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Forest pathology in Hawaii

Native Hawaiian forests are characterised by a high degree of endemism, including pathogens as well as their hosts. With the exceptions of koa (Acacia koa Gray), possibly maile (Alyxia oliviformis Gaud.), and, in the past, sandalwood (Santalum spp.), forest species are of little commercial value. On the other hand, these forests are immensely important from a cultural, ecological, and evolutionary standpoint. Forest disease research was lacking during the mid-twentieth century, but increased markedly with the recognition of ohia (Metrosideros polymorpha Gaud.) decline in the 1970s. Because many pathogens are themselves endemic, or are assumed to be, having evolved with their hosts, research emphasis in natural areas is on understanding host-parasite interactions and evolutionary influences, rather than disease control. Aside from management of native forests, attempts at establishing a commercial forest industry have included importation of several species of pine, Araucaria, and Eucalyptus as timber crops, and of numerous ornamentals. Diseases of these species have been introduced with their hosts. The attacking of native species by introduced pathogens is problematic - for example, Armillaria mellea (Vahl ex Fr.) Que??l. on koa and mamane (Sophora chrysophylla (Salisb.) Seem.). Much work remains to be done in both native and commercial aspects of Hawaiian forest pathology.

New Zealand Journal of Forestry Science

Ambrosia beetles (Coleoptera: Curculionidae) can directly transmit the fungal pathogens responsible for Rapid ʻŌhiʻa Death

The ecologically and culturally vital tree species, ʻōhiʻa lehua ( Metrosideros polymorpha ), is threatened by the fungal pathogens Ceratocystis lukuohia and Ceratocystis huliohia , the causal agents of the disease complex called Rapid ʻŌhiʻa Death (ROD). Four invasive ambrosia beetle (Coleoptera: Curculionidae: Scolytinae) species in the Xyleborini tribe colonize ROD Ceratocystis -infested ‘ōhiʻa and produce inoculum through their frass; however, the potential for direct transmission of the ROD fungi by these beetles was unknown. We fulfilled Leach's rules to support insect transmission of ROD by documenting the visitation of these ambrosia beetles to healthy ‘ōhiʻa trees, culturing C. lukuohia and C. huliohia from the ROD-associated beetles using three different collection methods at multiple study sites, and challenging healthy ʻōhiʻa seedlings with beetles that were exposed to both C. lukuohia and C. huliohia cultures. We documented all four invasive ROD-associated ambrosia beetle species including Xyleborinus saxesenii , Xyleborus affinis , Xyleborus ferrugineus , and Xyleborus perforans to regularly visit healthy ʻōhiʻa trees on sticky traps. Viable Ceratocystis propagules were isolated from all species, and C. lukuohia was most commonly isolated of the two ROD-causing fungi. Consistently across all collection techniques, ROD Ceratocystis spp. were detected on just under 3% of all assayed beetles, with the highest detection rate from X. affinis . All four beetle species were capable of directly transmitting both pathogens to healthy ʻōhiʻa seedlings with a high rate of transfer. Ceratocystis spp. are highly virulent pathogens in trees, and a single inoculation can result in tree death, therefore any direct transmission is a cause for concern. After meeting the criteria of Leach's rules, we propose that Xi. saxesenii , X. affinis , X. ferrugineus , and X. perforans are vectors of C. lukuohia and C. huliohia , particularly in areas of high ROD pressure and tree stress.

Hawaii

Wildfire risk as a socioecological pathology

Wildfire risk in temperate forests has become a nearly intractable problem that can be characterized as a socioecological “pathology”: that is, a set of complex and problematic interactions among social and ecological systems across multiple spatial and temporal scales. Assessments of wildfire risk could benefit from recognizing and accounting for these interactions in terms of socioecological systems, also known as coupled natural and human systems (CNHS). We characterize the primary social and ecological dimensions of the wildfire risk pathology, paying particular attention to the governance system around wildfire risk, and suggest strategies to mitigate the pathology through innovative planning approaches, analytical tools, and policies. We caution that even with a clear understanding of the problem and possible solutions, the system by which human actors govern fire-prone forests may evolve incrementally in imperfect ways and can be expected to resist change even as we learn better ways to manage CNHS.

Frontiers in Ecology and the Environment

Pathology of tissue loss in three key gorgonian species in the Mediterranean Sea

The Mediterranean is known for its marine biodiversity, especially gorgonian forests. Unfortunately, these are experiencing rapid declines due to climate change, manifested by repeated marine heat waves resulting in mass mortality events since the early 1990 s. To better understand why gorgonians are declining, more systematic approaches to investigate the exact causes are needed, and pathology may aid in this goal. We described gross and microscopic pathology of tissue loss in three key gorgonian species in the Mediterranean region, Paramuricea clavata , Eunicella cavolini , and Leptogorgia sarmentosa , that were all experiencing various degrees of acute to subacute tissue loss characterized by exposed axial skeleton sometimes partly colonized by epibionts and thinning of adjacent tissues. The most significant variety of lesions was seen in P. clavata followed by L. sarmentosa and E. cavolini. For all species, dissociation of gastrodermal cells was the dominant microscopic lesion followed by necrosis of the gastrodermis. Ciliates invading gastrodermis and associated with necrosis of polyps were seen only in E. cavolini. Epidermal tissue loss was seen only in L. sarmentosa , while P. clavata was distinguished by a prominent inflammatory response and unidentified dark round structures within the tentacle epidermis and gastrodermis with no host response. Further work to understand the cause of death in gorgonians is needed, particularly to elucidate the role of ciliates and environmental co-factors or infectious agents not visible on light microscopy, as well as applications of additional tools such as cytology.

Journal of Invertebrate Pathology

Why do trees die? Characterizing the drivers of background tree mortality

The drivers of background tree mortality rates—the typical low rates of tree mortality found in forests in the absence of acute stresses like drought—are central to our understanding of forest dynamics, the effects of ongoing environmental changes on forests, and the causes and consequences of geographical gradients in the nature and strength of biotic interactions. To shed light on factors contributing to background tree mortality, we analyzed detailed pathological data from 200,668 tree-years of observation and 3,729 individual tree deaths, recorded over a 13-yr period in a network of old-growth forest plots in California's Sierra Nevada mountain range. We found that: (1) Biotic mortality factors (mostly insects and pathogens) dominated (58%), particularly in larger trees (86%). Bark beetles were the most prevalent (40%), even though there were no outbreaks during the study period; in contrast, the contribution of defoliators was negligible. (2) Relative occurrences of broad classes of mortality factors (biotic, 58%; suppression, 51%; and mechanical, 25%) are similar among tree taxa, but may vary with tree size and growth rate. (3) We found little evidence of distinct groups of mortality factors that predictably occur together on trees. Our results have at least three sets of implications. First, rather than being driven by abiotic factors such as lightning or windstorms, the “ambient” or “random” background mortality that many forest models presume to be independent of tree growth rate is instead dominated by biotic agents of tree mortality, with potentially critical implications for forecasting future mortality. Mechanistic models of background mortality, even for healthy, rapidly growing trees, must therefore include the insects and pathogens that kill trees. Second, the biotic agents of tree mortality, instead of occurring in a few predictable combinations, may generally act opportunistically and with a relatively large degree of independence from one another. Finally, beyond the current emphasis on folivory and leaf defenses, studies of broad-scale gradients in the nature and strength of biotic interactions should also include biotic attacks on, and defenses of, tree stems and roots.

Ecology

Dieback of Acacia koa in Hawaii: Ecological and pathological characteristics of affected stands

Koa (Acacia koa) is an endemic Hawaiian tree that serves as a keystone species in the upper elevation forests of all the main islands. In the Mauna Loa Strip area of Hawaii Volcanoes National Park, mature koa stands are suffering from an unexplained dieback that has increased in severity since it was first noticed approximately 25 years ago. The dieback is often evident in patches, and generally spreads within stands in a radial fashion from a localized infection center. Entire crowns of affected trees become wilted, with foliage gradually progressing from an apparent healthy to a completely chlorotic condition. Although most trees die soon after the onset of symptoms, some trees are able to survive crown death by producing epicormic shoots on the lower portions of the trunk. Previously published studies reported that a vascular wilt fungus (Fusarium oxysporum f. sp. koae) was associated with koa seeds and the rhizosphere of healthy and dieback-affected koa stands. The purpose of this study was to characterize the stand structure, soil conditions, and physiological condition of dieback-affected trees, and to assess the possible role of F. oxysporum f. sp. koae in the current dieback stands. This fungus was isolated from branches of symptomatic koa in dieback-affected stands and roots from healthy and dieback-affected stands. Possible differences in the pathogenicity and virulence of F. oxysporum f. sp. koae isolates obtained from the roots of healthy koa in unaffected stands and those from branches of dieback-affected koa were determined by greenhouse inoculations of koa seedlings. Healthy koa saplings in stands unaffected by dieback were also inoculated to determine if disease symptoms could be induced by inoculation of injured roots in the field. Both branch and root isolates were pathogenic; with the percent mortality of inoculated seedlings ranging from 30 to 60% for all isolates. Disease severity between branch and root isolates was not significantly different. Wilt symptoms were not observed in field inoculations of koa saplings after 8 months. Investigation of the dieback stand structure determined that the larger size classes (i.e., DBH > 3 cm) were all affected equally, with the smallest size class (i.e., 0-3 cm DBH) having the lowest percentage of dieback at each site. Soil from dieback stands had higher water content and was more acidic than that from neighboring unaffected stands. Phyllodes on epicormic shoots of dieback trees had a reduced area, mass, and mass to area ratio. Phyllodes on dieback trees also had reduced stomatal conductance and higher (less negative) leaf water potential. Results from the greenhouse inoculations demonstrated that F. oxysporum f. sp. koae is pathogenic to koa, but the pathogen's activity in the field may be influenced by predisposing factors such as temperature fluctuations, water availability, soil type, and interactions with other soil organisms. F. oxysporum f. sp. koae appears to play an important role in regulating koa stand dynamics, growth, and physiology under environmental and edaphic conditions favorable for disease development. ?? 2002 Elsevier Science B.V. All rights reserved.

Forest Ecology and Management

Ceratocystis lukuohia-infested ambrosia beetle frass as inoculum for Ceratocystis wilt of ʻōhiʻa (Metrosideros polymorpha)

Metrosideros polymorpha (‘ōhi‘a) trees in Hawaiʻi are dying from two distinct diseases, collectively referred to as rapid ‘ōhi‘a death (ROD), caused by Ceratocystis lukuohia and Ceratocystis huliohia . Boring dust (frass) released when ambrosia beetles attack and colonize infected trees has been suspected as a transmission source. We sampled ambrosia beetle frass from six locations on Hawaiʻi Island and screened samples for Ceratocystis DNA and fungal viability. Ceratocystis DNA was detected in 79% of frass samples and 61% were viable. To assess the infectivity of C . lukuohia -colonized frass, M . polymorpha seedlings were wound-inoculated with frass in growth chamber trials. Wilt incidence was 40% in the first trial and 15% in the second. Frass particles naturally infested with C . lukuohia were treated with a cytoplasmic stain and microscopically examined; thick-walled chlamydospores were found in all samples. The chlamydospores appeared to be derived from aleurioconidia. Ceratocystis survival in frass was assessed in a baiting experiment conducted under varying environmental regimes; viability decreased with increasing temperature, frass age and decreasing humidity. After 6 months, 90% of the samples exhibited viable C . lukuohia at the lowest temperatures. Results confirmed that C . lukuohia chlamydospores are the fungal inoculum within ambrosia beetle frass and can induce wilt when introduced to M . polymorpha wounds. Although ambrosia beetles or other insects may serve as vectors, the present work supports the hypothesis that Ceratocystis species may be dispersed in frass, which can inform ongoing efforts to manage ROD across Hawaiʻi's native forests.

Hawaii

Mermithid parasitism of Hawaiian Tetragnatha spiders in a fragmented landscape

Hawaiian Tetragnatha spiders inhabiting small forest fragments on the Big Island of Hawaii are parasitized by mermithid nematodes. This is the first report of mermithid nematodes infecting spiders in Hawaii, and an initial attempt to characterize this host–parasite interaction. Because immature mermithids were not morphologically identifiable, a molecular identification was performed. A phylogenetic analysis based on 18S small ribosomal subunit nuclear gene sequences suggested that Hawaiian spider mermithids are more closely related to a mainland presumptive Aranimemis species that infects spiders, than to an insect-infecting mermithid collected on Oahu, HI, or to Mermis nigrescens , also a parasite of insects. Measured infection prevalence was low (ranging from 0 to 4%) but differed significantly among forest fragments. Infection prevalence was associated significantly with fragment area, but not with spider density nor spider species richness. Results suggest that mermithid populations are sensitive to habitat fragmentation, but that changes in infection prevalence do not appear to affect spider community structure.

Hawai'i

Potential causes for amphibian declines in Puerto Rico

We monitored 11 populations of eight species of Eleutherodactylus in Puerto Rico from 1989 through 2001. We determined relative abundance of active frogs along transects established in the Caribbean National Forest (El Yunque), Carite Forest, San Lorenzo, and in the vicinity of San Juan. Three species (Eleutherodactylus karlschmidti, E. jasperi, and E. eneidae) are presumed to be extinct and eight populations of six different species of endemic Eleutherodactylus are significantly declining at elevations above 400 m. Of the many suspected causes of amphibian declines around the world, we focused on climate change and disease. Temperature and precipitation data from 1970a??2000 were analyzed to determine the general pattern of oscillations and deviations that could be correlated with amphibian declines. We examined a total of 106 tissues taken from museum specimens collected from 1961a??1978 and from live frogs in 2000. We found chytrid fungi in two species collected at El Yunque as early as 1976, this is the first report of chytrid fungus in the Caribbean. Analysis of weather data indicates a significant warming trend and an association between years with extended periods of drought and the decline of amphibians in Puerto Rico. The 1970's and 1990's, which represent the periods of amphibian extirpations and declines, were significantly drier than average. We suggest a possible synergistic interaction between drought and the pathological effect of the chytrid fungus on amphibian populations.

Puerto Rico