A New Beetle-fungus Disease Complex Threatening Avocado
California Avocado Society 2011 Yearbook 94:120-126Akif EskalenDepartment of Plant Pathology and MicrobiologyRichard StouthamerDepartment of EntomologyUniversity of CaliforniaRiverside, CaliforniaA new beetle-fungus diseasecomplex threatening avocadoRecently a new beetle/disease complex was detected thatcauses a Fusarium dieback on avocado and other host plantsin and near Los Angeles County. The disease is caused by a new, yet unnamed Fusarium sp. that forms a symbiotic relationship with a recentlydiscovered Euwallacea sp. beetle, which serves as the vector. This beetle ismorphologically indistinguishable from the tea shot hole borer, Euwallacea fornicatus (an exotic Asianambrosia beetle). We suspect thatthis is a new species (for which wepropose the common name: Polyphagous Shot Hole Borer) basedon the large differences in DNAsequence between the beetle invading California, and beetlesfrom tea plantations in Sri Lankaand other Asian collection sites.The beetle discovered in CaliforFigure 1: An avocado tree in Israel exhib- nia is smaller than a sesame seediting symptoms of beetle/disease (Mendel(about 0.1 inch in length). Theet al. 2012)120identical new beetle species wasfound in Israel in 2009 in commercial avocado orchards where it hasbeen causing damage to avocado(Fig. 1,2).Unlike the Redbay Ambrosia beetle, the vector of the Laurelwilt disease which has been infesting avocado and other membersof the avocado family (Laureceae)in the Southeastern United States, Figure. 2: A failed main branch of avothe new beetle has been observed cado in Israel caused by the Fusarium dieon more than 100 different plant back (Mendel et al. 2012)species in California including many species common in the urban landscape and on such agriculturally important species as: avocado, olive,peach, guava, lychee, mango, persimmon, and pomegranate.The beetle and fungus have a symbiotic relationship. When thebeetle burrows into the tree, it inoculates the host plant with the fungus,which is carried in its mouthparts in a structure called a mycangia. Thefungus attacks the vascular tissue of the tree which brings water andnutrients from the roots to the rest of the tree eventually causing branchdieback. The beetle larvae live in galleries within the tree and feed on thefungus.The California Avocado Commission provided emergency funding to launch a research project since the beetle/fungus complex couldcause serious economic damage to the California avocado industry ifintroduced into commercial plantings. A team of University of California, Riverside (UCR) researchers are collaborating as they study thisproblem.Dr. Akif Eskalen and field specialist Alex Gonzalez from the Department of Plant Pathology and Microbiology are conducting a surveyin Southern California to determine the extent of the beetle infestationand the likely extent of Fusarium sp. infection in avocado trees and other host plants. They are also investigating Fusarium dieback incidencewithin the commercial avocado groves and landscape trees. Dr. RichardStouthamer, Paul Rugman-Jones and Tim Paine from the Departmentof Entomology are studying the biology and genetics of the beetle andare exploring insecticides that may be used to kill the beetle on the plant121
Akif EskalenDepartment of Plant Pathology and MicrobiologyRichard StouthamerDepartment of EntomologyUniversity of CaliforniaRiverside, CaliforniaA new beetle-fungus diseasecomplex threatening avocadoRecently a new beetle/disease complex was detected thatcauses a Fusarium dieback on avocado and other host plantsin and near Los Angeles County. The disease is caused by a new, yet unnamed Fusarium sp. that forms a symbiotic relationship with a recentlydiscovered Euwallacea sp. beetle, which serves as the vector. This beetle ismorphologically indistinguishable from the tea shot hole borer, Euwallacea fornicatus (an exotic Asianambrosia beetle). We suspect thatthis is a new species (for which wepropose the common name: Polyphagous Shot Hole Borer) basedon the large differences in DNAsequence between the beetle invading California, and beetlesfrom tea plantations in Sri Lankaand other Asian collection sites.The beetle discovered in CaliforFigure 1: An avocado tree in Israel exhib- nia is smaller than a sesame seediting symptoms of beetle/disease (Mendel(about 0.1 inch in length). Theet al. 2012)120identical new beetle species wasfound in Israel in 2009 in commercial avocado orchards where it hasbeen causing damage to avocado(Fig. 1,2).Unlike the Redbay Ambrosia beetle, the vector of the Laurelwilt disease which has been infesting avocado and other membersof the avocado family (Laureceae)in the Southeastern United States, Figure. 2: A failed main branch of avothe new beetle has been observed cado in Israel caused by the Fusarium dieon more than 100 different plant back (Mendel et al. 2012)species in California including many species common in the urban landscape and on such agriculturally important species as: avocado, olive,peach, guava, lychee, mango, persimmon, and pomegranate.The beetle and fungus have a symbiotic relationship. When thebeetle burrows into the tree, it inoculates the host plant with the fungus,which is carried in its mouthparts in a structure called a mycangia. Thefungus attacks the vascular tissue of the tree which brings water andnutrients from the roots to the rest of the tree eventually causing branchdieback. The beetle larvae live in galleries within the tree and feed on thefungus.The California Avocado Commission provided emergency funding to launch a research project since the beetle/fungus complex couldcause serious economic damage to the California avocado industry ifintroduced into commercial plantings. A team of University of California, Riverside (UCR) researchers are collaborating as they study thisproblem.Dr. Akif Eskalen and field specialist Alex Gonzalez from the Department of Plant Pathology and Microbiology are conducting a surveyin Southern California to determine the extent of the beetle infestationand the likely extent of Fusarium sp. infection in avocado trees and other host plants. They are also investigating Fusarium dieback incidencewithin the commercial avocado groves and landscape trees. Dr. RichardStouthamer, Paul Rugman-Jones and Tim Paine from the Departmentof Entomology are studying the biology and genetics of the beetle andare exploring insecticides that may be used to kill the beetle on the plant121
surface before they enter the tree. Reuben Hofshi, of the Hofshi Foundation, is also gathering information about the beetle/disease internationally, and has been instrumental in obtaining beetle specimens frominternational cooperators.According to survey data through July 2012, the easternmostlimit appears to be the 57 Freeway with everywhere west of it in LACounty infested. East of the 57 Freeway appears to be free of detections,thus far. South of the 57 Freeway is also infested, particularly areas inthe Orange County areas of Fullerton, Anaheim, and down to LagunaBeach. In between those cites, the Irvine area appears to be free of thebeetle. (Fig. 3).Figure 4: White ring of sugar exudatesurrounds the beetle entry/exit hole(A.Eskalen)Figure 5: Limb damage showingsugar exudate (A.Eskalen)Figure 6: A singe entry/exit hole covered Figure 7: Multiple entry/exit holes ofby sugar exudate. (A.Eskalen)beetle on an avocado branch (cv. Hass)(A.Eskalen)Figure 3: Map of complex distributionFusarium dieback symptomsSymptoms of the disease on avocado include a white powderysugar exudate that can be either dry, or surrounded by wet discolorationof the outer bark. (Fig. 4,5) This symptom can be in association with asingle or multiple beetle entry/exit holes (Fig. 6,7). While there is novisible injury to the bark at this early stage of colonization, examinationof the cortex and wood under the infested entry/exit hole bored by thebeetle, reveals brown discolored necrosis caused by the fungus (Fig. 8,9).122Figure 8: An entry/exit hole of the beetle Figure 9: Typical Fusarium dieback sympon Avocado bark (cv. Bacon) (A.Eskalen) toms, scraping the bark down to the woodbehind the beetle entry/exit hole revealsbrown necrotic tissue. (A.Eskalen).123
surface before they enter the tree. Reuben Hofshi, of the Hofshi Foundation, is also gathering information about the beetle/disease internationally, and has been instrumental in obtaining beetle specimens frominternational cooperators.According to survey data through July 2012, the easternmostlimit appears to be the 57 Freeway with everywhere west of it in LACounty infested. East of the 57 Freeway appears to be free of detections,thus far. South of the 57 Freeway is also infested, particularly areas inthe Orange County areas of Fullerton, Anaheim, and down to LagunaBeach. In between those cites, the Irvine area appears to be free of thebeetle. (Fig. 3).Figure 4: White ring of sugar exudatesurrounds the beetle entry/exit hole(A.Eskalen)Figure 5: Limb damage showingsugar exudate (A.Eskalen)Figure 6: A singe entry/exit hole covered Figure 7: Multiple entry/exit holes ofby sugar exudate. (A.Eskalen)beetle on an avocado branch (cv. Hass)(A.Eskalen)Figure 3: Map of complex distributionFusarium dieback symptomsSymptoms of the disease on avocado include a white powderysugar exudate that can be either dry, or surrounded by wet discolorationof the outer bark. (Fig. 4,5) This symptom can be in association with asingle or multiple beetle entry/exit holes (Fig. 6,7). While there is novisible injury to the bark at this early stage of colonization, examinationof the cortex and wood under the infested entry/exit hole bored by thebeetle, reveals brown discolored necrosis caused by the fungus (Fig. 8,9).122Figure 8: An entry/exit hole of the beetle Figure 9: Typical Fusarium dieback sympon Avocado bark (cv. Bacon) (A.Eskalen) toms, scraping the bark down to the woodbehind the beetle entry/exit hole revealsbrown necrotic tissue. (A.Eskalen).123
The vector beetleThe beetle is a new Euwallacea species relative of the tea shothole borer (an exotic Asian ambrosia beetle). It is very small and difficultto see (Fig. 10). The beetle holes penetrate approximately 0.4-1.57 inchinto the wood and there are often many entry/exit holes on an infestedtree. The exit hole on avocado is about 0.033 inch wide (Fig. 8). Females are black colored and about 0.07-0.1 inch long (Fig. 11), males arebrown colored and about 0.05 inch long (Fig. 12).Figure 10: Male (left) and female (right)beetles on a penny. (A. Eskalen).Figure 11: Female beetle Euwallacea sp., Figure 12: Male beetle Euwallacea sp.,0.07 to 0.1 inch long and black. (G. Ara- 0.05 inch long and brown. (G. Arakelian).kelian).(Platanus racemosa), Big leaf maple (Acer macrophyllum), California BayLaurel (Umbellularia californica), White alder (Alnus rhombifolia), Olive(Olea europaea), Peach (Prunus persica), Persimmon (Diospyros sp.), Mimosa (Albizia julibrissin), Liquid amber (Liquidambar styraciflua), Wisteria vine (Wisteria sinensis).What to do: Look for a single entry/exit holes with surrounding white powdery exudate on avocado. The symptoms may be different on different hosts such as different levels of staining and/or gumming. Scrape off the bark layer around the infected area to look fornecrotic tissue. Follow the gallery to look for the discolored necrosis caused bythe fungus. Certain plant species appear to be a preferred host for the beetlesuch as box elder, castor bean, English oak, sycamore, liquidambar, goldenrain tree and coast live oak. If such plants are present close to avocado groves, inspect them for evidence of thepresence or damage by the beetles. The highest beetle exit/entryholes density is found at the base of the plant. If no beetle holesare present, it may be a good idea to remove the plants1. If beetleholes are found we do not yet have a proven way in which theplants can be destroyed without allowing the beetles to disperse.Who to contact if you find the problem:If you suspect that you have found this beetle or seen symptomsof the Fusarium dieback in your grove or surrounding areas in other hostplants please contact either your local farm advisor, pest control advisor,county Ag Commissioner office, or the CAC office at 949-341-1955.You can also contact Dr. Akif Eskalen at the University of California,Riverside by either phone 951-827-3499, or email at akif.eskalen@ucr.edu for confirmation of the pathogen.Known HostsBox elder (Acer negundo), Castor bean (Ricinus communis), Avocado (Persea americana), Black locust (Robinia pseudoacacia), Englishoak (Quescus robur), Goldenrain (Koelreuteriae elegans), Coast live oak(Quercus agrifolia), Engelmann oak (Quescus engelmannii), Sycamore1 Editor Note: Please check with local authorities prior to removing protected species.124125
The vector beetleThe beetle is a new Euwallacea species relative of the tea shothole borer (an exotic Asian ambrosia beetle). It is very small and difficultto see (Fig. 10). The beetle holes penetrate approximately 0.4-1.57 inchinto the wood and there are often many entry/exit holes on an infestedtree. The exit hole on avocado is about 0.033 inch wide (Fig. 8). Females are black colored and about 0.07-0.1 inch long (Fig. 11), males arebrown colored and about 0.05 inch long (Fig. 12).Figure 10: Male (left) and female (right)beetles on a penny. (A. Eskalen).Figure 11: Female beetle Euwallacea sp., Figure 12: Male beetle Euwallacea sp.,0.07 to 0.1 inch long and black. (G. Ara- 0.05 inch long and brown. (G. Arakelian).kelian).(Platanus racemosa), Big leaf maple (Acer macrophyllum), California BayLaurel (Umbellularia californica), White alder (Alnus rhombifolia), Olive(Olea europaea), Peach (Prunus persica), Persimmon (Diospyros sp.), Mimosa (Albizia julibrissin), Liquid amber (Liquidambar styraciflua), Wisteria vine (Wisteria sinensis).What to do: Look for a single entry/exit holes with surrounding white powdery exudate on avocado. The symptoms may be different on different hosts such as different levels of staining and/or gumming. Scrape off the bark layer around the infected area to look fornecrotic tissue. Follow the gallery to look for the discolored necrosis caused bythe fungus. Certain plant species appear to be a preferred host for the beetlesuch as box elder, castor bean, English oak, sycamore, liquidambar, goldenrain tree and coast live oak. If such plants are present close to avocado groves, inspect them for evidence of thepresence or damage by the beetles. The highest beetle exit/entryholes density is found at the base of the plant. If no beetle holesare present, it may be a good idea to remove the plants1. If beetleholes are found we do not yet have a proven way in which theplants can be destroyed without allowing the beetles to disperse.Who to contact if you find the problem:If you suspect that you have found this beetle or seen symptomsof the Fusarium dieback in your grove or surrounding areas in other hostplants please contact either your local farm advisor, pest control advisor,county Ag Commissioner office, or the CAC office at 949-341-1955.You can also contact Dr. Akif Eskalen at the University of California,Riverside by either phone 951-827-3499, or email at akif.eskalen@ucr.edu for confirmation of the pathogen.Known HostsBox elder (Acer negundo), Castor bean (Ricinus communis), Avocado (Persea americana), Black locust (Robinia pseudoacacia), Englishoak (Quescus robur), Goldenrain (Koelreuteriae elegans), Coast live oak(Quercus agrifolia), Engelmann oak (Quescus engelmannii), Sycamore1 Editor Note: Please check with local authorities prior to removing protected species.124125
References:Arakelian, G. 2012. Tea Shot Hole Borer (Euwallacea fornicatus). County of Los Angeles, Department of Agricultural Commissioner/Weights and Measures.Danthanarayana, W. 1968. The distribution and host-range of the shothole borer (Xyleborus fornicatusEichh.) of tea. Tea Quarterly,39, 61–69.Eskalen, A., Gonzalez, A., Wang, H.W., Twizeyimana, M., Mayorquin,J.S., Lynch, S.C. 2012. First report of Fusarium sp. and its vector tea shot hole borer (Euwallacea fornicatus) causing Fusariumdieback on avocado in California. Plant Disease. n, A. and R. Stouthamer. 2012. Pest Alert: Fusarium dieback onCalifornia avocado trees vectored by Tea Shot Hole Borer (Euwallacea fornicatus. University of California, Riverside and California Avocado Commission (http://eskalenlab.ucr.edu/handouts/fusarium dieback english.pdf ).Mendel, Z., Protosav, A., Sharon,M., Zveibil, A., Ben Yahuda, S.,O’Donnell, K., Rabaglia,R., Wysoki,M., Freeman,S. 2012. AnAsian ambrosia beetle Euwallacea fornicatus and its novel symbiotic fungus Fusarium sp. pose a serous threat to Israeli avocadoindustry. Phytoparasitica, DOI 10.1007/s12600-012-0223-7O’Donnell, K., Sutton, D. A., Fothergill, A., McCarthy, D., Rinaldi, M.G., Brandt, M. E., et al. 2008. Molecular phylogenetic diversity,multilocus haplotype nomenclature, and in vitroantifungal resistance within the Fusarium solani species complex.Journal ofClinical Microbiology, 46, 2477–2490.Rabaglia, R. J., Dole, S. A., & Cognato, A. I. 2006. Review of AmericanXyleborina (Coleoptera: Curculionidae: Scolytinae) occurringnorth of Mexico, with an illustrated key. Annals of the Entomological Society of America, 99, 1034–1055.126Zvi Mendel1Stanley Freeman2Depts. of Entomology1, andPlant Pathology and Weed Research2Agricultural Research Organization,The Volcani Center, Bet Dagan 50250, IsraelThe current situation of‘ambrosia wilt’ in avocado plantationsin Israel caused by the shot hole borerand its fungal symbiont FusariumAvocado in Israel is a growing industry. The plantations coveran area of approximately 7,000 ha with ‘Hass’, ‘Ettinger’and ‘Pinkerton’ comprising the main varieties. During the 2010/11 season about 85,000 tons were produced (from a bearing area of 5,500 ha),with approximately 65% of the crop exported. For decades the use offrequent, prophylactic or responsive sprays has been avoided in avocadocultivation in Israel. However, the situation is now requiring the growersto consider the use of integrated pest management (IPM) schemes dueto the establishment of the shot hole borer.Host plant and symptom of injury of the shot hole borerThe shot hole borer, an ambrosia beetle, Euwallacea sp. near fornicatus, (Scolytinae) was first recorded in Israel in 2009 in an avocadoplantation in Glil Yam (central coastal area). Soon after the beetle and itsfungal symbiont Fusarium sp. nov. were found in other avocado plantations in that area and also in three other plant hosts, castor bean (Ricinuscommunis, a weed), box elder (Acer negundo) and Quercus pedunculifolia(Mendel et al. 2012). The latter 2 tree species are exotic ornamentaltrees. None of these four tree species are among the original hosts of the
identical new beetle species was found in Israel in 2009 in commer-cial avocado orchards where it has been causing damage to avocado (Fig 1,2) Unlike the Redbay Ambro-sia beetle, the vector of the Laurel wilt disease which has been infest-ing avocado and other members of the avocado family (Laureceae) in the Southeastern United States,
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