Miconia Biocontrol: Where Are We Going and When Will We Get There?

M. Tracy Johnson

Institute of Pacific Islands Forestry, Pacific Southwest Research Station, USDA Forest Service, Volcano, HI, Email: [email protected]

Abstract

We have made much progress in evaluating potential agents for biocontrol of , and several appear likely to be suitable for future introduction to Hawaiÿi. Unfortunately, none of them is an obvious silver bullet. We face the challenge of prioritizing the existing candidates and inventing the combination of agents that will achieve our goals. Now is an opportune moment to reassess our goals for miconia biocontrol and strategize how we will combine biocontrol and other management tools to successfully manage this weed for the long term. Our goals and choices for biocontrol should take into account the densities and distributions of existing miconia populations and projections for their expansion, as well as the sustainability and compatibility of other control methods. Critical areas of substantial uncertainty, where we may benefit from focused research, include: impacts and dispersal of biocontrol agents, potential interactions among agents, interactions between agents and their natural enemies, and dynamics of multi-melastome host use by agents. Some questions will be important to answer before releasing new agents; others may be essentially unanswerable until post-release. We should make our goals and strategies as explicit as possible, then adapt as we encounter future realities.

Summary of Biocontrol of Miconia to Date

The search for natural enemies of has been underway since 1993, when Robert Burkhart began his work in its native range. As exploratory entomologist of the Hawaiÿi Department of Agriculture (HDOA), he spent several months in Costa Rica and southern discovering and sampling a wide variety of sites. Considering the challenges of even locating the sparsely distributed M. calvescens – mainly restricted to lightly disturbed mid-elevation native forests – Burkhart uncovered a remarkable diversity of herbivores. Many live specimens were shipped back to Hawaiÿi, where some were reared to adulthood for later identification, but in the end none persisted long enough for detailed study within the HDOA quarantine. By 1994, he concluded explorations with the opinion that pathogens might perform better as biocontrol agents than the insect he observed attacking miconia (Burkhart 1995).

Subsequent entomological explorations from 1999 through 2007 revealed even more potential agents and important biological details. The work of Mohsen Ramadan (HDOA) in , Paul Hanson and students at the Universidad de Costa Rica, and Alec McClay in southern is summarized in these proceedings. Marcelo Picanço and students at the Universidade Federal de Viçosa in Minas Gerais, Brazil also added substantially to our knowledge of miconia’s insect fauna (Picanço et al. 2005). Much of this work based in the native range benefited from multiple years of intensive study by approximately 13 students and post-docs who selected one or more miconia as the focus of their research.

2009 International Miconia Conference Johnson • 1 Similarly, the pathogens of Miconia calvescens have been the focus of long-term studies under the direction of Robert Barreto since the mid 1990s. This work, mostly in Brazil but including considerable efforts elsewhere in the native range, has yielded the one biocontrol agent released to date, Colletotrichum gloeosporioides f. sp. miconiae, and a list of other pathogens with varying potential (Barreto et al. 2005, Seixas et al. 2007, Alves et al. 2009). Most promising among these are a fungus, Coccodiella miconiae, that may be more damaging than C. gloeosporioides, and a nematode, Ditylenchus gallaeformans, that deforms a variety of melastomes with impressive galls.

Priorities Among Candidate Agents

Given the intensity of work in Costa Rica and Brazil, with weekly or monthly field visits over a period of several years, we can be confident that very few additional herbivores remain undiscovered in these areas of the native range. From the dozens of insect species discovered, we can now define a short list of potential agents that appear likely to be host- specific and have substantial impact on miconia. The process of prioritizing insect species for further research has been based on qualitative observations of the damage they cause to in the native range, on their observed or expected host specificity, and on the likelihood of natural enemies already present in Hawaiÿi interfering with their population growth (Table 1).

Table 1. Potential for biological control among insects feeding on Miconia calvescens in its native range.

Interference Overall Impact Host Order Family Insect species Country1 by natural potential for on plant2 specificity3 enemies4 biocontrol5 Flower and/or fruit feeding Diptera Cecidomyiidae unidentified flower midge C 3 unknown 1 5 Coleophoridae Mompha sp. C 4 4 1 7 Lycaenidae Erora opisena C 5 unknown 2 6 Temecla paron C 5 unknown 2 6 Parrhasius polibetes C 5 2 2 5 Coleoptera Curculionidae Anthonomus monostigma C 4 5 1 8 Apion sp. B 4 4 1 7 Pedetinus halticoides C 3 unknown 1 5 Hymenoptera Braconidae Allorhogas sp. B 4 5 1 8 Leaf feeding Coleoptera Chrysomelidae Typophorus variabilis C 2 3 1 4 Margaridisa sp. C 2 unknown 1 4 Percolaspis sp. C 2 2 1 3 Curculionidae Exophthalmus jekelianus C 2 2 1 3 Penestes sp. C 2 2 1 3 Hymenoptera Argidae Atomacera petroa B,C 3 5 1 7 Formicidae Atta sexdens rubropilosa B,C 5 1 1 5 Lepidoptera Zanola impedita C 2 2 3 1 Arctiidae Melese sp. C 2 2 2 2 Salbia lotanalis B,C 5 4 3 6 Geometridae Isochromodes sp. C 2 unknown 3 2 unidentified C 2 unknown 3 2 Gracillariidae unidentified leaf miner C 1 unknown 2 2 Limacodidae Vipsophobetron marisa C 2 2 2 2 Isa diana C 2 2 2 2 Talima aurora C 2 2 2 2 Parasa imatata C 2 2 2 2 Euclea zygia C 2 2 2 2 Epiperola paida C 2 2 2 2

2009 International Miconia Conference Johnson • 2 Interference Overall Impact Host Order Family Insect species Country1 by natural potential for on plant2 specificity3 enemies4 biocontrol5 Natada sp. C 2 2 2 2 Lycaenidae Theritas mavors C 3 unknown 2 4 Druentia cf. inscita B 4 unknown 2 5 Antiblemma leucocyma B 4 4 3 5 Antiblemma sp. C 4 unknown 3 4 Plusiinae sp. C 2 unknown 3 2 Notodontidae Naprepa houla C 4 2 3 3 Rhuda difficilis C 3 unknown 3 3 Meragisa sp. C 3 unknown 3 3 Oecophoridae unidentified (2 species) C 2 unknown 3 2 Psychidae unidentified C 1 2 2 1 Pterophoridae unidentified C 1 unknown 2 2 Euselasia chrysippe C 5 5 2 8 Euselasia bettina C 5 5 2 8 Euselasia aurantia C 5 unknown 2 6 Anteros formosus micon C 2 unknown 2 3 Ancyluris inca C 2 unknown 2 3 Symmachia tricolor C 2 unknown 2 3 Saturniidae Hylesia continua C 3 2 3 2 Orthoptera Gryllidae unidentified crickets C 2 1 2 1 Thysanoptera Thripidae Heliothrips sp. B 2 2 2 2 Sap feeding Hemiptera Aleyrodidae unidentified whiteflies B,C 1 2 3 0 Clastopteriadae Clastoptera sp. C 1 2 1 2 Cicadellidae Empoasca sp. B 1 2 2 1 Scaphytopius sp. B 1 2 2 1 Membracidae Bolbonata sp. C 1 unknown 1 3 Micrutalis sp. C 1 unknown 1 3 Pseudococcidae unidentified mealybugs B,C 1 2 2 1 Psyllidae Diclidophlebia lucens C 2 5 2 5 Diclidophlebia smithi B 2 5 2 5 Stem feeding Coleoptera Buprestidae Agrilus sp. B 4 unknown 1 6 Cerambycidae Platyarthoron chilense C 4 unknown 1 6 Curculionidae Cryptorhynchus melastomae C 5 4 1 8 Naupactus spp. B 3 2 1 4 Copturus tricolor C 1 2 1 2 1 C=Costa Rica, B=Brazil 2 Impact (observed, or potential based on type of damage) scored 1=low, 2=low to moderate, 3=moderate, 4=moderate to high, 5=high. 3 Specificity (observed, or expectation based on related species) scored 1=very low, 2=low, 3=moderate, 4=high, 5=very high. 4 Risk of interference by natural enemies already in Hawaii scored as 1=low, 2=moderate, 3=high. 5 Overall potential calculated as Impact + Specificity - Interference (with unknown Specificity assigned a value of 3).

Although host specificity is widely recognized as a key trait of biocontrol agents, necessary for ensuring their environmental safety, potential effectiveness is also an important criterion to include early in the evaluation process. Too frequently, biocontrol agents have been developed for release because they are host-specific and easy to rear, but with little real potential for suppressing a target weed (McClay and Balciunas 2005). Our evaluation of miconia insects presents the example of two psyllid species in the Diclidophlebia, both of which appear to be highly specific to miconia and a few related melastomes (Burckhardt et al. 2005, Morais et al. 2008). However, neither species appears to seriously limit growth of miconia, even under conditions of heavy infestation of potted greenhouse plants. In addition, lab tests have shown that D. lucens is vulnerable to a common lady

2009 International Miconia Conference Johnson • 3 beetle that already limits other psyllids in Hawaiÿi (B. Wai unpub. data). Since miconia psyllids appear to have a low probability of developing into damaging populations in Hawaiÿi, they have been downgraded as potential agents in favor of other species with greater promise of impact on the target.

Some prospective agents can be judged by impacts of outbreaks witnessed on miconia in its native range. For example, in both Brazil and Costa Rica the leaf roller Salbia lotanalis (synonym: Ategumia lotanalis) occasionally causes defoliation to a degree that clearly impacts the host in the field, and larvae can severely damage potted plants under artificial conditions (Picanço et al. 2005, Badenes-Perez et al. 2008). However, for the great majority of insect species, population densities remain low on the dispersed miconia in the native range. In these cases the potential impacts of high infestations must be inferred by extrapolating the damage observed from individual feeding insects or insects confined artificially. Several of the species that feed on flowers and fruit fit in this category, since their populations are typically very low in the native range, but their impacts on individual inflorescences or fruit can be substantial (Badenes-Perez and Johnson 2007a, Badenes-Perez et al. in press). Explosive population growth would be required for these species to have meaningful impacts in Hawaiÿi and Tahiti, especially given the prolific seed production of miconia. Fortunately at least one factor, the year-round flowering and fruiting of miconia in Pacific islands (Meyer 1998), seems to favor the potential for population increases by these insects beyond levels seen in the native range, where miconia reproduction is highly seasonal (Picanço et al. 2005, Chacón 2007).

Interference by natural enemies can hamper weed biocontrol efforts using otherwise suitable agents, especially in Hawaiÿi, where a large number of introduced generalist and specialist enemies are active throughout the year (Goeden and Louda 1976, Julien and Griffiths 1998). Such interference has been cited often for limiting effectiveness of lepidopteran weed control agents in Hawaiÿi (Fullaway and Krauss 1945, Reimer and Beardsley 1986, Davis et al. 1992). Although the predictive power of such an analysis remains to be proven, it seems appropriate to take into account the relative likelihood of biotic interference when evaluating potential agents. For example, relatives of the leaf feeders Salbia lotanalis and Antiblemma leucocyma are known to be attacked by a variety of parasitoids in Hawaiÿi (Zimmerman 1958a, 1958b, Conant 2002, 2009). Given the prospects of heavy parasitism for these species, they have been scored as having relatively poor overall potential for miconia control (Table 1), in spite of demonstrated impacts and apparent specificity (Picanço et al. 2005, Badenes-Perez and Johnson 2008, Castillo 2009, E. Morais unpubl. data).

Strategy for Moving Forward With High Priority Agents

Since none of our prospective agents is obviously devastating to miconia within its native range, it is difficult to imagine that any single agent can provide complete control on its own. Instead, we can expect that combined action of several effective biocontrol agents will be necessary for management of miconia in the long term. Substantial complementary impacts by two or more agents are commonly a goal of biocontrol programs. Based on recent successes with biocontrol of acacias in South Africa, combinations of agents seem particularly appropriate for fast-spreading invasive (Hoffmann et al. 2002, Post et al. 2010). Our current strategy for miconia is to develop a suite of agents that attack different parts of the , including stems, and reproductive structures, with the goal of damaging miconia in multiple ways to lower its overall fitness (Badenes-Perez et al. 2008).

2009 International Miconia Conference Johnson • 4 Eight insect species emerge as the highest priorities for development of miconia biocontrol, scoring seven or eight out of a possible nine points maximum for overall potential (Table 1). A few additional species may be worth considering in the future if more information on their host specificity becomes available. Together with the two most promising miconia pathogens, there are ten potential new agents to consider (Table 2). Within this short list, future work must be further prioritized to make best use of limited resources and produce the greatest benefit in terms of miconia management.

Table 2. Overview of species with high potential for biocontrol of miconia. Agent Impact Remaining effort Potential drawbacks Euselasia chrysippe, Larvae moving as a Need method for mating Larvae and adults may be E. bettina family group consume adults in captivity, or must vulnerable to generalist whole leaves complete specificity tests predators with field-collected insects Atomacera petroa Larvae rasp surface of Need rearing methods, or Damage to older leaves may mature leaves, must complete specificity be similar to current effects of reducing area for tests with field-collected Colletotrichum; plants may photosynthesis insects compensate; larvae exposed Cryptorhynchus Adults damage leaves Assess risk of potential Long life cycle may slow melastomae and stems, larvae bore nontarget host use agent population increase stems Anthonomus Adults chew on Need fruiting trees for Dependence on developing monostigma immature fruit, larvae rearing and specificity fruit may limit agent consume developing tests populations seasonally seeds Apion sp. Larvae consume Need species identified Dependence on developing developing seeds /described, rearing fruit may limit agent methods, and fruiting trees populations seasonally for specificity tests Mompha sp. Larvae damage Need species described Some seed survival is likely; developing flower and fruiting trees for could be vulnerable to buds and bore inside rearing and specificity specialist parasitoids fruit tests Allorhogas sp. Larvae gall fruit, Need species described, Dependence on developing interfering with seed rearing methods, and fruit may limit populations development fruiting trees for seasonally specificity tests Coccodiella Fungus causes severe Conditions for infecting Requires conditions of high miconiae rust-like disease on plants in lab are very humidity; may be limited young leaves specific and demanding to seasonally or in drier areas execute Ditylenchus Nematodes gall new Need better understanding Galling on miconia may be gallaeformans growth, both of mechanisms for slight under field conditions vegetative and infection and dispersal (unlike heavy galling on reproductive )

The weevil Cryptorhynchus melastomae emerges as a clear favorite among stem-feeding species so far discovered (Table 1). Its long-lived adults cause extensive damage with their own feeding, and a boring larva can kill a young stem as it feeds inside (Reichert 2007). This species is undergoing intensive testing in quarantine to evaluate the limits of its host range. It appears to utilize mainly melastomes closely related to Miconia calvescens. Potential for use of plants outside this family must be carefully examined. If this agent is

2009 International Miconia Conference Johnson • 5 found suitable for release, its apparent preference for young stems may contribute to suppression of pre-reproductive trees in a way that complements current aerial herbicide treatments of mature miconia.

Among leaf-feeding insects, two closely related Euselasia species appear to have the most promise, although rearing these butterflies through their entire life cycle in captivity remains a significant challenge (Allen 2007, Nishida 2010). By comparison, the damage caused by the sawfly Atomacera petroa seems less likely to contribute significantly to miconia management, since their feeding is restricted to fully mature leaves which are already under some suppression by the fungus C. gloeosporioides (Badenes-Perez and Johnson 2007b). Although lepidopteran, Euselasia caterpillars could experience relatively low levels of predation and parasitism in Hawaiÿi because of their group defensive behavior and because riodinids appear to share few parasitoids with other Lepidoptera. As is the case for almost all natural enemies of miconia, populations of these caterpillars would have to increase far beyond the levels typically seen in their native range to have meaningful impacts on miconia in the Pacific.

Insects that attack reproductive structures of a weed are unlikely to provide complete control by themselves, but they may contribute importantly to managing dispersal (Post et al. 2010). This is particularly true in the case of the tremendously prolific miconia, for which preventing spread beyond current infestations is a central goal of management. Four insect species have high potential for impacts on reproduction (Table 1). While all four are likely to have narrow host ranges, gaps in our knowledge of each species make it difficult to assess which of them has the greatest probability of lowering reproduction of miconia. The undescribed Mompha sp. is attractive for its ability to attack flower buds as well as developing fruit (Alfaro-Alpizar unpub. data). In this regard, it could have greater impact than its relative Mompha trithalama, which is already established in Hawaiÿi for clidemia biocontrol but only destroys seeds in developing fruit. Of the fruit-feeding weevils, we have more information about Anthonomus monostigma (Chacón 2007), and this species would likely be easier logistically than Apion sp. to develop further. The Allorhogas sp. is the least understood of all, but as a gall former, it may have the greatest potential for diverting miconia’s resources away from seed production (Badenes-Perez and Johnson 2007a). Research in the near future will attempt to describe the biology of Allorhogas in greater detail. Meanwhile we must overcome the technical challenge of rearing any of these species. Miconia inflorescences cut from a decompose in a few days, so carrying captive insects through their complete life cycle will require work with mature trees in a controlled environment.

The two pathogens both merit full evaluation and are discussed in detail elsewhere in these proceedings. Severe impacts of the galling nematode on some melastomes make it an impressive candidate, although it is quite rare on Miconia calvescens and much more damaging to Clidemia species in its native range. Because Coccodiella miconiae can attack young leaves and produce extensive damage, its impacts might exceed those of the fungus Colletotrichum gloeosporioides, particularly if it is introduced without hyperparasites. There is little doubt that both these pathogens will be found sufficiently host-specific for release in Hawaiÿi, and Coccodiella miconiae is likely so tightly coevolved that it would be restricted to M. calvescens throughout the invaded Pacific. Use of the nematode for biocontrol outside of Hawaiÿi will require assessment of risk to native melastomes which are found in some Pacific Islands.

2009 International Miconia Conference Johnson • 6 Areas of Uncertainty

While appropriate pre-release tests can determine host specificity of weed biocontrol agents with a high degree of confidence (Pemberton 2000, Sheppard et al. 2005), predicting impacts of agents on a weed target remains a major challenge (McClay and Balciunas 2005). Factors such as climate, weed distribution and phenology, plant defenses, natural enemies and competitors of agents, plant community interactions, and management decisions can affect the outcome of a particular biocontrol introduction in ways that may vary in time and space. Although perfect prediction of such complex interactions will probably always lie beyond the capabilities of science, it does seem reasonable to try to anticipate and avoid known barriers to successful biocontrol. Evaluating the potential for interference by natural enemies of miconia agents is one example of pre-release assessment appropriate to our efforts in Hawaiÿi. Other factors likely to control population dynamics of each agent in Pacific Island environments are important to consider case by case. We want to select agents with high intrinsic growth rates, well adapted to local climate and host plant conditions, and unlikely to compete against each other. Avoiding competition might be a reason to select only the most promising agent from each guild (stem, leaf, flower/fruit feeders), and may be important to consider when evaluating the fruit-feeding insects.

Adding to the complexity of forecasting impacts is the likelihood that some of our potential agents have a host range extending beyond miconia to other melastomes. Release of agents against miconia outside of Hawaiÿi will require careful consideration of potential non-target effects on native species. For example, in the Society Islands and New Caledonia there are a few endemic melastome species for which possible impacts of biocontrol agents might need to be avoided or mitigated. This is not a problem in Hawaiÿi in terms of non-target effects because all melastomes are alien and generally weedy here. However, use of other melastome hosts could mean that dynamics of agent populations would not be tightly linked with miconia, but rather their populations and impacts might be driven mainly by interactions with other common melastomes. Multi-melastome host use so far seems likely for the stem borer Cryptorhynchus melastomae, for which Melastoma candidum (M. septemnervium), among others, may be a significant alternate host (Johnson unpub. data). Ideally, the agent might contribute to management of multiple weed species, but the actual consequences of such interactions are difficult to predict. For such agents it will be important to monitor impacts on multiple targets, and this will add to the effort required in post-release evaluations.

Dispersal ability of biocontrol agents is another critical determinant of their impacts on target weeds (e.g. Pratt et al. 2003). In Hawaiÿi where active management focuses on containing the spread of miconia, agents ideally might contribute to this task by rapidly colonizing new miconia plants, even when they occur far from established populations. Actual dispersal rates of agents are not likely to be known in advance of release, although they can be estimated from what is known for similar species. The dispersal dynamics of both agents and miconia will determine how biocontrol and other management tools can be integrated. If agents are relatively slow spreading, then we may have to decide between redistribution of biocontrols versus mechanical/herbicidal treatments of incipient miconia populations.

The challenge of integrating biocontrol and other management tools is especially significant in the case of miconia. In the short term, simply establishing biocontrol agents and then assessing their populations may be difficult, since there are relatively few areas in Hawaiÿi where miconia is not under some level of periodic management. If biocontrol agents can be released and their effectiveness measured, then we face a transition from almost complete reliance on mechanical/herbicidal methods to partial reliance on 2009 International Miconia Conference Johnson • 7 biocontrol for management objectives. The ultimate balance between use of biocontrol and other tools will depend on how effective biocontrol can be, what resources are available for other control methods, and what our long-term objectives for miconia management are. These are questions of substantial complexity, but amenable to modeling with the right combination of expertise. Modeling miconia invasion dynamics and control tactics has been underutilized as a tool in Hawaiÿi. Deliberate investment in this approach could greatly facilitate management planning and integration of biocontrol with other control methods.

Goals for Development of Biocontrol

Setting explicit goals for biocontrol of miconia is the best way to chart a future course that can be periodically evaluated and changed as necessary. We currently have a good selection of potential agents and some information with which to prioritize their development. Goals over the next several years include:  Develop a suite of agents attacking stems, leaves and fruits of miconia, selecting the most damaging among available host-specific species. At present these include the stem borer Cryptorhynchus, one of two species of leaf-feeding butterflies (Euselasia), one or more of four flower/fruit-feeding insects, and two pathogens that infect leaves and new growth.  Anticipate and avoid agents with critical vulnerability to natural enemies present in Hawaiÿi and avoid competition or interference among agents.  Understand post-release population dynamics of agents by monitoring dispersal and host use patterns in the field. Post-release monitoring is essential for informing managers and the public about the utility and limits of biocontrol.  Develop spatial-temporal modeling of miconia dispersal and control methods as a decision-making tool to integrate biocontrol with existing management.

The pace of future progress toward biocontrol of miconia is mainly dependent on funding. Although in Hawaiÿi we have minimally adequate facilities and expertise to pursue the goal of developing several miconia agents, we are well below critical mass for accomplishing this goal rapidly and comprehensively. Existing programs funded modestly ($100-200K per year) might be expected to result in release of 4-5 agents over the next decade, with a minimal commitment to post-release monitoring and little or no effort toward use of modeling or other decision tools. On the other hand, a more intensive effort employing additional researchers would cost more (perhaps $400-500K per year) but accomplish additional goals in less time. Scaling up to include expertise from colleagues outside Hawaiÿi, for example from Australia, probably is the most effective way we can improve on current efforts.

Acknowledgements

Cliff Smith, with colleagues at the University of Hawaiÿi Pacific Cooperative Studies Unit, initiated and oversaw collaboration between Hawaiÿi and key researchers abroad for many years. Without his energy and vision, miconia biocontrol might have stalled long ago. Professors Robert Barreto and Marcelo Picanço at the Universidade Federal de Viçosa, Brazil, and Paul Hanson at the Universidad de Costa Rica and their many colleagues have committed lab space and years mentoring students interested in miconia biocontrol research. The students have included Eduardo Chacón, Kenji Nishida, Pablo Allen, Luis Madrigal, Manuel Alfaro-Alpizar, Alex Castillo, Beth Reichert, Mayrelith Artavia and Ana Araya in Costa Rica; and Altair Semeão, Elisangela Morais, Emerson de Barros, Gerson Silva, Jander Rosado, Shaiene Moreno and Claudine Seixas in Brazil. Edgar Rojas and

2009 International Miconia Conference Johnson • 8 others helped direct the very productive program in Costa Rica. Francisco Badenes-Perez and Anna Dietrich bridged the projects in Brazil and Costa Rica and helped evaluate several potential new agents. Alec McClay discovered the galling nematode under our noses and gave us a first look at miconia in Mexico. Fellow biocontrol researchers at the Hawaiÿi Department of Agriculture, especially Bob Burkhart, Eloise Killgore, Mohsen Ramadan, Marianne Chun, Ken Murai, Pat Conant, Lionel Sugiyama, Walter Nagamine and Bernarr Kumashiro, collected and processed our earliest records of miconia enemies. Wendell Sato, Fran Calvert, Brenner Wai, Erin Raboin and Sam Brooks have conducted initial quarantine evaluations for the USDA Forest Service. Julie Denslow and other colleagues in Hawaiÿi have supported our Forest Service biocontrol program in many critical ways. Financial support has been provided by the government of French Polynesia, USGS Pacific Island Ecosystems Research Center, National Park Service, Hawaiÿi Department of Agriculture, Hawaiÿi Council, Forest Service International Programs, and Forest Service Special Technology Development Program.

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