Leaf Caused by uredinis on Native and Nonnative Rubus Species in Hawaii

DONALD E. GARDNER, Research Plant Pathologist, National Park Service Cooperative Park Studies Unit, University of Hawaii, Honolulu 96822, and CHARLES S. HODGES, Chief Plant Pathologist, Institute of Pacific Islands Forestry, Pacific Southwest Forest and Range Experiment Station, U.S. Forest Service, Honolulu, HI 96813 reports. ABSTRACT This investigation was conducted to Gardner, D. E., and Hodges, C. S. 1983. Leaf rust caused by on native and document the present distribution of K. nonnative Rubus species in Hawaii. Plant Disease 67:962-963. uredinis in Hawaii and its impact on native and nonnative Rubus species. attacks Rubuspenetrans, a Kuehneola uredinis is widely distributed in Hawaii, where it primarily noxious exotic species. Other nonnative Rubus species are not infected, however. R. macraei and MATERIALS AND METHODS R. hawaiiensis,both valued endemic species, are mildly and inconsequentially attacked by the rust. R. penetrans is universally distri Additional key words: biocontrol, Rubus ellipticus, R. glaucus, R. rosaefolius throughout Hawaii, occurring commonly on all major islands at elevations above 1,000 m. We have observed light to heavy infection by the uredinial state of K. Although individual residents of effective in some areas, but additional uredinis wherever this host has been Hawaii value exotic Rubus species for control measures are essential to prevent found. Infection on R. penetrans results their fruit, these plants have no economic further spread of this weed. in scattered and confluent pale lemon- usefulness as commercial crops and are We are currently investigating the use yellow hypophyllous uredinia with considered noxious weed pests in forest of plant pathogens, specifically rust associated yellowing or reddening of and pasture lands. The aggressive fungi, as biocontrol agents for Rubus corresponding upper leaf surface areas. invasion of such species into native spp. Rust diseases are typically host- Necrosis and defoliation may follow forests of Hawaii's state and national specific, often highly destructive, and moderate to heavy leaf infection. parks and other preserves poses a serious have been used with success elsewhere in Severely infected thickets frequently threat to the rare and fragile ecosystems biocontrol programs against other appear unthrifty and sparsely foliated, these areas were established to protect. noxious Rubus species (4). Cane and leaf but we have not observed evidence that On the other hand, endemic Hawaiian rust(Kuehneolauredinis(Lk.)Arth.)had the disease by itself is capable of Rubus species are highly valued as been suggested in earlier biocontrol significantly limiting the spread of R. components of such ecosystems and are considerations as a potentially effective penetrans. No stem or fruit infection has strenuously protected by such conservation agent worthy for testing in our program. been noted. agencies as the National Park Service At that time, however, little information We have observed the telial, sperm- (NPS). Control approaches for exotic was available concerning the status of this ogonial, and aecial states of K. uredinis in Rubus species must therefore be carefully rust in Hawaii. the autumn on R. penetrans. The telia evaluated for their potential detrimental K. uredinis was first reported in Hawaii were hypophyllous, whitish, and waxy in effects on native as well as undesirable by Stevens in 1925 (6) from a single appearance. Teliospores(14-22X80-100 target plants. location on the island of Maui, heavily gim) with four to seven cells were The most pervasive noxious Rubus attacking R. villosus. This Rubus species common. Those observed were often species in Hawaii is R. penetrans Bailey is not currently recognized as such in fragmented and incomplete. Teliospore (prickly Florida blackberry). The Hawaii and was probably used as a walls were colorless. The aecia were of R. penetrans is somewhat synonym of R. penetrans (H. St. John, uredinoid, epiphyllous, orange-yellow, unsettled. St. John (5) acknowledges that personalcommunication).R. villosus was and surrounded the prominent sperm- this species has been variously placed in thought by Stevens to have been recently ogonia, which occurred on reddish leaf synonymy with three other species but introduced to Hawaii because it was spots (1). Aeciospores were verrucose refers to it as a cultigen that does not localized in distribution and no published and measured 16-19 X 20-23 #.m. exactly match any of these. Other reference to its presence in Hawaii could Urediniospores resembled aeciospores in investigators (D. Hall, personal be found. Stevens (6) further reported morphology but were more strongly communication) consider R. penetrans that R. macraei Gray, a native Hawaiian echinulate and measured 17-21 X 22-28 synonymous with R. argutus Link, a species, was free of rust infection even rem. Dimensions of 18-24 X 85-110 gm native to the southeastern United States. though it grew near the infected R. for teliospores, 18-19 X 19-23 #im for Past attempts to control R. penetrans in villosus. aeciospores, and 16-19 X 21-27/•.m for Hawaii have involved importation and The status of K. uredinis in Hawaii has urediniospores were given for K. uredinis release of various species of phytophagous not been discussed in the literature since by Arthur (1). Although locally common, insects (3). These have been partially Stevens' report. Herbarium specimens the spermogonial, aecial, and telial states (USDA National Collections, occur infrequently compared with the Accepted for publication 28 February 1983. Beltsville, MD) indicate, however, that uredinial state, which can be found ______the uredinial state of the rust was throughout the year. The publication costs of this article were defrayed in part collected on R. hawaiiensis Gray, a Irregular white patches of nonviable, by page charge payment. This article must theretore be hereby marked "advertisement" in accordance with 18 second native species, on the island of apparently hyperparasitized uredini- U.S.C. §1734 solely to indicate this fact. Kauai in 1928 and on R. penetrans on ospores similar to those reported by Oahu in 1951. A reference to K. uredinis Fischer and Johnson (2) on Rubus This article is inthe public domain and not copy- on R. hawaiiensis in the USDA plant species in western Washington are rightable. It may be freely reprinted with cus- tomary crediting of the source. The American disease index (7) is apparently based on commonly seen and can be confused with Phytopathological Society, 1983. herbarium records or unpublished telial sori on casual observation. 962 Plant Disease/Vol. 67 No. 9 Table 1. Infectivity of Kuehneola uredinis on native and noxious exotic Rubus species in Hawaii in the greenhouse (Table 1). We consider Rubus species Greenhouse Detached leaf it likely that both native Species infection'Field infection b infectionc are only slightly susceptible to K. Seives iuredinis, with infectivity determined by Native subtle factors of the environment or of R. hawaiiensis +(l)d _(1) host or pathogen physiology. Thus, it is Nonnative probable that R. macraei may also be under suitable R. ellipticus attacked in the field R. glaucus - conditions. R. penetrans +f + + Aside from R. penetrans, our inocula- R. rosaefolius - - tions produced no infection on any of the aNatural infection observed on the islands of Hawaii, Maui, and Kauai. other nonnative species tested. These bPlants were inoculated by spraying with aqueous urediniospore suspensions of approximately 4 X results are consistent with our field 10' spores per milliliter to presaturation and maintained under high (90-100%) relative humidity. observations of these plants (Table 1). Results were observed after about 3 wk. Our observations indicate that although cLeaves were inoculated as in the greenhouse and maintained in moist petri dishes at room K. uredinis exerts some impact on R. temperature or incubated at 17-19 C. Results were observed after about 3 wk. penetrans, the rust is not capable of d+(l) Indicates light leaf infections (ie, one to several 1.0-mm-diameter discrete uredinial sori severely curtailing this host in Hawaii without epiphyllous symptoms). under normal conditions. The pathogen, Indicates no infection. however, has become well established on + Indicates a range of infection severity including heavy infection (ie, extensive colonization of leaf R. penetransand is distributed throughout undersurfaces by convergent uredinia and corresponding epiphyllous symptoms). theRh pnrans islands. It mayand actditi is in conjunctionuted th with other purposely introduced biocontrol R. hawaiiensis, the more common of X 10 spores per milliliter. Inoculum was agents in impairing the growth and vigor the two native species, is also attacked by collected from various R. penetrans- of this species. The aggressiveness and K. uredinis in the field, although to a infested sites on the islands of Hawaii and competitive advantage of R. penetrans in significantly lesser extent than is R. Maui. Detached leaves of each species favor of native plants may thereby be penetrans. Infection of R. hawaiiensis were also tested by placing leaves of lessened. Although the two endemic results only in one to several small (1 mm comparable age and developmental state Rubus species show some susceptibility diam.) individual, hypophyllous uredinial in moist petri dishes and spraying them to K. uredinis, the minimal effect on these sori per leaf that remain discrete. Aside with spore suspensions of similar species indicates a genetic differential from the uredinia themselves, no concentrations. The leaves were main- between these and R. penetrans that is symptoms or other detrimental results of tained at room temperature in the regarded positively. The possibility that infection are manifest on R. hawaiiensis. laboratory or at 17-19 C in a lighted various races of K. uredinis, such as those In agreement with Stevens' report (6), we incubator, known among other rusts, exist in Hawaii have not observed K. uredinis on R. was not considered in this study. macraei in the field. This species is rare, RESULTS AND DISCUSSION Attention may be directed toward the however, and only infrequently encoun- R. penetrans became readily infected determination of such races if future tered. under greenhouse conditions, with observations indicate that they occur. Likewise, we have observed no uredinia developing about 3 wk after Further work toward the biocontrol of infection on R. ellipticus Sm. (yellow inoculation. Expanding or mature leaves undesirable Rubus species in Hawaii with Himalayan raspberry), R. rosaefolius appeared most susceptible, whereas other rusts, introduced for this purpose, Sm. (roseleaf raspberry), or R. glaucus newly formed unexpanded leaves were is planned for the future. Benth. (wild raspberry) in Hawaii, even in not infected. The disease was perpetuated areas where infection on nearby R. in the greenhouse only under conditions ACKNOWLEDGMENTS Smith for his contribution penetransplants is heavy. We have found of high humidity. As in the field, infection to Wethis thankproject. Clifford We also acknowledge the technical assistance of D eaKageler and Calen Miyahara. no conclusive published report of K. was limited to undersurfaces of the leaves and yellowing, reddening, and necrosis uredinis attacking any of these three LITERATURE CITED Rubus species elsewhere, although R. occurred on corresponding upper areas. occurred in the States and Canada. Purdue Research Foundation. ellipticus and R. rosaefolius are implied No stem or fruit infection as hosts of this rust in the USDA plant greenhouse. Detached leaves of this West Lafayette, IN. 438 pp. disease index (7). Like R. penetrans, these species also became infected (Table 1), 2. Fischer, 0. W., and Johnson. F. 1950. Cane and aresecis ndeirale ocuring xotc withureini liewie apeaingabot 3 leaf rust, Kuehneola uredini~s (link) Arth.. of aresecis ndeirale ocuring xotc withureini liewie apeaingabot 3 blackberries in western Washington. Phyto- within or near Hawaii's NPS and other wk after inoculation. Light and temper- pathology 40:199-204. natural areas. ature conditions of incubation did not 3. Gardner, D. E., and Davis, C. J. 1982. The To assess the infectivity of K. uredinis appear critical to this development, prospects for biological control of nonnative on each of the mentioned native and R. macraei became lightly infected in plants in Hawaiian national parks. Nat. Park Serv., CPSU/ Univ. Hawaii Tech. Rep. 45. 55 pp. nonnative Rubus species under uniform the greenhouse even though natural 4. Hasan, 5. 1980. Plant pathogens and biological conditions, rooted cuttings of each infection in the field had not been found control of weeds. Rev. Plant Pathol. 59:349-356. species were established in pots in the in our limited observation. Individual, 5. St. John, H. 1973. List and Summary of the Flowering Plants in the Hawaiian Islands. Mem. greenhouse on a mist bench or were scattered uredinial soniwere produced on 1. Pacific Tropical Botanical Garden, tLawai, HI. covered with plastic bags for 48 hr after several leaves without apparent negative 519 pp. inoculation to ensure high (90-100%) impact on the plants. Detached leaves of 6. Stevens, F. L. 1925. Hawaiian Fungi. Bernice P. relative humidity. The plants were R. macraei did not become infected. Bishop Museum Bull. 19. Bishop Museum, spraed o pesauraionwitfrshl Conerslydetche levesof . ~ Honolulu. 189 pp. spraedwih t prsatuatinfeshl Conersly, etahed eavsofR. U.us. Department of Agriculture. 1960. index of collected urediniospores in water hawaiiensis became lightly infected but Plant Diseases in the United States. Agric. suspensions at concentrations of about 4 no infection was obtained on this species Handbk. 165. 531 pp.

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