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Potential Effects of Laurel Wilt on the Flora of North America
Joel M. Gramling

Southeastern Naturalist, Volume 9, Issue 4 (2010): 827–836

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2010 SOUTHEASTERN NATURALIST 9(4):827–836 Potential Effects of Laurel Wilt on the Flora of North America Joel M. Gramling* Abstract - Laurel Wilt is caused by a recently identified fungal pathogen infecting plants in the Lauraceae. Laurel Wilt is transmitted by Xyleborus glabratus (Redbay Ambrosia Beetle), which was recently introduced to the southeastern United States from Asia. As the insect expands its range in the US, so too has the disease. A query of the NatureServe Explorer database was used to identify the conservation status of native plant species and recognized plant communities that may be affected by Laurel Wilt. Laurel Wilt affects at least nine plant species, and all fifteen species in the Lauraceae currently found in North America may ultimately prove to be hosts for the disease. Four of the twelve native lauraceous species had been identified as vulnerable to extirpation or extinction, prior to the introduction of Laurel Wilt. There are 55 plant communities in the US and Canada that have a member of the Lauraceae as a dominant or diagnostic species. The majority of these plant communities have been identified as vulnerable. Agricultural industries that are based upon Persea americana (Avocado) cultivation in Florida and California are threatened by Laurel Wilt as well. Given the potential impact of this disease on lauraceous plants and their associated communities, these taxa and assemblages should be monitored for the arrival and impact of Laurel Wilt. Introduction In 2002, an unknown ambrosia beetle was collected in a funnel trap at Port Wentworth near Savannah, GA (Rabaglia 2003). Shortly thereafter, dead and dying Persea borbonia (L) Spreng. (Redbay) were observed on nearby Hilton Head Island, SC. Specimens of the beetle were collected from dying Redbays and subsequently identified as Xyleborus glabratus Eichhoff (Redbay Ambrosia Beetle), a native of eastern Asia (Rabaglia 2006). The Redbay Ambrosia Beetle hosts a fungal symbiont that is introduced into susceptible plant species when the female beetle begins excavating galleries to lay its eggs (Harrington et al. 2008). The fungus, Raffaelea lauricola T.C. Harr., Fraedrich, & Aghayeva sp. nov. (Harrington et al. 2008), is consumed by larvae of the beetle, and causes Laurel Wilt (Fraedrich et al. 2008). In Redbay and Persea palustris (Raf.) Sarg. (Swampbay), the disease induces rapid wilting followed by branch and stem die-back. Within weeks, previously healthy individuals of Redbay and Swampbay die but retain their brown, necrotic leaves (Fraedrich et al. 2008, Harrington et al. 2008). Laurel Wilt has spread quickly across the south Atlantic coastal region since it was first observed on Hilton Head Island, SC. As of August 2009, Laurel Wilt had been observed in 11 counties in southeastern South Carolina, 24 counties in eastern Georgia, 22 counties across Florida, and in Jackson County, MS (USFS 2009). Koch and Smith (2008) calculated that the Redbay Ambrosia Beetle is spreading at a rate of 54.8 km/yr. Based on *Department of Biology, The Citadel, 171 Moultrie Street, Charleston, SC 29409; joel.gramling@citadel.edu. 828 Southeastern Naturalist Vol. 9, No. 4 the distributions of P. borbonia sensu lato (including P. palustris) and Sassafras albidum (Nutt.) Nees (Sassafras, another host), it was estimated that the northern and southern limits of P. borbonia sensu lato would be affected by Laurel Wilt within the next decade and its western limits would be affected by 2040 (Koch and Smith 2008). Anthropogenic movement of the vector, which has been suspected in range extensions in Florida and Mississippi, is increasing this rate of spread. Better understandings of the disease’s host range and epidemiology would be needed to predict its ultimate impact on native plant communities and new host species. Most Laurel Wilt research has focused on the pathogen, its vector, and their interactions with host plants. Studies on the communities that have and will be impacted are needed to fully understand the potential ecological impact of this disease. To this end, a list of vulnerable plant species and plant communities in the US and Canada has been produced. Monitoring them for the arrival and impact of Laurel Wilt is discussed below. Methods The NatureServe Explorer database (http://www.natureserve.org/explorer) was used to identify the plant species of the Lauraceae family that occur in the US and Canada. It was also used to identify ecological communities in which a member of the Lauraceae is a dominant or diagnostic species. Plant communities included in this study followed the IVCS/NVCS conventions (Grossman et al. 1998). The IVCS/NVCS system is a hierarchy of classifi- cations from which the plant association level was chosen for this project. A plant association represents “the basic unit of inventory for biodiversity assessment” and is defined as “a plant community of definite floristic composition, presenting a uniform physiognomy, and growing in uniform habitat conditions” (Grossman et al. 1998; see also Flahault and Schroter 1910). Henceforth, the term “community” will refer directly to association-level classifications in the IVCS/NVCS. NatureServe assigns a conservation status for both plants and plant communities. A global conservation status was given using the following ranked scale: G5 = secure, G4 = apparently secure, G3 = vulnerable, G2 = imperiled, and G1 = critically imperiled. Species or communities ranked as G5 are “common, widespread and abundant”, while those assigned a rank of G1 are “at very high risk of extinction due to extreme rarity (often 5 or fewer populations), steep declines, or other factors” (NatureServe 2010). A literature review was used to identify lauraceous species that have been artificially or naturally affected by Laurel Wilt. Results Twelve native plant species and three introduced species from the Laurel family may be at risk (Table 1). Four of these native plant species are considered globally imperiled or vulnerable to extirpation or extinction: Lindera melissifolia (Walter) Blume (Pondberry), Lindera subcoriacea B.E. Wofford (Bog Spicebush), Litsea aestivalis (L.) Fern. (Pondspice), and Persea humilis Nash (Silkbay) (Table 1). There are 56 native plant communities in 2010 J.M. Gramling 829 the United States that have a member of the laurel family as a dominant or diagnostic plant species (NatureServe 2010; Appendix A), of which nine are critically imperiled (G1), 17 are imperiled (G2), and 11 are vulnerable (G3) communities (Table 2, Appendix A). Without significant conservation efforts, critically imperiled and imperiled communities (i.e., communities ranked as G1 or G2) are poised for global extinction (NatureServe Explorer 2010). There were no plant communities dominated or diagnosed by Lauraceous species attributed to Canada. The Laurel Wilt pathogen, R. lauricola, has been isolated from dead or wilted individuals of Cinnamomum camphora (L.) J. Presl. (Camphortree), Persea americana P. Mill. (Avocado), Redbay, Swampbay, Sassafras, Silkbay, Pondspice, and Pondberry (Fraedrich et al. 2008; Mayfield et al. 2008c; Smith et al. 2009a, 2009b; R.C. Ploetz, Tropical Research and Education Center, University of Florida, Homestead, fl, unpubl. data). Under experimental conditions, R. lauricola has also caused Laurel Wilt on Lindera benzoin (L.) Blume (Northern Spicebush), Umbellularia californica (Hook. & Arn.) Nutt. (California Laurel), Avocado, Redbay, and Swampbay (Fraedrich 2008, Fraedrich et al. 2008, Mayfield et al. 2008b). In total, seven Table 1. Conservation rankings and distribution of lauraceous species in the United States and Canada. Species in bold print have been affected by Laurel Wilt under natural conditions. G5 = secure, G4 = apparently secure, G3 = vulnerable, G2 = imperiled, G1 = critically imperiled, NN = non-native species. Species States or Species rank Region Provinces Common names Cassytha filiformis G4G5 Tropical 3: fl, HI, TX Fine-stem Lovevine Cinnamomum camphora NN Southeast and 7: AL, fl, GA, Camphortree Tropics HI, LA, SC, TX Laurus nobilis NN West 1: CA Bay Laurel Licaria triandra G4G5 Tropical fl1: flGulf Licaria Lindera benzoin† G5 Eastern US 34: 33 states and Northern Spicebush ON, Canada Lindera melissifolia* G2 Southeast 8: AL, AR, GA, LA, Pondberry MO, MS, NC, SC Lindera subcoriacea G2/G3 Southeast 8: AL, fl, GA, LA, Bog Spicebush MS, NC, SC, VA Litsea aestivalis G3 Southeast 7: fl, GA, LA, Pondspice MD, NC, SC, VA Nectandra coriacea G5 Tropical fl1: flLancewood Persea americana NN Tropical 3: CA, fl, HI Avocado Persea borbonia G5 Southeast 9: AL, AR, fl, Redbay GA, LA, MS, NC, SC, TX, VA Persea humilis G3 flScrub 1: flSilkbay Persea palustris G5 Southeast 11: AL, DE, fl, Swampbay GA, LA, MD, MS, NC, SC, TX, VA Sassafras albidum G5 Eastern US 34: 33 states and Sassafras ON, Canada Umbellularia californica† G5 West 2: CA, OR California Laurel †Species that have developed Laurel Wilt after artificial inoculation. *Extirpated in Florida. 830 Southeastern Naturalist Vol. 9, No. 4 species have been naturally affected by the disease and two additional species, Northern Spicebush and California Laurel, have proven susceptible after artificial inoculation with R. lauricola (Table 1). Discussion The flora of North America is highly vulnerable to Laurel Wilt. To date, Laurel Wilt has impacted plant communities from coastal South Carolina to south Florida, in particular those in which Redbay is/was prevalent. The longterm survival rates of Redbay and other susceptible species in the affected areas are not known, but may be poor given the high mortality observed in the short term (Fraedrich et al. 2008; J.M. Gramling and D.J. Gustafson, unpubl. data). Large caliper trees (those of a reproductive age) are killed most rapidly, and smaller diameter individuals usually succumb eventually (Fraedrich et al. 2008; J.M. Gramling and D.J. Gustafson, unpubl. data). Further study of seed production and survivorship in the presence of Laurel Wilt is needed for all species in this family in the southeastern United States. The long-term survival of non-native lauraceous species is not clear, but it may differ from that of US endemics, especially when the species originated in the beetle’s native range. For example, Camphortree, an east Asian endemic, is susceptible but often recovers to the extent that symptoms of the disease are not visible in years that follow disease onset (Smith et al. 2009b). Understanding which species are resistant to Laurel Wilt and how different species tolerate infection by R. lauricola will help predict how this disease will impact native and non-native lauraceous plants and the communities in which they reside. Avocado is a tropical American member of the Lauraceae. California and Florida are the top producers of avocadoes in the US, where they make up Table 2: Summary of conservation rankings for communities associated with Lauraceous species in the United States. Species in bold print have been affected by Laurel Wilt under natural conditions. G5 = secure, G4 = apparently secure, G3 = vulnerable, G2 = imperiled, G1 = critically imperiled, NN = non-native species. Total = total number of communities. # of communities for each rank Species Region Total G5 G4 G3 G2 G1 NR Cassytha filiformis Tropical 0 0 0 0 0 0 0 Cinnamomum camphora‡ Southeast and tropics 0 0 0 0 0 0 0 Laurus nobilis‡ West 0 0 0 0 0 0 0 Licaria triandra Tropical fl0 0 0 0 0 0 0 Lindera benzoin Eastern US 13 0 5 3 1 1 3 Lindera melissifolia Southeast 1 0 0 0 1 0 0 Lindera subcoriacea Southeast 0 0 0 0 0 0 0 Litsea aestivalis Southeast 0 0 0 0 0 0 0 Nectandra coriacea Tropical fl2 0 0 0 0 2 0 Persea americana‡ Tropical 0 0 0 0 0 0 0 Persea borbonia Southeast 7 0 0 0 6 1 0 Persea humilis flscrub 0 0 0 0 0 0 0 Persea palustris Southeast 17 0 2 2 8 5 0 Sassafras albidum Eastern US 4 1 0 0 1 0 2 Umbellularia californica West 12 0 6 6 0 0 0 ‡Species that are not native to the United States and Canada. 2010 J.M. Gramling 831 90% and 9% of domestic avocado production, respectively (USDA 2008). Although Laurel Wilt has not invaded commercial production areas in California or Florida, the available evidence indicates that the crop is susceptible, and that the US avocado industry could be at risk (Mayfield et al. 2008b, Ploetz et al. 2009). The invasion of commercial areas in the US and tropical America (Mexico is the world’s most important producer) could result in dramatic fiscal losses and further impact natural plant communities. The ultimate effects of Laurel Wilt may not be observed until the pathogen and vector shift from invaders to residents in a given area. Invasions of plant pathogens are often driven by the availability of a host population in the short-term, whereas their persistence may be confronted with diminished host numbers and a variety of demographic or environmental factors (Gilligan and van den Bosch 2008). As Persea spp. and other host plants in the Southeast are reduced in number and in average individual size, populations of the Redbay Ambrosia Beetle will likely decline. Hanula et al. (2008) demonstrated that population densities of the beetle were lower in areas in which host densities had declined due to Laurel Wilt; peak populations declined as the age of infestations increased. Continued study of the X. glabaratus - R. lauricola - Lauraceous species complex should incorporate both recently invaded sites and areas already affected by the disease. This work may be complicated by the differing susceptibilities of the different host species to Laurel Wilt, their varying levels of attractiveness to the Redbay Ambrosia Beetle, and the distributions and sizes (i.e., patchiness) of their populations (Park et al. 2002). Monitoring and modeling the effects of Laurel Wilt must consider all of the above factors. Laurel Wilt was first observed on native flora of North America in 2003, but the disease’s full impact will not be realized until it has spread to all areas in which susceptible species grow. Exotic forest tree diseases, such as Dutch Elm Disease and Chestnut Blight, have had far-ranging ecological impacts on other native plant and animal species (Anderson et al. 2004). Laurel Wilt not only threatens members of the Lauraceae, but also the integrity of plant communities of which they are components. Exotic tree diseases can alter habitat suitability within a community and indirectly affect other species via changes in light availability, increases in litter and woody debris, changes in the soil, and other factors (Dobson and May 1986). Associated faunal species may be impacted by the spread of Laurel Wilt as well. The availability of fruits and nectar from host species will likely decrease as the average size of individuals declines. The greatest faunal impacts may be seen in Papilio palamedes Drury (Palamedes Swallowtail Butterfly), whose larvae are only known to feed on plants in the laurel family, and Papilio troilus L. (Spicebush Swallowtail), which uses Sassafras and Northern Spicebush as its preferred host plants (Scott 1986). The decline of these and other associated insects could have unforeseen effects on the plants that they pollinate or other animals with which they interact. Non-native Lauraceous species found in North America may also impact the relationship between the Laurel Wilt pathogen and native plant species. In general, a patchy or discontinuous distribution of a species helps to buffer local populations from diseases (Dobson and May 1986). The distribution of 832 Southeastern Naturalist Vol. 9, No. 4 ornamental plants (such as Camphortree) and crop plants (such as Avocado) does not necessarily mirror the distribution of native plant species and thus could provide alternative hosts across the landscape that act as stepping stones between native plant populations. For the four plant species that have been classified as globally imperiled or vulnerable to extinction, their proximity to either broadly distributed native Lauraceous species or cultivated non-native Lauraceous species may be of particular importance in assessing their potential to be further reduced in number by Laurel Wilt. Laurel Wilt is a relatively new disease and much is still unknown about how it will impact the flora of North America. More research is needed to better understand the effects of Laurel Wilt on plant communities over time. The large-scale losses associated with previous exotic fungal pathogens in North America, including Dutch Elm Disease and Chestnut Blight, provide precedents for a potentially catastrophic toll that Laurel Wilt could take on members of the Lauraceae. A concerted effort is needed to monitor and mitigate the spread of this disease as it continues to impact our native plant communities. Acknowledgments D.J. Gustafson , A.E. Gramling, R.C. Ploetz and two anonymous reviewers provided helpful comments on earlier versions of this manuscript. This research was supported in part by a grant from The Citadel Foundation. Literature Cited Anderson, P.K., A.A. Cunningham, N.G. Patel, F.J. Morales, P.R. Epstein and P. Daszak. 2004. Emerging infectious diseases of plants: Pathogens pollution, climate change, and agrotechnology drivers. Trends in Ecology and Evolution 19:535–544. Dobson, A.P., and R.M. May. 1986. Disease and conservation. Pp. 345–365, In M.E. Soule (Ed.). Conservation Biology: The Science of Scarcity and Diversity. Sinauer Associates, Sunderland, MA. 584 pp. Flahault, C., and C. Schroter. 1910. Rapport sur la nomenclature phytogeopraphique. Proceedings of the Third International Botanical Congress, Brussels 1:131–164. Fraedrich, S.W. 2008. California Laurel is susceptible to laurel wilt caused by Raffaelea lauricola. Plant Disease 92:1469. Fraedrich, S.W., T.C. Harrington, R.J. Rabaglia, M.D. Ulyshen, A.E. Mayfield III, J.L. Hanula, J.M. Eickwort, and D.R. Miller. 2008. A fungal symbiont of the Redbay Ambrosia Beetle causes a lethal wilt in Redbay and other Lauraceae in the southeastern United States. Plant Disease 92:215–224. Gilligan, C.A., and F. van den Bosch. 2008. Epidemiological models for invasion and persistence of pathogens. Annual Review of Phytopathology 46:385–418. Grossman, D.H., D. Faber-Langendoen, A.S. Weakley, M. Anderson, P. Bourgeron, R. Crawford, K. Goodin, S. Landaal, K. Metzler, K.D. Patterson, M. Pyne, M. Reid, and L. Sneddon. 1998. International Classification of Ecological Communities: Terrestrial Vegetation of the United States. Vol. 1. The National Vegetation Classification System: Development, Status, and Applications. The Nature Conservancy, Arlington, VA. 127 pp. 2010 J.M. Gramling 833 Hanula, J.L., A.E. Mayfield III, S.W. Fraedrich, and R.J. Rabaglia. 2008. Biology and host associations of Redbay Ambrosia Beetle (Coleoptera: Curculionidae: Scolytinae), exotic vector of Laurel Wilt killing Redbay trees in the southeastern United States. Journal of Economic Entomology 101:1276–1286. Harrington, T.C., S.W. Fraedrich and D.N. Aghayeva. 2008. Raffaelea lauricola, a new ambrosia beetle symbiont and pathogen on the Lauraceae. Mycotaxon 104:399–404. Koch, F.H., and W.D. Smith. 2008. Spatio-temporal analysis of Xyleborus glabratus (Coleoptera: Circulionidae: Scolytinae) invasion in eastern US forests. Environmental Entomology 37(2):442–452. Mayfield, A.E. III, E.L. Barnard, J.A. Smith, S.C. Bernick, J.M. Eickwort, and T.J. Dreaden. 2008a. Effect of propiconazole on Laurel Wilt disease development in Redbay trees and on the pathogen in vitro. Arboriculture and Urban Forestry 34:317–324. Mayfield, A.E. III, J.E. Pena, J.H. Crane, J.A. Smith, C.L. Branch, E.D. Ottoson, and M. Hughes. 2008b. Ability of the Redbay Ambrosia Beetle (Coleoptera: Curculionidae: Scolytinae) to bore into young Avocado (Lauraceae) plants and transmit the Laurel Wilt pathogen (Raffaelea lauricola). Florida Entomologist 91:485–487. NatureServe. 2010. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.0. NatureServe, Arlington, VA. Available online at http:// www.natureserve.org/explorer. Accessed 1 April 2010. Park, A.W., S. Gubbins, and C.A. Gilligan. 2002. Extinction times for closed epidemics: The effects of host spatial structure. Ecology Letters 5:747–755. Ploetz, R.C., G. Bender, R. Bostock, R. Bulluck, J. Crane, M. Draper, A. Eskalen, G. Evans, B. Faber, S. Fraedrich, J. Hanula, C. Harmon, T. Harrington, A. Palmateer, J. Pena, R. Schnell, J. Smith, K. Smith, M. Wingfield. 2009. A national plant disease recovery plan for Laurel Wilt of Avocado. Phytopathology 99:S102. Rabaglia, R.J. 2003. Pest Report: Xyleborus glabratus Available online at http://spfnic. fs.fed.us/exfor/data/pestreports.cfm?pestidval=148&langdisplay=english. Accessed 10 September 2009. Rabaglia, R.J., S.A. Dole, and A.I. Cognato. 2006. Review of American Xyleborina (Coleoptera: Curculionidae: Scolytinae) occurring north of Mexico, with an illustrated key. Annals of the Entomological Society of America 99:1034–1056. Scott, J.A. 1986. The Butterflies of North America: A Natural History and Field Guide. Stanford University Press, Stanford, CA. 583 pp. Smith, J.A., T. J. Dreaden, A.E. Mayfield III, A. Boone, S.W. Fraedrich and C. Bates. 2009a. First Report of Laurel Wilt Disease Caused by Raffaelea lauricola on Sassafras in Florida and South Carolina. Plant Disease 93:1079. Smith, J.A., L. Mount, A.E. Mayfield III, C.A. Bates, W.A. Lamborn, and S.W. Fraedrich. 2009b. First report of Laurel Wilt Disease caused by Raffaelea lauricola on Camphor in Florida and Georgia. Plant Disease 93:198. US Department of Agriculture, Foreign Agriculture Service (USDA). 2008. January 2008 Market news: Avocado update. Available online at http://www.fas.usda.gov/ htp/horticulture/Avocados/2008_Avocado_Article.pdf. Accessed 10 Sept 2009. US Forest Service (USFS) 2009. Distribution of counties with Laurel Wilt Disease by year of initial detection. Available online at http://www.fs.fed.us/r8/foresthealth/ laurelwilt/dist_map.shtml. Accessed 10 September 2009. 834 Southeastern Naturalist Vol. 9, No. 4 Appendix 1. Lauraceous community associations, their global rank, the states in which they are found, and the key lauraceous species they contain. Community (association name) CEGL code Rank States Lauraceous species Alnus serrulata - Lindera benzoin / Osmunda regalis var. spectabilis - Carex tetanica Shrubland CEGL008408 G1 VA Lindera benzoin Bursera simaruba - Swietenia mahagoni - Lysiloma latisiliquum / Nectandra coriacea - CEGL007003 G1 flNectandra coriacea Coccoloba diversifolia Forest Ficus aurea - Sideroxylon foetidissimum - Bursera simaruba / Eugenia foetida - Guapira CEGL007001 G1 flNectandra coriacea discolor - Nectandra coriacea Forest Quercus virginiana - Sabal palmetto - Persea borbonia / Myrcianthes fragrans - CEGL007033 G1 flPersea borbonia Ardisia escallonoides - Psychotria nervosa Forest Liquidambar styraciflua / Persea palustris Forest CEGL004481 G1 NC Persea palustris Magnolia virginiana - Persea palustris - Chrysobalanus icaco / Acrostichum danaeifolium - CEGL007015 G1 flPersea palustris Nephrolepis exaltata Forest Magnolia virginiana - Persea palustris - Chrysobalanus icaco / Cladium mariscus ssp. CEGL007016 G1 flPersea palustris jamaicense Woodland Persea palustris / Morella cerifera Maritime Forest CEGL004635 G1 NC, SC Persea palustris Quercus falcata - Pinus taeda - (Fagus grandifolia, Quercus nigra) / Persea palustris Maritime CEGL007540 G1 NC, VA Persea palustris Forest Alnus serrulata - Lindera benzoin / Scutellaria lateriflora - Thelypteris noveboracensis Shrubland CEGL003909 G2 NC Lindera benzoin Quercus lyrata - Quercus palustris / Acer rubrum var. drummondii / Itea virginica - CEGL004778 G2 AR, MO Lindera melissifolia Cornus foemina - (Lindera melissifolia) Forest Fagus grandifolia - Magnolia grandiflora / Ilex opaca - (Persea borbonia) / Mitchella repens CEGL007459 G2 GA, SC Persea borbonia Forest Quercus alba - Quercus hemisphaerica / Prunus caroliniana - Persea borbonia - CEGL007959 G2 TX Persea borbonia Viburnum acerifolium Forest Quercus fusiformis - Persea borbonia Forest CEGL002117 G2 TX Persea borbonia Quercus geminata - Quercus myrtifolia - Serenoa repens - Persea borbonia Shrubland CEGL003821 G2 flPersea borbonia Quercus hemisphaerica - Quercus geminata / Persea borbonia - Osmanthus americanus Forest CEGL004787 G2 fl, GA Persea borbonia Quercus virginiana - (Pinus elliottii var. elliottii, Sabal palmetto) / Persea borbonia - CEGL007032 G2 AL, fl, Persea borbonia Callicarpa americana Forest GA, NC, SC (Pinus elliottii) / Cyrilla racemiflora - Persea palustris - Magnolia virginiana - CEGL004974 G2 AL, MS Persea palustris Smilax laurifolia Shrubland 2010 J.M. Gramling 835 Community (association name) CEGL code Rank States Lauraceous species Chamaecyparis thyoides / Persea palustris / Lyonia lucida - Ilex coriacea Forest CEGL006146 G2 NC, VA Persea palustris Cyrilla racemiflora - Persea palustris - Magnolia virginiana Shrubland CEGL004449 G2 NC Persea palustris Quercus virginiana - Quercus hemisphaerica - Pinus taeda - Quercus falcata / Persea CEGL007026 G2 NC,SC Persea palustris palustris Forest Quercus virginiana - Quercus hemisphaerica - Pinus taeda / Persea (borbonia, palustris) - CEGL007027 G2 NC, SC, Persea palustris Ilex vomitoria Forest VA Taxodium ascendens / Chrysobalanus icaco - Ficus aurea - Persea palustris Forest CEGL007416 G2 flPersea palustris Taxodium distichum - Nyssa aquatica - Persea palustris Forest CEGL004681 G2 flPersea palustris Taxodium distichum / Persea palustris - Fraxinus caroliniana - Chrysobalanus icaco / CEGL007440 G2 flPersea palustris Blechnum serrulatum Forest Prunus serotina - Sassafras albidum - Amelanchier canadensis - Quercus velutina / Smilax CEGL006145 G2 CT, DE, Sassafras albidum rotundifolia Forest MA, NH, NJ, NY, RI Liriodendron tulipifera - Platanus occidentalis - Betula lenta / Lindera benzoin / Circaea CEGL006255 G3 MD, VA Lindera benzoin lutetiana ssp. canadensis Forest Quercus alba - Quercus rubra - Acer saccharum - Carya cordiformis / Lindera benzoin Forest CEGL002058 G3 AR, Lindera benzoin MO, OK Magnolia virginiana - Persea palustris / Lyonia lucida Forest CEGL007049 G3 fl, GA, Persea palustris SC Pinus serotina / Gordonia lasianthus - Persea palustris Saturated Woodland CEGL007996 G3 fl, GA Persea palustris Acer macrophyllum - Pseudotsuga menziesii - Umbellularia californica / Polystichum CEGL000518 G3 OR Umbellularia californica munitum Forest Pseudotsuga menziesii - Tsuga heterophylla / Lithocarpus densiflorus - Umbellularia CEGL000087 G3 OR Umbellularia californica californica Forest Quercus agrifolia - Umbellularia californica / Ceanothus oliganthus Woodland CEGL002870 G3 CA Umbellularia californica Quercus agrifolia - Umbellularia californica Woodland CEGL002869 G3 CA Umbellularia californica Umbellularia californica - Juglans californica / Ceanothus spinosus Forest CEGL002876 G3 CA Umbellularia californica Umbellularia californica - Platanus racemosa Forest CEGL002877 G3 CA Umbellularia californica Liquidambar styraciflua - (Quercus alba, Acer saccharum) / Carpinus caroliniana / Lindera CEGL007826 G3 AR, OK Lindera benzoin benzoin Forest Fagus grandifolia - Liriodendron tulipifera - Carya cordiformis / Lindera benzoin / Podophyllum CEGL006055 G4 DC, DE, Lindera benzoin peltatum Forest MD, NJ, VA, WV 836 Southeastern Naturalist Vol. 9, No. 4 Community (association name) CEGL code Rank States Lauraceous species Liquidambar styraciflua - Liriodendron tulipifera / Lindera benzoin / Arisaema triphyllum Forest CEGL004418 G4 DC, GA, Lindera benzoin MD, NC, TN, VA Platanus occidentalis - Celtis laevigata - Fraxinus pennsylvanica / Lindera benzoin - Ilex CEGL007730 G4 SC Lindera benzoin decidua / Carex retroflexa Forest Platanus occidentalis - Celtis laevigata - Liriodendron tulipifera / Lindera benzoin - Arundinaria CEGL008429 G4 AL, GA, Lindera benzoin gigantea / Amphicarpaea bracteata Forest KY, TN Gordonia lasianthus - Magnolia virginiana - Persea palustris / Sphagnum spp. Forest CEGL007044 G4 fl, GA, Persea palustris NC, SC Quercus laurifolia - Quercus lyrata / Carpinus caroliniana - Persea palustris / Vaccinium CEGL004737 G4 NC, SC Persea palustris elliottii Forest Pseudotsuga menziesii - Umbellularia californica / Toxicodendron diversilobum Forest CEGL000073 G4 CA, OR Umbellularia californica Pseudotsuga menziesii / Umbellularia californica / Frangula californica ssp. californica Forest CEGL003167 G4 CA Umbellularia californica Pseudotsuga menziesii / Umbellularia californica / Polystichum munitum Forest CEGL003168 G4 CA Umbellularia californica Quercus chrysolepis - Umbellularia californica Forest CEGL008604 G4 CA Umbellularia californica Umbellularia californica - Quercus agrifolia / Toxicodendron diversilobum Forest CEGL003175 G4 CA Umbellularia californica Umbellularia californica / Polystichum munitum Forest CEGL003174 G4 CA Umbellularia californica Acer rubrum - Fraxinus (pennsylvanica, americana) / Lindera benzoin / Symplocarpus CEGL006406 G4 CT, DC, Lindera benzoin foetidus Forest DE, IN, MA, MD, NH, NJ, NY, PA, RI, VA, VT Sassafras albidum - Quercus spp. Forest CEGL004096 G5 VA Sassafras albidum Liriodendron tulipifera - Fraxinus spp. / Lindera benzoin - Viburnum prunifolium / Podophyllum CEGL006314 NR NJ, PA Lindera benzoin peltatum Forest Quercus alba - Quercus rubra - Quercus prinus - Acer saccharum / Lindera benzoin Forest CEGL002059 NR OH, PA Lindera benzoin Quercus palustris - (Fraxinus nigra) / Lindera benzoin / Carex bromoides Forest CEGL007399 NR WV Lindera benzoin Quercus coccinea - Quercus velutina / Sassafras albidum / Vaccinium pallidum Forest CEGL006375 NR CT, MA, Sassafras albidum NH, NJ, NY, RI Quercus stellata - Sassafras albidum / Smilax glauca Woodland CEGL006372 NR NJ, NY Sassafras albidum