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Acacia Koa: a Review of Its Diseases and Associated Fungi

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Acacia Koa: a Review of Its Diseases and Associated Fungi ~ Koa:ADecadeofGrowth. Acacia koa: A Review of its Diseases and Associated Fungi Donald E. Gardner, Biological Resources Division, USGS Department of Botany, University of Hawai'i at Manoa Abstract leguminous species, koa is thought to account signifi- Acacia koa (koa), among the most prominent over- cantly for the nitrogen content of otherwise nitrogen- story species of native Hawaiian forests, is affected by a poor volcanic forest soils (Whitesell 1990). Koa is also number of diseases, including those caused by rust fungi, prominently represented in early Hawaiian legends and wood-rotting fungi, root-infecting fungi, and diseases culture, and economically is considered the most valu- categorized as seedling blights, foliar infections, and able of the common native timber species. Koa wood, vascular wilts. A number of fungi occurring saprophyti- sometimes referred to as "Hawaiian mahogany," can be cally on koa substrates are also known. Symptoms of highly polished to emphasize its deep reddish colora- other apparent diseases and disorders of unknown ori- tion and wavy grain, and is used for furniture, paneling, gin, or whose cause has not been clearly demonstrated, and such woodworking crafts as bowls and ukuleles. are often manifest as leaf spots or other foliar abnor- St. John (1979) recognized three distinct species as malities. Endemic pathogens which have evolved with representing the genus Acacia in Hawai'i: A. koa Gray, their host may be responsible for many of the hereto- A. koaia Hbd., and A. kauaiensis Hbd. Furthermore. fore observed disease conditions, and as such usually within A. koa three varieties were recognized: A. koa do not threaten koa populations. However, a condition var. koa, found on all major Hawaiian islands; A. koa tentatively referred to as "koa decline" may represent var.latifolia (Benth.) St. John (=A. koa var. hawaiiensis an exception. This condition is characterized by slow to Rock); and A. koa var. waianaiensis St. John; the latter rapid wilt and death of apparently healthy, vigorous trees two restricted to the islands of Hawai'i and O'ahu, re- of all ages, occurring in more or less well defined dis- spectively. Although the current treatment of A. koa ease centers. This phenomenon is particularly promi- (Wagner et al. 1990) combines all of these forms within nent in upper-elevation koa forests on the slopes of A. koa, the distinction among them is still apparently Mauna Loa on the island of Hawai'i. Other apparent sufficient to delimit the host ranges of host-specific decline problems on O'ahu and elsewhere in the Islands pathogens, such as the rust fungi. Reference to St. John's have been reported, some associated with insects. The classification is therefore useful to this discussion. relationship of these observations to the koa decline on Several observations of diseases and pathogenic the Big Island has yet to be determined, and the ques- fungi occurring on koa have been individually reported tion as to whether "koa decline" represents a complex in the literature as they have been observed over the years of disease conditions or can be more closely attributed (Raabe et al. 1981), or have not been formally published. to a single, specific cause requires further study. It was of interest to assemble such published informa- Koa (Acacia koa Gray), endemic to the Hawaiian tion, together with as yet unpublished observations of Islands, is the second most abundant overstory species fungi and disease conditions ofkoa, to provide the basis in upper-elevation forests. It forms nearly pure stands for a discussion of the pathology of this pivotal species. in montane forests such as those on the slopes of Mauna Loa between approximately 4000 and 7000 ft (1230 and Diseases caused by rust fungi 2150 m) (Whitesell 1990). Koa is distinctive in the for- Rust fungi, members of the order Uredinales, are so mation of crescent-shaped phyllodes which, in older called because of the often brownish to orange appear- trees, assume the function of the pinnately compound ance of the spore masses. These fungi produce perhaps true leaves produced in juveni le stages of development. the most widespread and prominent diseases of koa, As an endemic tree, koa is of critical ecological collectively referred to as koa rust. Stevens (1925), origi- importance, forming habitat for numerous native birds, nally described a single fungus, Uromyces koae Arth., insects, and other flora and fauna. As a nitrogen-fixing as the cause of the disease, but five species of rust fungi. 56 Koa:ADecadeq(Grou,th ~ four of which are considered endemic and one indig- koa var. latifolia and is limited in distribution on the enous, are currently recognized on koa (Hodges and island of Hawai'i, but visible in large koa trees on the Gardner 1984, Gardner 1991), each with distinctive upper Saddle Road. This rust forms large (up to 1 m symptoms. These include three species now placed in tall) witches' brooms comprised of hypertrophied but the genus Atelocauda, A. koae (Arth.) Cumm. & Y. abnormally reduced, much narrowed phyllodes almost Hiratsuka, A. digitata (Wint.) Cumm. & Y. Hiratsuka, circular in cross section, giving a "shoe-string" appear- and A. angustiphylloda Gardner, and two species of ance (Gardner, in press). As with A. digitata, brooms Endoraecium, E. acaciae Hodges & Gardner and E. are covered with powdery, brown spore masses. Al- hawaiiense Hodges & Gardner. Atelocauda koae occurs though limited in distribution, large numbers of brooms commonly on Acacia koa var.latifolia on Hawai'i. and (100+) may occur in certain mature, presumably sus- on A. koa var. koa on Kaua'i, O'ahu, and Maui. It has ceptible trees. Host trees appear to tolerate infection by not been reported on Moloka'i, nor has it been found A. angustiphylloda, but loss of vigor and local tissue on A. koaia anywhere. Atelocauda koae is confined to death may occur. young tissue, occurring most frequently on newly de- Endoraecium acaciae, the most common of koa veloping true leaves, phyllodes, and shoots of saplings, rusts, has been found onA. koa var. koa on Maui, Kaua'i, but also found on new shoots of older trees. It is evident and Hawai'i, and on A. koaia on Hawai'i, Kaua'i, and as raised, powdery, brown leaf spots, 1-10 mm diam- Moloka'i. It has not been found on A. koa var.latifolia. eter, occurring singly or in groups, on both leaf surfaces. This fungus produces conspicuous, profusely branched Heavy infection, in which spots coalesce to form large, brooms up to 30 em long comprised of much-reduced, irregular powdery blotches, can cause severe distortion flattened phyllodes covered with spore masses. In ap- of leaves, shoots, and small branches, leading to stem parent contrast to other koa rusts, E. acaciae may cause deformation (Hodges and Gardner 1984, Chen et al. severe damage to the host, and heavy infections pro- 1996). ducing multiple brooms may kill the entire tree. Brooms Atelocauda digitata is known from all major islands of E. hawaiiense are similar to those of E. acaciae but except Lanai, although its presence on this island in are smaller, being seldom taller than 12 cm, and rela- upland forests would not be surprising. On Acacia koa tively inconspicuous. This fungus appears to be limited var. latifolia, the conspicuous stages of this fungus cause in distribution, having been found only at certain sites "witches' -brooms" up to approximately 15 em tall, com- on O'ahu on A. koa var. waianaiensis and A. koa var. prised of clustered, misshapen, abnormally thickened koa. An insufficient number of trees heavily infected shoots developing from infected branches. The diseased with this fungus has been found to assess its potential tissue may be covered by powdery, brown spore masses for host damage. The two species of Endoraecium are that are easily rubbed off. Removal of witches' brooms further separated from one another by differences in is ineffective in controlling the disease, because the fun- spore morphology. gus invades branch tissues systemically. From one to Wind-distributed spores of the koa rust fungi are many brooms may occur in a tree. Less conspicuous thought to be the principal means of long- and short- stages of A. digitata occur as small, brown, raised sporu- distance dispersal, although Leeper arid Beardsley (1973) lating pustules up to approximately 1 mm diameter on noted the possibility that the koa psyllid Acizzia the surface of otherwise normal phyllodes. Occasion- uncatoides (Ferris & Klyver) [= Psylla uncatoides (Ferris ally such spots are surrounded by larger chlorotic spots, & Klyver)] may at least in part account for short-dis- causing them to be more conspicuous. Atelocauda tance dispersal where populations of these insects are digitata kills branch portions distal to the broom, but high. Wherever koa rust diseases are observed, evidence entire trees are typically not killed by this fungus alone. can be found of variability in susceptibility to infection On Acacia koa var. koa on Maui and O'ahu A. digitata among neighboring individual trees. Koa breeding pro- produces hypertrophy of flower and shoot tissue, result- grams directed at selecting such resistance may be the ing in malformation, but it does not produce witches' most effective control approach for rust diseases. brooms typical of those on Acacia koa var. latifolia .Hodges and Gardner 1984). Wood-rotting fungi Atelocauda angustiphylloda is confined to Acacia A number of diseases of koa caused by wood rot- 57 ~ Koa:A Decade of Grounb ting, "higher" fungi (i.e., basidiomycetes) have been dicate the true incidence of disease in a stand (Larsen described. These fungi are characterized by external et al. 1985). Rather than killing their hosts directly, production of fleshy or woody fruiting bodies which are pathogenic heart and root-rotting fungi destroy timber often conspicuous and indicate the presence of already usefulness and predispose trees to wind-throw and well-established internal infection.
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