metcakf﹠HortonTricentrus aleuritis Chou,

rare

A

Leaves, Stems

U

occasional

A, N

Leaves, Stems

O

ALeaves, Stems

P

Homoptera
Cicadidae
Ledridae
Aphrophoridae
Membracidae
Cicadellidae

Bothrogonia (O.)acuminateyang ＆ Li
Unidentified

commonrare

A

Leaves, Stems

U

Cercopidae

Paphnutius ruficeps (Melichar)occasionalA

Leaves. StemsU

Cosmoscarta exultans( Walker)
Paphnutius semirufus (Haupt)

rare

A

Leaves, stems

U

rare

A

Leaves, stems

U

Acraea issoria（Hübner）

rare

A

LeavesO*

Lepidoptera
Acraeidae*

Thyatiridae

Gaurena delattini Werny

commonA, L

LeavesU

Noctuidae

Anomis mesogona (Walker)

common

A, L, P

LeavesP*

Lymantriidae

Euproctis flava (Bremer)

rare

A, L

LeavesP*

Lycaenidae

rare

A, L, P

LeavesO

Choreutidae

Sinthusa chandrana grotei(Moowe)Choreufis sp.

rare

A, L, P

Tortricoidae

Epinotia sp.

Pyralidae

Unidentified

rare

L

Unidentified

common

A, L, P

Xenocatantops brachycerus(Will.)
Tegra novaehollandiae
viridinotata (Stal)

commoncommonLeavesU

Leaves, buds

U

stems

U

Leaves, buds

U

A, N

LeavesP*

A, N

LeavesP*occasional A, L, P

Hymenoptera
Argidae
Orthoptera
Catantopidae
TettigoniidaeaL, larva; P, pupa; A, adult; N, nymph.

bP, polyphagous species from other families; O, oligophagous species occurring mainly on Polygonaceae; M, monophagous on Rubus

ellipticus, U, unknown.

Important insects
Chlamisus setosus (Bowditch) and Chlamisus sp. (Coleoptera: Chlamisinae)
The warty leafbeetle, Chlamisus setosus (Bowditch) (Fig. 3), was recorded on R. ellipticus var.
obcordatus in Yunnan Province and some species from this genus Chlamisus were also recorded
as monophagous or oligophagous (Reu Jr. and Del-Claro 2005; Tan et al. 1980). In our survey,
larvae and adults of Chlamisus sp. were found on the R. ellipticus var. obcordatus from Kunming,
Dali, and Simao, Baoshan city in Yunnan province. However, from our preliminary host range testin Wuhan Botanical Garden, it could be oligophagous by feeding on some plants in Rubus otherthan its original host (unpublished data). It has the whole life cycle on R. ellipticus var. obcordatus,with one generation per year. We found it mates and lays eggs in field during summer andhibernate during winter in our lab, but no emergence in Wuhan Botanical Garden. It makes small
feeding holes on the back of leaves. Although the damage by this leafbeetle on R. ellipticus var.

obcordatus is not serious, its feeding scars could be helpful for some fungal infection. This beetle
could be considered as a biological control agents for further study.

Fig. 3 Chlamisus larva

Fig. 4 Chaetoenema adult

Flea beetles (Coleoptera: Alticinae)
Two species in genus Chaetoenema (Coleoptera: Alticinae) (Fig. 4) were collected in the same site,
Qinghai Lake in Tengchong county, where 5-6 species in Rosaceae naturally distributed. These 2
flea beetles have very similar size and color, and collected on R. ellipticus and another Rubus sp.
(close to Rubus alceifolius). We could not distinguish them in field at first, so we collected them
together, and then conducted preliminary host range test in laboratory. The feeding holes on R.
ellipticus observed in lab were more intensive than on the Rubus sp. in our study site andlaboratory test got the host range result as oligophagous. But more study should be taken to knowwhether they have respective or common host range.
Weevils (Coleoptera: Curculionidae)
Weevils have been considered as potential successful biological control agents of many weeds formany years for their narrow host range. In our study, four species in Family Curculionidae wereconsidered to be potential candidates. They are: Involvulus sp., Ptochus sp., Sympiezomias
menlongensis Chao and Apoderus minimus Roelofs. Involvelus sp. is shiny bronzed colorful,feeding on the buds of R.ellipticus. We also collected it on Rubus niveus Thunb. The weevil inserts
its mouthpart to the base of buds, and then the buds wither. It can be found in wide range but
mostly in hills near Yunnan Agricultural University. Ptochus sp. and Sympiezomias menlongensis
Chao are grey. They were very common in several sites, but it just has limited number in each site.
Apoderus minimus Roelofs, a dark orange leaf-rolling weevil, was collected in a wide range inYunnan province. It makes small holes in raspberry leafs. Further testing is needed to determine
the host range of these potential candidates.
Epinotia sp. (Lepidoptera: Tortricoidae)
The larvae of the leaf-rolling moth, Epinotia sp. (Lepidoptera: Tortricoidae) (Fig. 5-6), werecollected in several sites from Yunnan province and reared to adults in laboratory in Wuhan
Botanical Garden (WBG). The rolling impacts of the larvae on the leaves were very heavy inmany exploring sites, where they can be found in nearly all the top of the raspberry twigs, theseactions of the larvae can limit the growth of branches. After being reared in our laboratory, somelarvae emerged to adults while some are parasitized by unidentified hymenopterans during larval
and pupa stage. No moth mating and oviposition was found in laboratory. Other species in genus
Epinotia were recorded as forest or agriculture pests in some literatures and some are report asmonophagous or oligophagous (Liu and Bai 1977; Muenster-Swendsen 1991; Su et al. 2003).
More field exploration and laboratory tests are needed to determine if the leaf-rolling moth could
be a biological control candidate.

Fig. 5 Epinotia Larvae

Fig. 6 A Epinotia larva in buds

Stem borer (Lepidoptera: Pyralidae)
Larvae of unidentified stem borer (Fig. 7-8) were found in both Sichuan and Yunnan Provinces.
The larvae were white with light brown head. The damage of the borer was heavy. Fresh or brownwaste of larvae stuffed the hole of the stem. We collected some stems with borer, but failed to rear
adults in the laboratory. Larvae in Pyralidae have been noted as good candidates for weed
biocontrol (Goolsby et al. 2003), thus this insect may warrant further study.

Fig. 7 A stem borer

Fig. 8 Feeding damage in stem

Sawfly (Hymenoptera: Argidae)
Larvae of the sawfly (Fig. 9) were found to feed on R. ellipticus in both Sichuan and Yunnan
province. They were reared to pupae and adults in WBG. Pupae were covered in brown cocoons.
Adults are shiny metallic black/blue. Most of the species recorded in this family may have narrow
host range (Badenes-Perez and Johnson 2007; Bruzzese 1982; Goulet 2007; Sahragard andHeydari 2001), so further study on identification and host range should be conducted.

Metallic wood-boring beetles (Coleoptera: Buprestidae)
A large number of beetles with 5 morphotypes were collected in the Yunnan and Sichuan province.
They fed on the leaves, and their impacts are significant in some sites (Fig. 10). Some beetles inthis family may have narrow host range and have been considered as biological control agents(Bruzzese 1980; Kaya and Kovanc 2004 ; Lei and Zhou 1998). More study on identification andhost range of these beetles should be considered.

Fig. 9 Sawfly

Fig. 10 Buprestid adult in stem

Important fungal pathogen
Sixty five fungal species in 19 families of 3 phylum are associated with Rubus, (Table 3), some ofthem may be potential candidates for biological control Rubus ellipticus in Hawaii. There are 32
species from Ascomycota, 32 from Basidiomycota and 1 from Oomycota. Among those fungus, 19
species were directly reported from R. ellipticus. All fungi data of in this paper are from publishedliterature or online database.

Table 3. Literature review: species of pathogenic microorganisms on the genus Rubus in East Asia
DamageDistribution a

Coryneopsis rubi

Twigs,

Europe; North America (USA); Asia

(Westend.) Grove

branches,(China, India); Africa (Ethiopia, South

leaves, Shoots.

Africa); Australia; New Zealand

LeavesGX(China );Japan; Burma

Phylum/ Family Species

HostRangeb

References

Ascomycota
Amphisphaeriaceae

Clavicipitaceae

Aschersonia tamurai

O

2007)

P

Henn.
Cercoseptoria

Cercoseptoria(Bai 2003; Cybernome(Cybernome 2007; Dai
1979; Index 2007)

Uncertain

India

U

(Bioscience 2007;
Cybernome 2007)

heteromalla (Syd.)
Kamal & Narayan
Dothideaceae

Septoria darrowii

LeavesTW (China)

ZellerO(Dai 1979; Index 2007)(R. triathus R.
hispidus)Elsinoaceae

Elsinoë veneta(Burkh.) Jenkins

Twigs, leaves

JL (China); Greece; Chile; the UnitedStates; the British Isles; Ireland; Australia

O(Bioscience 2007;
Cybernome 2007; Dai
1979)Phylum/ Family Species

DamageDistribution a

HostRangeb

References

Hyaloscyphaceae

Stems

Jamaica; Spain

P (R. ellipticus)(Cybernome 2007; Farr etVerticillium

Roots, stems,

China; Ukraine; Uzbekistan; Armenia;

albo-atrum Reinke &twigs, leaves,

Russia; Canada; Cuba; Turkmenistan;

Bertholdbuds

Venezuela; Brazil; Japan; Chile; Georgia;

Polydesmia fructicola
Korf

Hypocreaceaeal. 2007; Zhuang 1987)
P(Cybernome 2007; Dai
1979; Index 2007)

France; Puerto Rico; Australia; the United
Kingdom; Spain; Argentina; the former
USSR
Hyaloscyphaceae

Calycellina

Stems

India

M (R .ellipticus) (Lowen and Dumont 1984;
Thind and Sharma 1980)

tetraspora K.S.
Thind & M.P.
SharmaLeotiaceae

Strossmayeria

Stems

British Isles;

Jamaica;

Slovakia

P(R. ellipticus)(Cybernome 2007; Zhuang
1987)

basitricha (Sacc.)
DennisStems, culms

Jamaica

P(R. ellipticus)

Stems, twigs

Jamaica, Venezuela

P (R. ellipticus ) (Zhuang 1987)

Velutarina

Stems,

Argentina;

rufo-olivacea (Alb. &twigs ,branches

United Kingdom;

Appendiculella

Leaves andChina; Uganda; South Africa; India;

calostroma (Desm.)

Stems

Ecuador; Colombia; Costa Rica; Sao

Strossmayeria(Iturriaga and Korf 1990)

jamaicensis (Seaver)
Iturr. & Korf
Unguiculariopsis
ravenelii (Berk. &M.A. Curtis) W.Y.
Zhuang & Korf apud
W.-y. Zhuang
British Isles;

Jamaica;Ukraine;P (R. ellipticus)

USA(Cybernome 2007)(Zhuang 1987)

Schwein.) Korf
Meliolaceae

P (R. ellipticus)(Cybernome 2007; Dai
1979; Hu 1996)

Tome and Principe Democratic Republic;

Höhn.

Venezuela; Brazil; Japan; Chile; Puerto
Rico; Australia; Guadeloupe; the UnitedStates; the United Kingdom; thePhilippines; Vietnam; Ireland.
Meliola rubi F.

LeavesPhilippine

M (R. moluccanus) (Cybernome 2007)

LeavesPhilippine; India

O (R. ellipticus, R. (Cybernome 2007; Farr et

Stevens & Roldan ex
Hansf.
Meliola rubiella
Hansf.
Meliola formosensis

LeavesTW(China)

moluccanus)al. 2007; Patil et al. 2004)

O(Cybernome 2007; Dai

W. Yamam.
Mycosphaerellaceae

Cercospora rubicola
Thüm.

1979; Hu 1999)
LeavesYN, GX (China); Georgia

O (R.
cochinchinensis R.
caucasigenus)(Liu and Guo 1998)

Phylum/ Family Species
Mycosphaerella

DamageDistribution a

HostRangeb

References

LeavesHB, SC, GZ, (China); Russia; India;

P (R. ellipticus)(Index 2007; Liu and Guo

confusa F.A. Wolf

1998)

Latvia; Chile; Georgia; Puerto Rico;
United Kingdom

Mycosphaerella

Leaves, stems,

JS, JL, FJ, LN, HLJ, SD, ZJ, XJ, HeN,

fragariae (Tul.)fruit stalks,

GD, SC (China); Ukraine; Cuba;

Lindauberry caps

Turkmenistan; the Dominican Republic;

P(Cybernome 2007; Dai
1979)

Venezuela; Chile; Georgia; Puerto Rico;
Jamaica; the British Isles; Ireland; Japan
Mycosphaerella rubi

Leaves, shoots

Roark
Pseudocercosporella

China; Ukraine; Venezuela; Georgia;

O

Puerto Rico; Azerbaijan.
Leaves

Nepal(Cybernome 2007; Dai
1979)

M (R. ellipticus) (Index 2007; Singh et al.
1997)

arcuata S.K. Singh,
P.N. Singh & Bhalla
Pseudocercospora

Patellariaceae

LeavesChina; Zaïre; Sudan; India; Ethiopia;

heteromalla (Syd.)

Uttar Pradesh; Bhutan; Uttaranchal; New

Deighton

Zealand; United Kingdom; Autrilia

RhytidhysteronStems, shoots

China; India; Cuba; Colombia; theO (R. ellipticus, (Cybernome 2007; Index
R. idaeus,)

2007; Liu and Guo 1998)

P(Cybernome 2007; Dai
1979)

Dominican Republic; Venezuela; Brazil;

rufulum (Spreng.)

Chile; Haiti; Jamaica; Trinidad; Tobago;
Virgin Islands (United States)
Phaeosphaeriaceae

Stagonospora

LeavesJL (China)

brevispora (Ellis &O (R. ideaus, R. (Dai 1979; Index 2007)
occidentalis)

J.J. Davis) Castellani& E.G. Germano
Hendersonia vulgaris

Twigs

HB(China); Ukraine

P

Desm.
Sclerotiniaceae(Cybernome 2007; Dai
1979)

Botryotinia

Stems, fruits,

China; Ukraine; Uzbekistan; Israel; Iran;fuckeliana (de Bary)

flowers, twigs,

Russia; Congo; South Africa; India; Cuba;

Whetzelshoots, leaves

Kazakhstan; Colombia; Tanzania;

P(Cybernome 2007; Dai
1979)

Ethiopia; Cyprus; Dominican Republic;
Venezuela; Brazil; Italy; Moldova; New
Zealand; Japan; Chile; Georgia; Poland;
Zimbabwe; Australia; Estonia; USA;
Kenya; United Kingdom; Netherlands;
Portugal; Azerbaijan; Argentina
Lambertella

Stems, petiole

Jamaica; India

aurantiaca V.P.

P

(Cybernome 2007; Index(R. ellipticus)

2007; Zhuang 1987)

P (R .ellipticus)(Cybernome 2007; Index

Tewari & D.C. Pant
Lambertella

Uncertain

Jamaica;corni-maris Höhn.
Incertae sedis

Acrothecium rubi
Sawada2007; Zhuang 1987)
LeavesTW (China)

O (R.
conduplicatus, R.
trianthus)(Dai 1979; Index 2007)

Phylum/ Family Species
Chalara verruculosa

DamageDistribution a

HostRangeb

Stems

MalawiM (R. ellipticus) (Bioscience 2007; Index

Stems

MalawiM (R. ellipticus) (Bioscience 2007;

B. Sutton
Sporendocladia

References

2007; Sutton 1993)

Cybernome 2007)

kionochaetoides B.
SuttonVenturiaceae

Coleroa chaetomium

Leaves(Kunze) Rabenh

Ukraine; Germany; the Czech Republic;

O (R. ellipticus, R. (Bioscience 2007;

Georgia; United Kingdom; India

caesius, R. idaeus) Cybernome 2007)

Basidiomycota
Incertae sedis

Caeoma cheoanum

LeavesGX, GZ (China)

LeavesJapan; Australia

M (Rubus sp.)

(Dai 1979; Index 2007)

Cummins
Phragmotelium

M (R. parvifolius) (Berlese et al. 1888;barnardii (Plowr. &Cybernome 2007; Sydow

G. Winter) Syd.

1897)

Phragmotelium

LeavesTW (China)

O(Cybernome 2007; Dai
1979)

formosanum (Hirats.)
Thirum.
Phragmotelium

LeavesHLJ (China )O(Cybernome 2007; Dai
1979)okianum (Hara)
Thirum.
Phragmotelium

LeavesTW (China); India

O(Dai 1979; Index 2007)

LeavesTW (China); India

P (R. ellpiticus)(Farr et al. 2007; Onorubi-fraxinifolii (Syd.
& P. Syd.) Thirum
Uredo spinulosa Y.
Ono
Phragmidiaceae

Arthuriomyces

1982; Patil et al. 2004)
LeavesHLJ (China) ,Russia, United states, NorthO(Dai 1979; Index 2007)

Americapeckianus (Howe)
Cummins & Y.
Hirats.
Arthuriomyces

LeavesSC (China )LeavesSC, GX, GZ, GD, TW, FJ, HN (China)

P (R. pileatus)(Index 2007)

rubicola J.Y. Zhuang
Gerwasia rubi Racib.

O(Cybernome 2007; Dai
1979)

Hamaspora

LeavesTW, GZ, ShX, JX, GD, GX, HN(China);acutissima P. Syd. &Papua New Guinea; Philippines;

Syd

Indonesia; Australia ; Papua New Guinea;

O(Dai 1979; Farr et al. 2007)

Japan; New Guinea; South Africa
Hamaspora

LeavesYN, TW, GZ (China), Philippine

benguetensis Syd.

O (R. ellipticus R. (Cybernome 2007; Dailambertianus, R. 1979)
taiwanianus)

Hamaspora

LeavesTW, ZJ, GX, SC (China)

O(Dai 1979)

LeavesNew Guinea

O(Cybernome 2007; Dai

hashiokai Hirats. f.
Hamasporarubi-sieboldii

1979; Farr et al. 2007)

Phylum/ Family Species

DamageDistribution a

LeavesGZ, HN, SC, GX (China)

HostRangeb

References(Kawagoe) Dietel
Hamaspora sinica

O(Cybernome 2007; Dai
1979)

F.L. Tai & C.C.
Cheo
Hamaspora tairai

LeavesHB(China)

M (Rubus sp.)

(Dai 1979)

LeavesTW (China)

M (R kawakamii) (Cybernome 2007; Dai

Hirats.
Hamaspora

1979)

taiwaniana Hirats. f.
& Hashioka
Phragmidium

Leaves

ShaX, TW, (China)

O(Cybernome 2007; Dai
1979)arisanense Hirats. &HashiokaPhragmidium

LeavesTW, HeB, LN, JL, AH (China)

Ogriseum Dietel
Phragmidium

1979)
LeavesNorth AmericaO

occidentale Arthur
Phragmidium(Cybernome 2007; Dai(Cybernome 2007; Index
2007)

LeavesChina, India

M (R. ellipticus) (Cybernome 2007; Index
2007)

orientale Syd. & P.
Syd
Phragmidium

LeavesZJ, AH, GZ (China); Japan

LeavesSC (China)

LeavesShaX (China); Ukraine; Russia; the Czech

O(Dai 1979; Index 2007)

rubi-thunbergii
KusanoPhragmidium

M (R. eucalyptus) (Index 2007; Wei 1988)

rubi-eucalypti S.X.
Wei
Phragmidium
rubi-idaei (DC.) P.

Republic; Georgia; Switzerland; United

Karst.

States; United Kingdom

Phragmidium

LeavesGeorgiaO

2007)

M (R. saxatilis)

rubi-saxatilis LiroPhragmidium(Cybernome 2007; Index(Cybernome 2007; Index
2007)

LeavesChina

M (R. parvifolius) (Cybernome 2007; Index
2007)

rubi-parvifolii Liou
& WangPhragmidium

LeavesJapan

M (R. oldhami)(Cybernome 2007; Index
2007)

rubi-oldhami
Togashi & Y. Maki
Phragmidium rubi

LeavesUkraine, Russia, Georgia

P(Pers.) G. WinterPhragmidium(Cybernome 2007; Index
2007)

LeavesShaX (China)

O(Cybernome 2007; Dai
1979)shensianum F.L. Tai& C.C. Cheo
Phragmidium
sikangense Petr.

Leaves

ShaX, SC (China);

O(Cybernome 2007; Dai
1979)Phylum/ Family Species
Phragmidium

DamageDistribution a

LeavesHeB (China)sinicum F.L. Tai &HostRangeb
M (R.
crataegifolius)

References(Cybernome 2007; Dai
1979)

C.C. Cheo
Phragmidium

LeavesYN (China); Ukraine; France; Greece;

violaceum (Schultz)

Chile; Georgia; the United Kingdom;

G. Winter

Azerbaijan

O(Cybernome 2007; Dai
1979)

Leaves

ShaX, SC (China); JapanO(Dai 1979)

Phytophthora

Leaves, stems,

China; Sweden; British Columbia; Italy;P(Index 2007; Yun 1998)

citricola Sawada

twigs

Zimbabwe; Switzerland; Portugal;

Phragmidium
yamadanum Hirats.
Oomycota
Pythiaceae

Northern America; Yugoslavia; India;
South Africa; New Zealand; Sicilia; NewSouth Wales; Ireland; United Kingdom;France; Great Britain; Southern SouthAmerica; South Australia Western
Australia; United StatesaChinese Provinces abbreviated as in Table 1.

bP, polyphagous species from other families and R ellipticus listed in parentheses for the known ones; O, oligophagous species occurring

mainly on Rubus and plant names listed in parentheses for the known ones; monophagous on one weed and plant name listed inparentheses for the known one, U, unknown. All information based on literature records and online database.
.

Potential candidates in phylum Ascomycota
Six species in Ascomycota were listed in this literature. Two species are leaves spot
Pseudocercosporella arcuata S. K. Singh, P. N. Singh & Bhalla and Pseudocercospora
heteromalla (Syd.) Deighton (Ascomycota: Mycosphaerellaceae), which were reported as
monophagous and oligophagous respectively, P. heteromalla also reported on Rubus idaeus(Cybernome 2007; Index 2007; Singh et al. 1997). Chalara verruculosa B. Sutton andSporendocladia kionochaetoides B. Sutton (Ascomycota: Incertae sedis) were found on R.
ellipticus from Malawi in 1993(Bioscience 2007; Index 2007; Sutton 1993). Calycellina
tetraspora K.S. Thind & M.P. Sharma (Ascomycota: Hyaloscyphaceae) was found on stems ofour target weed in India (Lowen and Dumont 1984; Thind and Sharma 1980). Coleroa
chaetomium (Kunze) Rabenh could be oligophagous on R. ellipticus, R. caesius; R. idaeus, which
epiphyllous, superficial, solitary or clustered pseudothecium parasitizes on live leaves (Ellis and
J.P. 1997).
Potential candidates in phylum Basidiomycota
Two rust species, Phragmidium orientale Syd. & P. Syd and Hamaspora benguetensis Syd.
(Basidiomycota: Phragmidiaceae) were recorded as monophagous and oligophagous respectively
on R. ellipticus, the later also was found on R. lambertianus, R. taiwanianus and some Rubus sp.

Preliminary host range test of the warty leaf beetle and flea beetle
The warty leaf beetle (Chlamisus spp.) was recorded as monophagous in some literature. We tested

its host specificity in no-choice test in lab, using about 20 plant species, of which, 13 species arein the genus of Rubus. Our tests indicated both the warty leaf beetle and flea beetle may beoligophagous (Table 4).
Table 4. Preliminary host range tests of Chlamisus spp. and Chaetocnema spp.
Chlamisus

Chlamisus

Pop. 1

Pop. 2

Rubus ellipticus Smith√√

Rubus niveus Thunb.

√√

Rubus pinfaensis Lévl.et Vant.

√Light feeding

√

√

√

Light feeding

Rubus irenaeus Focke

Light feeding√

Light feeding

×

Light feeding×

Rubus sumatranus Miq.

×

√

×

×

×

Rubus tephrodes Hance√√

Light feeding

×

×

×

Rubus amabilis Focke

×

×

×

×

×

×

Rubus reflexus Ker

×

×

×

×

×

×

Rubus lambertianus Ser.

√×

Light feeding

×

×

×

Rubus sp.

√

Light feeding Light feeding

××

Plant species

Rubus coreanus

Chlamisus Chaetocnema Chaetocnema Chaetocnema
Pop. 3Pop. 1

Pop. 2√√√light feeding light feeding light feeding

Pop. 3√×

Light feeding

×

Light feeding√

Light feeding

Rubus trianthus Focke

××

Light feeding××

Rubus sp.

×

×

×

×

×

Vaccinium virgatum Ait.

×

×

×

×

×

×

Potentilla fulgens Wall

×

×

×

×

×

×

Sanguisorba officinalis L.

×

×

×

×

×

×

Diospyros kaki Thunb.

×

×

×

×

×

×

Cerasus cerassoides

×

×

×

×

×

×

Eupatorium adenophorum

×

×

×

×

×

×

Rosa chinensis Jacq.

×

×

×

×

×

×

Rosa omeiensis Rolfe

×

×

×

×

×

×

Miq.var.tomentosus Card.

Die out

Discussion and future directions
This literature and field survey has revealed numerous herbivorous arthropods and pathogenic
microorganisms associated with Rubus ellipticus in its native range. Some of them appear
specialized and have been reported or observed to have an impact of the target species.
The most promising insect candidates, in our study, for control of the invasive R. ellipticus appear
to be the warty beetles, the flea beetles, the stem borer, the leaf-rolling moth, and the sawfly.
However, more controlled screening of their host specificity is a necessary next step. Considering
the fragile of ecology in Hawaii, all candidate species should be subjected to further risk
assessment. More importantly, investigations of the impact of these insect species upon R.
ellipticus, and of their host specificity, are needed in order to evaluate their potential as bio-control
agents.
Literature review (Tables 1 and 3) provides more clues to screen potential natural enemies. If

herbivores have a wider distribution than the range of the target plant, it may suggest thecandidates as not host-specific. Comparing Table 1 with Table 2, few common herbivores could befound in both tables, which indicates the herbiviory diversity on Rubus taxa in the original region
and the current limited knowledge of the herbiviory on Rubus ellipticus. Buprestidae has 5
unidentified morphotype in our survey but only one species was recorded in literature (Lei andZhou 1998), the distribution of the species matched that of Rubus ellipticus well in China, but it
was recorded in Japan. Thus, further study is needed to show if they are the same species. The
distribution of warty leafbeetles, Chlamisus setosus (Bowditch) (Coleoptera: Chlamisinae) foundin Yunnan province matches well with the literature record (Tan et al. 1980). The leaf-rolling moth,Epinotia sp. (Lepidoptera: Tortricoidae) could be found from a large range in Yunnan Province,
which does not match to the distribution of the moth Epinotia ustulana Hübner, which is a pest on
fruit trees (Liu and Bai 1977; Muenster-Swendsen 1991; Su et al. 2003).
Historically, fungi have been used as biological controls for Rubus and some of them have beeneffective (Gardner and Davis 1982; Goeden et al. 1974). However, out of 7 insects and 3 fungi rustspecies imported to New Zealand in 1920s none showed significant impact on wild blackberry,and only one rust, Phragmidium violaceum, with narrow host range was considered for further
study (Pennycook 1997). Rusts in genus Phragmidium are also potential choice in control of R.
ellipticus, 12 species list in this paper are from China, one report from R. ellipticus, the other fungi
species listed also could be considered.
References
Aslan MM and Uygun N (2005) Aphids (Homoptera: Aphididae) of Kahramanmaraß Province, Turkey. Turk J Zool 29:
201-209
Badenes-Perez FR and Johnson MT (2007) Ecology, host specificity and impact of Atomacera petroa Smith(Hymenoptera: Argidae) on Miconia calvescens DC (Melastomataceae). Biological Control 43: 95-101
Bai J (2003) Flora Fungorum Sinicorum. Science Press, Beijing, China, 372 pp
Berlese AN, De-Toni JB and Fischer E (1888) Sylloge Fungorum, 746 pp
Bioscience C (2007) HERB. IMI ON-LINE, CABI Databases.
Bruun H (2006) Prospects for Biocontrol of Invasive Rosa rugosa. BioControl 51: 141-181
Bruzzese E (1980) The phytophagous insect fauna of Rubus spp. (Rosaceae) in Victoria, a study on the biologicalcontrol of blackberry (Rubus fruticosus L. Agg.). pp 1-6
Bruzzese E (1982) The host specificity ofHartigia albomaculatus (Hym.: Cephidae] and its potential effectiveness in the
biological control of European blackberry. BioControl 27: 335-342
Cai R (1979) Economic Insect Fauna of China. Science Press, Beijing, China, 166 pp
Chen J (1985) The List of the Noctuidae in Guizhou Province. pp 43-60
Chen Y (1983) Noctuidae. In: Wang P, Wang L, Fang C, Bai J, Zhu H, Liu Y and Liu X (eds) Iconographia
Heterocerorum Sinincorum, pp 237-390, Science Press, Beijing, China
Cybernome (2007) Fungi.
Dai F (1979) Sylloge Fungorum Sinicorum. Science Press, Beijing, China, 1527 pp
Database E (2007) Essig Museum of Entomology Collections. University of California, Berkeley
Ding J, Reardon R, Wu Y, Zheng H and Fu W (2006a) Biological control of invasive plants through collaboration
between China and the United States of America: a perspective. Biological Invasions 8: 1439-1450
Ding J, Wu Y, Zheng H, Fu W, Reardon R and Liu M (2006b) Assessing potential biological control of the invasive plant,tree-of-heaven, Ailanthus altissima. pp 547 - 566, Taylor &amp; Francis
Ellis MB and J.P. (1997) Microfungi on Land Plants: An Identification Handbook
Evans KJ and Bruzzese E (2003) Life history of Phragmidium violaceum in relation to its effectiveness as a biologicalcontrol agent of European blackberry. Australasian Plant Pathology 32: 231-239
Farr DF, Rossman AY, Palm ME and McCray EB (2007) Fungal Databases. Systematic Mycology and Microbiology
Laboratory, ARS, USDA
Gardner DE (1992) Plant pathogens as biocontrol agents in native Hawaiian ecosystems. In: Stone CP, Smith CW and
Tunison JT (eds) Alien Plant Invasions in Native Ecosystems of Hawaii: Management and Research, pp 432-451
Gardner DE and Davis CJ (1982) The prospects for biological control of non-native plants in Hawaiian National Parks
In: Smith CW (ed) Tachnical report 45, p 55, Cooperative national park resources studies unit university of Hawaii
at Manoa,Department of Botany Honolulu, Hawaii 96822, University of Hawaii at Manoa Cooperative Agreement
No. CX 8000 20001
Gardner DE, Hodges CS, Killgore E and Anderson RC (1997) An evaluation of the rust fungus Gymnoconia nitens as apotential biological control agent for alien Rubus species in Hawaii. Biological Control 10: 151-158
Gerrish G, Stemmermann L and Gardner DE (1992) The distribution of Rubus species in the state of Hawaii. Technical
Report 85, p 14, Cooperative National Park Resources Studies Unit, University of Hawaii at Manoa; Western
Region, National Park Service, Honolulu, Hawaii 96822
Goeden RD, Andres LA, Freeman TE, Harris P, Pienkowski RL and Walker CR (1974) Present Status of Projects on the
Biological Control of Weeds with Insects and Plant Pathogens in the United States and Canada. Weed Science 22:490-495
Goolsby JA, Wright AD and Pemberton RW (2003) Exploratory surveys in Australia and Asia for natural enemies of Old
World climbing fern, Lygodium microphyllum: Lygodiaceae. Biological Control 28: 33-46
Goulet H (2007) Sawflies (Hymenoptera: Symphyta).

Assessment of species diversity in the mixedwood plains

ecozoneHasan S (1980) Plant pathogens and biological control of weeds. Rev. Plant Pathol. 59: 349-356
HIRANO Y (1998) A LIST OF COLEOPTERA FROM KANAGAWA PREFECTURE, CENTRAL JAPAN.
Howarth DG, Gardner DE and Morden CW (1997) Phylogeny of Rubus Subgenus Idaeobatus (Rosaceae) and its
Implications Toward Colonization of the Hawaiian Islands. Systematic Botany 22: 433-441
Hu Y (1996) Flora Fungorum Sinicorum. Science Press, Beijing, China, 270 pp
Hu Y (1999) Flora Fungorum Sinicorum. Science Press, Beijing, China, 252 pp
Index Fp (2007) Index Fungorum. CABI Bioscience Databases
Iturriaga T and Korf RP (1990) A monograph of the discomycete genus Strossmayeria (Leotiaceae), with comments on
its anamorph, Pseudospiropes (Dematiaceae). Mycotaxon 36: 383-454
Jacobi JD and Warshauer FR (1992) Distribution of six alien plant species in upland habitats on the island of Hawai'i. In:
Stone CP, Smith CW and Tunison JT (eds) Alien Plant Invasions in Native Ecosystems of Hawai'i, pp 155-188,
Univ. Hawaii Coop. Natl. Park Resour. Stud. Unit. Honolulu: Univ. Hawaii Pr
Kaya M and Kovanc B (2004 ) The species of Coleoptera occurred in raspberry areas in Bursa. Ondokuz Mays
Üniversitesi, Ziraat Fakültesi Dergisi 19: 1-7
Lei C and Zhou Z (1998) Insect Records of Hubei, China. Hubei Science and Technology Publishing House, Wuhan,
China., 650 pp
Liu X and Guo Y (1998) Flora Fungorum Sinicorum. Science Press, Beijing, China, 473 pp
Liu Y and Bai J (1977) Economic Insect Fauna of China. Science Press, Beijing, China, 93 pp
Liu Y, Xiuqiong L and Bai J (1983) Tortricidae. In: Wang P, Wang L, Fang C, Bai J, Zhu H, Liu Y and Liu X (eds)
Iconographia Heterocerorum Sinincorum, pp 28-56, Science Press, Beijing, China
Lowe S, Browne M, Boudjelas S and Poorter MD (2005) 100 of the World's Worst Invasive Alien Species: A Selection

from the Global Invasive Species Database. Invasive Species Specialist Group.
Lowen R and Dumont KP (1984) Taxonomy and Nomenclature in the Genus Calycellina (Hyaloscyphaceae). Mycologia
76: 1003-1023
Lu L and Boufford DE (2003) Rubus L. In: Wu Z and Peter HR (eds) Flora of China(Pittosporaceae throughConnaraceae), pp 196-288, Science Press and Missouri Botanical Garden Press, Beijing and St. Louis
Markin GP, Lai PY and Funasaki GY (1992) Status of biological control of weeds in hawaii and implications for
managing native ecosystems. In: Stone CP, Smith CW and Tunison JT (eds) Alien Plant Invasions in Native
Ecosystems of Hawaii: Management and Research pp 466-482, University of Hawaii Press, Honolulu, Hawaii
96822
Muenster-Swendsen M (1991) The effect of sublethal neogregarine infections in the spruce needleminer, Epinotia
tedella (Lepidoptera: Tortricidae). Ecological Entomology 16: 211-219
Oehrens E (1977) Biological control of the blackberry through the introduction of rust, Phragmidium violaceum, in
Chile. FAO Plant Prot. Bull 25: 26-28
Ono Y (1982) Transactions of the British Mycological Society 79: 426
Patil AR, Patil TM and Patil MS (2004) Studies on rust fungi - VI. J. Mycol. Pl. Pathol. 34: 826-832
Pennycook SR (1997) Blackberry in New Zealand. In: Groves RH, Williams J and Corey S (eds) Towards an integrated
management system for blackberry (Rubus fruticosus L. agg.), pp 151-199, Plant Protection Quarterly, Charles
Sturt University, Albury, New South Wales
Reu Jr. WF and Del-Claro K (2005) Natural History and Biology of Chlamisus minax Lacordaire (Chrysomelidae:
Chlamisinae). Neotropical Entomology 34: 357-362
Sahragard A and Heydari R (2001) Biology and spatial distribution of rose sawfly, Arge rosae (Hym.: Argidae) in Guilan.
Journal of Entomological Society of Iran 21: 25-37
Santos GL, Cuddihy LW and Stone CP (1991a) Control of Yellow Himalayan Raspberry (Rubus ellipticus Sm.) with cut
stump herbicide treatments in Hawaii Volcanoes National Park. In: Smith CW (ed) Technical Report 80, p 25,
Cooperative National Park Resources Studies Unit, University of Hawaii at Manoa; Western Region, National Park
Service, Honolulu, Hawaii 96822,
Santos GL, Kageler D, Gardner DE, Cuddihy LW and Stone CP (1992) Physical and chemical control herbicidal control
of selected alien plant species in Hawaii volcanoes national park. In: Stone CP, Smith CW and Tunison JT (eds)
Alien plant invasions in native ecosystems of Hawaii: management and research, pp 341-376
Santos GL, Stone CP and Cuddihy LW (1991b) Control of Yellow Himalayan Raspberry (Rubus Ellipticus Sm.) with
Cut Stump Herbicide Treatments in Hawaii Volcanoes National Park. Cooperative National Park Resources Studies
Unit, University of Hawaii at Manoa, Dept. of Botany
Singh SK, Singh PN and Bhalla K (1997) Novel taxa of Pseudocercosporella from Nepal Himalaya. MycologicalResearch 101: 542-544
Stratton L (1996) The Impact and Spread of Rubus ellipticus in 'Ola'a Forest Tract Hawaii Volcanoes National Park.
Technical Report 107, p 35, Cooperative National Park Resources Studies Unit, University of Hawaii at Manoa;
Western Region, National Park Service, Honolulu, Hawaii 96822
Su M, liu X, Yang F, Zhang Q and Xu S (2003) Experiment on the control of Epinotia rubiginosana with harmlesspesticides. Forest Pest and Disease 22: 9-11
Sun S, Wu K, Chen F and Wang L (1998) Lists of Insects in the forest areas of Changbai Mountain I: Geometridae. pp28-31
Sutton BC (1993) Mitosporic fungi from Malawi. Mycological Papers 167: 93
Sydow P (1897) Sylloge Fungorum, 543 pp
Tan J, Yu P, Li H and Wang S (1980) Economic Insect Fauna of China. Science Press, Beijing, China, 213 pp
Thind KS and Sharma MP (1980) Journal of the Indian Botanical Society 59: 350

Thomas MB and Reid AM (2007) Are exotic natural enemies an effective way of controlling invasive plants? Trends in
Ecology & Evolution 22: 447-453
US Forest Service PIEaRP (2007) Online resource at http://www.hear.org/pier/Wei S-x (1988) A taxonomic study of the genus Phragmidium of China. Mycosystema 1: 179-210
Wu K, Sun S, Chen F, Wang L and Yu B (1999) A List of the Insects in the Forest areas of Changbai Mountains II:
Noctuidae. pp 34-36, 40
Xi J, Pan H, Chen Y and Zhang X (2002) A List of Geometridae in Jilin Province. Journal of Jilin Agricultural
University 24 (5): 53-57
Xue D and Zhu H (1999) Fauna Sinica. Insecta. Science Press, Beijing, China, 1079 pp
Yu P, Wang S and Yang X (1996) Economic Insect Fauna of China. Science Press, Beijing, China, 324 pp
Yun Y (1998) Flora Fungorum Sinicorum. Science Press, Beijing, China, 530 pp
Zhang G and Zhong T (1983) Economic Insect Fauna of China. Science Press, Beijing, China, 387 pp
Zhang S (1985) Economic Insect Fauna of China. Science Press, Beijing, China, 242 pp
Zhao Y and Chen Y (1980) Economic Insect Fauna of China. Science Press, Beijing, China, 184 pp
Zheng H, Wu Y, Ding J, Binion D, Fu W and Reardon R (2005) Invasive Plants Established in the United States that are
Found in Asia and Their Associated Natural Enemies. USDA Forest Service - Forest Health Technology EnterpriseTeam
Zhou Y (1994) Monographia Rhopalocerrum Sinensium. Henan Scientific and Technological Publishing House,
Zhengzhou, Henan, China, 854 pp
Zhu H and Chen Y (1962) Economic Insect Fauna of China. Science Press, Beijing, China, 172 pp
Zhu H, Yang J, Lu J and Chen Y (1964) Economic Insect Fauna of China. Science Press, Beijing, China, 183 pp
Zhu HCH-f and Wang L (1996) Fauna Sinica. Insecta. Beijing, China, Science Press, 302 pp
Zhuang W-y (1987) Notes on some inoperculate discomycetes in Jamaica. Mycotaxon 29: 393-397