Ambrosia beetles pests of Avocado

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Ambrosia beetles pests of Avocado Daniel Carrillo & Marc Hughes Ploetz, Crane (TREC) Cave (IRREC) Stelinski (CREC) Kendra, (USDA-ARS) Cooperband (USDA-APHIS)- Tropical Research and Education Center

Ambrosia beetles are fungal farmers Mycangia specialized saclike organ selectively maintain and transport fungi during dispersal

Red Bay Ambrosia Beetle, Xyleborus glabratus, primary vector of the laurel wilt disease in natural forests Tropical Research and Education Center

Causal agent : Raffaelea lauricola Moves systematically within the host (Lauraceae) and causes vascular wilt

Tylose formation: outgrowths on cells of xylem vessels Control (mock inoculated) Xylem dysfunction Inch, S.A.

Distribution of Counties with Laurel Wilt Disease* by year of Initial Detection * Laurel Wilt Disease is a destructive disease of redbay (Persea borbonia), and other species within the laurel family (Lauraceae) caused by a vascular wilt fungus (Raffaelea lauricola) that is vectored by the redbay ambrosia beetle (Xyleborus glabratus).the pathogen has been confirmed through laboratory analysis of host samples collected in the counties highlighted. -90-80 Initial Detection of Xyleborus glabratus - May 2002 Port Wentworth, GA Information Provided By: Alabama Dana McReynolds Stones Dana.McReynolds@forestry.alabama.gov Arkansas Chandler Barton Chandler.Barton@arkansas.gov Florida Jeff Eickwort Jeffrey.Eickwort@freshfromflorida.com Georgia Chip Bates cbates@gfc.state.ga.us Louisiana Brent Cutrer mcutrer@ldaf.state.la.us Mississippi Gulf of Mexico John J. Riggins jriggins@entomology.msstate.edu North Carolina Rob Trickel rob.trickel@ncagr.gov South Carolina David Jenkins Djenkins@scfc.gov Texas Shane Harrington sharrington@tfs.tamu.edu 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 Native to Taiwan, Japan & South East Asia Updated: February 15, 2018 0 25 50 100 150 200 Miles

Xyleborus glabratus, Redbay Ambrosia Beetle (RAB) infected ~ 0.5 billion native lauraceous trees with R. lauricola in the southeastern U.S. Persea borbonia Red bay Persea humilis Silk bay Sassafras albidum Persea palustris Swamp bay

Redbay A. wilt in upper crown B. Complete wilt of canopy Photo: Hughes et al. 2015

Ambrosia Beetle Boring A. Frass toothpicks or tubes B. Accumulated frass at tree base Photo: Hughes et al. 2015

Avocado is not a good host for Xyleborus glabratus Avocado (Persea americana)

~44,000 lost to LW Pathogen spreading in the apparent absence of X. glabratus Alternative vectors?

Several species of AB can carry R. lauricola

species n= No. beetles carrying R. lauricola probability of a beetle carrying R. lauricola CFUs Mean ± SEM CFU Range Xyleborus glabratus 50 43 0.86 a 2783.3 ± 281.9 a 0-7800 Xyleborus affinis 41 5 0.12 c 1 ± 0.6 c 0-20 Xyleborus volvulus 39 20 0.51 b 28.4 ± 10.6 b 0-100 Xyleborus ferrugineus 118 70 0.59 b 33 ± 7.4 b 0-118 Xyleborinus gracilis 52 26 0.50 b 100.6 ± 34 b 0-1240 Xyleborinus saxeseni 68 2 0.03 c 1.5 ± 1 c 0-60 Xylosandrus crassiusculus 39 1 0.03 c 2.6 ± 2.6 c 0-100 Ambrosiodmus devexulus 25 0 - - - Ambrosiodmus lecontei 41 0 - - - Raffaelea lauricola

Two can transmit R. lauricola to avocado

Xyleborus bispinatus (~X. ferrugineus) Can develop and reproduce feeding exclusively on R. lauricola N # with R. lauricola % of beetles with R. lauricola CFU mean CFU range Swampbay 118 70 59 60 0-118 Avocado logs 5 5 100 40 0-80 Avocado logs R. 20 18 90 53 0-320 Traps 35 6 17.1 4.7 0-60

Xyleborus volvulus Carries R. lauricola passively N # with R. lauricola % of beetles with R. lauricola CFU mean CFU range Swampbay 39 20 51 28 0-100 Avocado logs 53 10 19 30 0-1140 Avocado logs R. Saucedo 20 2 3 12 0-20 Traps 117 3 2.6 0.4 0-20

Notoriously difficult to control Feed on fungi not on plants >99% of time hidden inside the tree No management options other than sanitation control beetles inside trees

Monitoring- Scouting Early detection and rapid removal 160 140 120 100 80 60 40 20 0 Logs Carrying R. lauricola Chips Chipping wood is an effective way of killing beetles inside the trees

Chemical Ecology of RAB Attractants Fungal volatiles (Kuhns et al. 2013) Sesquiterpenes (Kairomones) α-copaene, Cubeb, and Eucalyptol (Kendra et al; Kuhns et al.) Martini et al. 2017 Repellents: methyl salicylate (MeSA) & verbenone (Hughes et al.2017)

Insecticides have very limited use: Do not kill ambrosia beetles that are inside the tree. Broadcast sprays do not suppress ambrosia beetle populations. Low persistence - estimated efficacy 2-3 weeks when applied with a sticker.

Fungicides Alamo and Tilt (propiconazole) Macro-infusion process Requires professional help

Biological Control Multiple potential parasitoids and predators associated with infested logs but could not determine if they were AB parasites.

Biological Control Entomopathogenic Fungi Beauveria bassiana Augment beetle pathogens and increase beetle mortality

Host Resistance Propagation of redbay survivors from severely affected sites Screen for resistance to LW pathogen Tolerance redbays in development Hughes and Smith 2014, Native Plants Journal

Euwallacea fornicatus species complex Originally thought to be one species: Tea Shot Hole Borer Several cryptic species, three of which are found in the US: Polyphagous SHB (California) Kuroshio SHB (California) Tea SHB (Florida, Hawaii)

Primary nutritional symbionts- Fusarium fungi (AFC) Fusarium dieback Polyphagous SHB (California) - AF 2 Fusarium euwallaceae Kuroshio SHB (California) AF 12 Fusarium sp. Tea SHB (Florida) AF 6, AF 8, AF 9 Fusarium sp. O'Donnell K, Sink S, Libeskind-Hadas R, Hulcr J, Kasson MT, Ploetz RC, Konkol JL, Ploetz JN, Carrillo D, Campbell A, Duncan RE, Liyanage PNH, Eskalen A, Na F, Geiser DM, Bateman C, Freeman S, Mendel Z, Sharon M, Aoki T, Cossé AA, and Rooney AP. 2015. Discordant phylogenies suggest repeated host shifts in the Fusarium - Euwallacea ambrosia beetle mutualism. Fungal Genetics and Biology 82:277-290.

Early signs: Sugar volcanoes Late signs: Frass - sawdust

Damage Photo: Akif Eskalen attack and kill medium and small branches Interior-shaded braches first, later outer branches base of the branch first, later all the branch can lead to the death of individual branches or, in severe cases, the entire tree

Impact Urban Forests ~30% of street trees in So. CA are susceptible spp. Tree removal costs about $1000 per tree Danger of falling branches National Forests, State Forests Many native, threatened, or endangered species are highly susceptible (California sycamore, red willow, white alder, coast live oak, etc.) Riparian dominant spp. at risk http://eskalenlab.ucr.edu/distribution.html

64 Hosts Support Beetle Reproduction in California 1.Box Elder (Acer negundo)* 2.Big Leaf Maple (Acer macrophyllum)* 3. Evergreen Maple (Acer paxii) 4. Trident Maple (Acer buergerianum) 5. Japanese Maple (Acer palmatum) 6. Castorbean (Ricinus communis) 7. California Sycamore (Platanus racemosa)* 8. Mexican Sycamore (Platanus mexicana) 9. Red Willow (Salix laevigata)* 10. Arroyo Willow (Salix lasolepis)* 11. Avocado (Persea americana) 12. Mimosa (Albizia julibrissin) 13. English Oak (Quercus robur) 14. Coast Live Oak (Quercus agrifolia)* 15. London Plane (Platanus x acerifolia) 16. Cottonwood (Populus fremontii)* 17. Black Cottonwood (Populus trichocarpa)* 18. White Alder (Alnus rhombifolia)* 19. Titoki (Alectryon excelsus) 20. Engelmann Oak (Quercus engelmannii)* 21. Cork Oak (Quercus suber) 22. Valley Oak (Quercus lobata)* 23. Coral Tree (Erythrina coralloides) 24. Blue Palo Verde (Cercidium floridum)* 25. Palo Verde (Parkinsonia aculeata)* 26.Moreton Bay Chestnut (Castanospermum australe) 27. Brea (Cercidium sonorae) 28. Mesquite (Prosopis articulata)* 29. Weeping Willow (Salix babylonica) 30. Chinese Holly (Ilex cornuta) Eskalen and Lynch, 2017. Unpublished data 31. Camelia (Camellia semiserrata) 32. Acacia (Acacia spp.) 33. Japanese Wisteria (Wisteria floribunda) 34. Black Willow (Salix gooddingii)* 35. Tree of Heaven (Ailanthus altissima) 36. Kurrajong (Brachychiton populneus) 37. Black Mission fig (Ficus carica)** 38. Japanese Beech (Fagus crenata) 39. Dense Logwood (Xylosma avilae) 40. Mule Fat (Baccharis salicina)* 41. Black Poplar (Populus nigra) 42. Carrotwood (Cupaniopsis anacardioides) 43. California Buckeye (Aesculus californica)* 44. Canyon Live Oak (Quercus chrysolepis)* 45. Kentia Palm (Howea forsteriana) 46. King Palm (Archontophoenix cunninghamiana) 47. Tamarix (Tamarix ramosissima 48- Red Flowering Gum (Eucalyptus ficifolia)** 49- American Sweetgum (Liquidambar styraciflua) 50- Honey Locust (Gleditsia triacanthos) 51. Brazilian Coral Tree (Erythrina falcata) 52. Purple Orchid Tree (Bauhinia variegata)** 53. Council Tree (Ficus altissima)** 54. Tulip Wood (Harpullia pendula) 55. Chinese Flame Tree (Koelreuteria bipinnata)** 56. Laurel-leaf Snailseed tree (Cocculus laurifolius)** 57. Southern Magnolia (Magnolia grandiflora)** 58. Jacaranda (Jacaranca mimosifolia)** 59. Coast coral tree (Erythrina caffra)** 60. Australian blackwood (Acacia melanoxylon) 61. Sweet Bay (Magnolia virginiana)** 62. African Tulip Tree (Spathodea campanulata)** 63. Strawberry snowball tree (Dombeye cacuminum)** 64. Chinese Wingnut (Pterocarya stenoptera)** http://eskalenlab.ucr.edu/distribution.html 19 CA Native (*) Canker associated (**)

Surveyed Distribution TSHB in the avocado growing region of Florida Persea americana Mill., (Lauraceae) Lysiloma latisiliquum (L.) Bentham (Fabaceae) Annona muricata L. (Annonaceae) Albizia lebbeck (L.) Bentham (Fabaceae) Manguifera indica (L.) Anacardiaceae Delonixregia Sarg. (Fabaceae) Persea palustris (Lauraceae)

Chemical Ecology Quercivorol Pheromone" of the ambrosia beetle Platypus quercivorus (Kashigawi et al 2006) Fungal Kairomone (1S,4R)-p-Menth-2-en-1-ol (Quercivorol) Carrillo et al. 2015. Attraction of Euwallacea nr. fornicatus to lures containing quercivorol. Florida Entomologist 98:780-782. http://dx.doi.org/10.1653/024.098.0258 Dodge et al. 2017. Quercivorol as a lure for the polyphagous and Kuroshio shot hole borers, Euwallacea spp. nr. fornicatus (Coleoptera: Scolytinae), vectors of Fusarium dieback. PeerJ 5:e3656. DOI 10.7717/peerj.3656

Chemical Ecology Tea Shot Hole Borer (Florida) Quercivorol & α-copaene Quercivorol α-copaene 2017

Management? Sanitation Insecticide - injections (California) Emamenctin Benzoate, Imidacloprid Repellents? Biological control, parasitoids?

Thank you! Questions? Tropical Research and Education Center