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.
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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