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Leaf Rust Caused by Kuehneola uredinis on Native and Nonnative ...

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Table 1. Infectivity of <str<strong>on</strong>g>Kuehneola</str<strong>on</strong>g> <str<strong>on</strong>g>uredinis</str<strong>on</strong>g> <strong>on</strong> native <strong>and</strong> noxious exotic Rubus species in Hawaii in the greenhouse (Table 1). We c<strong>on</strong>sider<br />

Species<br />

Seives<br />

infecti<strong>on</strong>'<br />

Field<br />

i<str<strong>on</strong>g>uredinis</str<strong>on</strong>g>,<br />

Greenhouse<br />

infecti<strong>on</strong> b<br />

Detached leaf<br />

infecti<strong>on</strong>c<br />

it likely that both native Rubus species<br />

are <strong>on</strong>ly slightly susceptible to K.<br />

with infectivity determined <str<strong>on</strong>g>by</str<strong>on</strong>g><br />

<strong>Native</strong><br />

R. hawaiiensis<br />

N<strong>on</strong>native<br />

+(l)d _(1)<br />

subtle factors of the envir<strong>on</strong>ment or of<br />

host or pathogen physiology. Thus, it is<br />

probable that R. macraei may also be<br />

R. ellipticus<br />

R. glaucus -<br />

R. penetrans +f + +<br />

R. rosaefolius - -<br />

aNatural infecti<strong>on</strong> observed <strong>on</strong> the isl<strong>and</strong>s of Hawaii, Maui, <strong>and</strong> Kauai.<br />

bPlants were inoculated <str<strong>on</strong>g>by</str<strong>on</strong>g> spraying with aqueous urediniospore suspensi<strong>on</strong>s of approximately 4 X<br />

attacked in the field under suitable<br />

c<strong>on</strong>diti<strong>on</strong>s.<br />

Aside from R. penetrans, our inoculati<strong>on</strong>s<br />

produced no infecti<strong>on</strong> <strong>on</strong> any of the<br />

other n<strong>on</strong>native species tested. These<br />

results are c<strong>on</strong>sistent with our field<br />

10' spores per milliliter to presaturati<strong>on</strong> <strong>and</strong> maintained under high (90-100%) relative humidity.<br />

Results were observed after about 3 wk.<br />

cLeaves were inoculated as in the greenhouse <strong>and</strong> maintained in moist petri dishes at room<br />

temperature or incubated at 17-19 C. Results were observed after about 3 wk.<br />

d+(l) Indicates light leaf infecti<strong>on</strong>s (ie, <strong>on</strong>e to several 1.0-mm-diameter discrete uredinial sori<br />

without epiphyllous symptoms).<br />

Indicates no infecti<strong>on</strong>.<br />

observati<strong>on</strong>s of these plants (Table 1).<br />

Our observati<strong>on</strong>s indicate that although<br />

K. <str<strong>on</strong>g>uredinis</str<strong>on</strong>g> exerts some impact <strong>on</strong> R.<br />

penetrans, the rust is not capable of<br />

severely curtailing this host in Hawaii<br />

under normal c<strong>on</strong>diti<strong>on</strong>s. The pathogen,<br />

however, has become well established <strong>on</strong><br />

+ Indicates a range of infecti<strong>on</strong> severity including heavy infecti<strong>on</strong> (ie, extensive col<strong>on</strong>izati<strong>on</strong> of leaf<br />

undersurfaces <str<strong>on</strong>g>by</str<strong>on</strong>g> c<strong>on</strong>vergent uredinia <strong>and</strong> corresp<strong>on</strong>ding epiphyllous symptoms).<br />

R. penetrans <strong>and</strong> is distributed throughout<br />

the Rh pnrans isl<strong>and</strong>s. It may <strong>and</strong> act diti is in c<strong>on</strong>juncti<strong>on</strong> uted th with<br />

R. hawaiiensis, the more comm<strong>on</strong> of<br />

the two native species, is also attacked <str<strong>on</strong>g>by</str<strong>on</strong>g><br />

K. <str<strong>on</strong>g>uredinis</str<strong>on</strong>g> in the field, although to a<br />

significantly lesser extent than is R.<br />

penetrans. Infecti<strong>on</strong> of R. hawaiiensis<br />

results <strong>on</strong>ly in <strong>on</strong>e to several small (1 mm<br />

diam.) individual, hypophyllous uredinial<br />

sori per leaf that remain discrete. Aside<br />

from the uredinia themselves, no<br />

symptoms or other detrimental results of<br />

infecti<strong>on</strong> are manifest <strong>on</strong> R. hawaiiensis.<br />

In agreement with Stevens' report (6), we<br />

have not observed K. <str<strong>on</strong>g>uredinis</str<strong>on</strong>g> <strong>on</strong> R.<br />

macraei in the field. This species is rare,<br />

however, <strong>and</strong> <strong>on</strong>ly infrequently encountered.<br />

Likewise, we have observed no<br />

infecti<strong>on</strong> <strong>on</strong> R. ellipticus Sm. (yellow<br />

Himalayan raspberry), R. rosaefolius<br />

Sm. (roseleaf raspberry), or R. glaucus<br />

Benth. (wild raspberry) in Hawaii, even in<br />

areas where infecti<strong>on</strong> <strong>on</strong> near<str<strong>on</strong>g>by</str<strong>on</strong>g> R.<br />

penetrans plants is heavy. We have found<br />

no c<strong>on</strong>clusive published report of K.<br />

<str<strong>on</strong>g>uredinis</str<strong>on</strong>g> attacking any of these three<br />

Rubus species elsewhere, although R.<br />

ellipticus <strong>and</strong> R. rosaefolius are implied<br />

as hosts of this rust in the USDA plant<br />

disease index (7). Like R. penetrans, these<br />

are secis ndeirale ocuring xotc<br />

are secis ndeirale ocuring xotc<br />

within or near Hawaii's NPS <strong>and</strong> other<br />

natural areas.<br />

To assess the infectivity of K. <str<strong>on</strong>g>uredinis</str<strong>on</strong>g><br />

<strong>on</strong> each of the menti<strong>on</strong>ed native <strong>and</strong><br />

n<strong>on</strong>native Rubus species under uniform<br />

c<strong>on</strong>diti<strong>on</strong>s, rooted cuttings of each<br />

species were established in pots in the<br />

greenhouse <strong>on</strong> a mist bench or were<br />

covered with plastic bags for 48 hr after<br />

X 10 spores per milliliter. Inoculum was<br />

collected from various R. penetransinfested<br />

sites <strong>on</strong> the isl<strong>and</strong>s of Hawaii <strong>and</strong><br />

Maui. Detached leaves of each species<br />

were also tested <str<strong>on</strong>g>by</str<strong>on</strong>g> placing leaves of<br />

comparable age <strong>and</strong> developmental state<br />

in moist petri dishes <strong>and</strong> spraying them<br />

with spore suspensi<strong>on</strong>s of similar<br />

c<strong>on</strong>centrati<strong>on</strong>s. The leaves were maintained<br />

at room temperature in the<br />

laboratory or at 17-19 C in a lighted<br />

incubator,<br />

RESULTS AND DISCUSSION<br />

R. penetrans became readily infected<br />

under greenhouse c<strong>on</strong>diti<strong>on</strong>s, with<br />

uredinia developing about 3 wk after<br />

inoculati<strong>on</strong>. Exp<strong>and</strong>ing or mature leaves<br />

appeared most susceptible, whereas<br />

newly formed unexp<strong>and</strong>ed leaves were<br />

not infected. The disease was perpetuated<br />

in the greenhouse <strong>on</strong>ly under c<strong>on</strong>diti<strong>on</strong>s<br />

of high humidity. As in the field, infecti<strong>on</strong><br />

was limited to undersurfaces of the leaves<br />

<strong>and</strong> yellowing, reddening, <strong>and</strong> necrosis<br />

occurred <strong>on</strong> corresp<strong>on</strong>ding upper areas.<br />

No stem or fruit infecti<strong>on</strong> occurred in the<br />

greenhouse. Detached leaves of this<br />

species also became infected (Table 1),<br />

withureini liewie apeaingabot 3<br />

withureini liewie apeaingabot 3<br />

wk after inoculati<strong>on</strong>. Light <strong>and</strong> temperature<br />

c<strong>on</strong>diti<strong>on</strong>s of incubati<strong>on</strong> did not<br />

appear critical to this development,<br />

R. macraei became lightly infected in<br />

the greenhouse even though natural<br />

infecti<strong>on</strong> in the field had not been found<br />

in our limited observati<strong>on</strong>. Individual,<br />

scattered uredinial s<strong>on</strong>iwere produced <strong>on</strong><br />

several leaves without apparent negative<br />

other purposely introduced bioc<strong>on</strong>trol<br />

agents in impairing the growth <strong>and</strong> vigor<br />

of this species. The aggressiveness <strong>and</strong><br />

competitive advantage of R. penetrans in<br />

favor of native plants may there<str<strong>on</strong>g>by</str<strong>on</strong>g> be<br />

lessened. Although the two endemic<br />

Rubus species show some susceptibility<br />

to K. <str<strong>on</strong>g>uredinis</str<strong>on</strong>g>, the minimal effect <strong>on</strong> these<br />

species indicates a genetic differential<br />

between these <strong>and</strong> R. penetrans that is<br />

regarded positively. The possibility that<br />

various races of K. <str<strong>on</strong>g>uredinis</str<strong>on</strong>g>, such as those<br />

known am<strong>on</strong>g other rusts, exist in Hawaii<br />

was not c<strong>on</strong>sidered in this study.<br />

Attenti<strong>on</strong> may be directed toward the<br />

determinati<strong>on</strong> of such races if future<br />

observati<strong>on</strong>s indicate that they occur.<br />

Further work toward the bioc<strong>on</strong>trol of<br />

undesirable Rubus species in Hawaii with<br />

other rusts, introduced for this purpose,<br />

is planned for the future.<br />

ACKNOWLEDGMENTS<br />

We thank Clifford Smith for his c<strong>on</strong>tributi<strong>on</strong><br />

to this project. We also acknowledge the technical<br />

assistance of D ea Kageler <strong>and</strong> Calen Miyahara.<br />

LITERATURE CITED<br />

States <strong>and</strong> Canada. Purdue Research Foundati<strong>on</strong>.<br />

West Lafayette, IN. 438 pp.<br />

2. Fischer, 0. W., <strong>and</strong> Johns<strong>on</strong>. F. 1950. Cane <strong>and</strong><br />

leaf rust, <str<strong>on</strong>g>Kuehneola</str<strong>on</strong>g> uredini~s (link) Arth.. of<br />

blackberries in western Washingt<strong>on</strong>. Phyto-<br />

pathology 40:199-204.<br />

3. Gardner, D. E., <strong>and</strong> Davis, C. J. 1982. The<br />

prospects for biological c<strong>on</strong>trol of n<strong>on</strong>native<br />

plants in Hawaiian nati<strong>on</strong>al parks. Nat. Park<br />

Serv., CPSU/ Univ. Hawaii Tech. Rep. 45. 55 pp.<br />

4. Hasan, 5. 1980. Plant pathogens <strong>and</strong> biological<br />

c<strong>on</strong>trol of weeds. Rev. Plant Pathol. 59:349-356.<br />

5. St. John, H. 1973. List <strong>and</strong> Summary of the<br />

Flowering Plants in the Hawaiian Isl<strong>and</strong>s. Mem.<br />

1. Pacific Tropical Botanical Garden, tLawai, HI.<br />

519 pp.<br />

inoculati<strong>on</strong> to ensure high (90-100%) impact <strong>on</strong> the plants. Detached leaves of<br />

relative humidity. The plants were R. macraei did not become infected.<br />

spraed o pesaurai<strong>on</strong>wit frshl C<strong>on</strong>erslydetche levesof .<br />

spraed wih t prsatuatin feshl C<strong>on</strong>ersly, etahed eavsofR.<br />

collected urediniospores in water hawaiiensis became lightly infected but<br />

suspensi<strong>on</strong>s at c<strong>on</strong>centrati<strong>on</strong>s of about 4 no infecti<strong>on</strong> was obtained <strong>on</strong> this species<br />

~<br />

6. Stevens, F. L. 1925. Hawaiian Fungi. Bernice P.<br />

Bishop Museum Bull. 19. Bishop Museum,<br />

H<strong>on</strong>olulu. 189 pp.<br />

U. us. Department of Agriculture. 1960. index of<br />

Plant Diseases in the United States. Agric.<br />

H<strong>and</strong>bk. 165. 531 pp.<br />

Plant Disease/September 1983 963

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