Juglans cinerea

L.

Butternut

G3Vulnerable Found in 18 roadless areas NatureServe Explorer →
G3VulnerableGlobal Rank
EndangeredIUCN
HighThreat Impact
Butternut (Juglans cinerea). Photo by Michael Kondratowicz, CC BY-NC 4.0, via iNaturalist.
Michael Kondratowicz, CC BY-NC 4.0
Butternut (Juglans cinerea). Photo by Michael Kondratowicz, CC BY-NC 4.0, via iNaturalist.
Michael Kondratowicz, CC BY-NC 4.0
Butternut (Juglans cinerea). Photo by warrenmickley, CC BY-NC 4.0, via iNaturalist.
warrenmickley, CC BY-NC 4.0
Butternut (Juglans cinerea). Photo by warrenmickley, CC BY-NC 4.0, via iNaturalist.
warrenmickley, CC BY-NC 4.0
Butternut (Juglans cinerea). Photo by Tom Norton, CC BY 4.0, via iNaturalist.
Tom Norton, CC BY 4.0
Identity
Unique IDELEMENT_GLOBAL.2.154365
Element CodePDJUG02030
Record TypeSPECIES
ClassificationSpecies
Classification StatusStandard
Name CategoryVascular Plant
IUCNEndangered
Endemicoccurs (regularly, as a native taxon) in multiple nations
KingdomPlantae
PhylumAnthophyta
ClassDicotyledoneae
OrderJuglandales
FamilyJuglandaceae
GenusJuglans
Other Common Names
butternut (EN) Noyer cendré (FR)
Concept Reference
Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
Conservation Status
Rank MethodLegacy Rank calculation - Biotics v1
Review Date2019-12-20
Change Date2019-12-20
Edition Date2019-12-20
Edition AuthorsOliver, L.
Threat ImpactHigh
Range Extent200,000-2,500,000 square km (about 80,000-1,000,000 square miles)
Number of Occurrences> 300
Rank Reasons
Juglans cinerea occurs infrequently in forest stands throughout most of the eastern United States and Canada. The abundance and condition are both in decline due to butternut canker disease, with no known remedy. Even with the canker evident and widespread, there are a large number of occurrences persisting and resistant trees, though rare, are found in many areas of the range. Declines are estimated at 44% rangewide. A number of measures are taking place to halt extinction: locating resistant butternut, collecting seed for germplasm, and hybridizing with the Japanese walnut (Juglans ailantifolia). Reintroductions are in consideration in areas where there may be disease resistance.
Range Extent Comments
Juglans cinerea occurs throughout the central and eastern United States and southeastern Canada. It extends from New Brunswick, Canada south to North Carolina; to the north into Minnesota, and west to Missouri. Other isolated populations are known from Arkansas, Mississippi, Alabama, Georgia and South Carolina (Morin et al. 2018).
Threat Impact Comments
The major threat to Juglans cinerea throughout its entire range is susceptibility to the butternut canker disease caused by the fungus Ophiognomonia clavigignenti-juglandacearum (= Sirococcus clavigignenti-juglandacearum). The fungus disrupts nutrient flow through cambium areas, which is generally fatal. It may take trees more than 40 years to die, but in many cases, death has rapidly followed infection. Following dieback, this species does not leave live rootsprouts and usually does not leave viable seed. The wood, which is highly prized for cabinet-making and other types of woodworking, is in great demand. "Pre-emptive salvage" cutting, in which any remaining healthy trees are cut in the attempt to get full value for them before they become diseased, is common (at least in the Great Lakes states). The salvage of non-infected trees, however reduces the pool of potentially disease resistant individuals. Woodlot management is also a a form of threat to this species, since it may not involve providing the types of disturbance (e.g., soil disturbance) needed by this shade-intolerant species to successfully reproduce and establish new individuals. Unless management practices are altered to allow for open, disturbed areas, it is unlikely that there will be significant butternut reproduction (Skilling 1993).

The canker kills trees of all ages (Morin et al. 2018).

In many areas healthy butternut trees have been found growing adjacent to diseased trees and these presumably resistant trees have remained healthy over a 12 year period (Ostr and Woeste 2004).

Butternut canker disease can be identified by the following characteristics or conditions: (1) trees with dying branches or dead tops, and tufts of shoots (epicormic shoots) below the dead portions; (2) discolored bark that in spring exudes an inky-black, thin fluid from cracks in the cankers, and in summer has sooty bark patches that commonly have a white margin; and (3) cankers of various types. In early stages, cankers are sunken and elongate, originating at leaf scars, buds, and in various tree wounds. In latter stages, cankers are visible on older stem and branch portions, many loosely covered with shredded bark and bordered by successive callus layers. Infected trees are also characterized by dark brown to blackish wood portions in elliptical patterns beneath the bark. Dieback occurs through single or coalescing cankers girdling branches, and branch suckers and stem sprouts quickly become infected and die.

An additional threat is hybridization with Juglans ailantifolia (heartnut), a species from Japan which is marketed as a nut tree (McDaniel 1979).
Ecology & Habitat

Description

Juglans cinerea is a deciduous tree reaching nearly 100 ft. in height. Its bark is grayish-brown, with smooth ridges. The pith of the stems is transversely partitioned into chambers which are dark chocolate brown. The leaves are compound with 11-17 leaflets that have long, pointed tips. Leaflets from the middle of the leaf are the largest. The flowers are in dense, drooping catkins (resembling cats' tails) which protrude from the buds in autumn and lengthen in the spring. The nut resembles a short cylinder (1.5-2.75 inches long) with 2 halves and 2 or 4 ridges running end to end along its rough surface. Both the nut and the husk remain closed at maturity. Seed production begins when the tree has reached 20 years of age and peaks when it is 30-60 years old. Seeds are produced every year, but larger seed crops occur every 2-3 years. Germination of seeds usually takes place in the spring from seeds that fell the previous season. (Gleason and Cronquist 1991, Ostry et al. 1994).

Diagnostic Characteristics

There are some pubescence differences which will help to distinguish Butternut from the Black Walnut, even when the distinctive dark pith is not exposed. In J. cinerea, there is often a pad of dense small hairs extending transversely along the upper margin of the old leaf scars; in J. nigra, this pad is absent, although the circular area of bud pubescence is confusing, and some specimens are ambiguous. The underside of the leaflets in J. cinerea is +/- densely covered with mostly stellate hairs, while in J. nigra the pubescence is sparser and mostly of simple hairs. The pubescence of cinerea, including that on the fruit, is more clammy than that of J. nigra. Persons familiar with the species in the field will also know differences in fragrance of the foliage and fruit and in stain from the husks--all difficult to expresss in words (Voss, 1980).

Habitat

Juglans cineara typically grows in rich mesophytic forests, lower slopes, ravines, and various types of bottomland, including banks and terraces of creeks and streams, and floodplain forests (Fernald 1950, Gleason and Cronquist 1991). This species achieves its best growth in well-drained bottomland and floodplain soils.

The following provides additional information on the specific habitats of J. cinerea in several states from different parts of the species' range:

The habitat in Ohio is mesic river terraces and ravines (Cusick 1992).

Michigan habitat includes streambanks, swamp forests, and upland beech-sugar maple, oak-hickory, and mixed hardwood stands (Voss 1985). In southern Michigan sites, associates may include such species as Ulmus americana (American elm), Acer saccharinum (silver maple), Fraxinus quadrangulata (blue ash), Betula alleghaniensis (yellow birch), Tilia americana (basswood), and many woodland and swamp herbs. In some areas, butternut occurs occasionally as a roadside tree.

The habitat in Illinois and Wisconsin is mesic and riparian hardwood forests, where associates of the overstory include Acer saccharum (sugar maple), Carpinus caroliniana (blue beech), Celtis occidentalis (hackberry), Fagus grandifolia (American beech), Prunus serotina (black cherry), Quercus alba (white oak), Quercus macrocarpa (bur oak), Quercus rubra (red oak), Tilia americana (basswood), and Ulmus americana (American elm). The herbaceous associates of this habitat include Claytonia virginiana (spring beauty), Dicentra cucullaria (squirrel corn), Hydrophyllum appendiculatum (appendaged waterleaf), Isopyrum biternatum (false meadow-rue), and Solidago caesia (woodland goldenrod). In wooded floodplain habitats, associate plant species include Celtis occidentalis, Cryptotaenia canadensis (honewort), Elymus virginicus (Virginia rye grass), Fraxinus pennsylvanica (red ash), Galium aparine (cleavers), Geum canadense (geum), Hydrophyllum virginianum (Virginia waterleaf), Laportea canadensis (wood nettle), Ranunculus septentrionalis (swamp buttercup), Toxicodendron radicans (poison ivy), Rudbeckia laciniata (cutleaf coneflower), Ulmus americana, and Viola sororia (woolly blue violet). A characteristic habitat in the Chicago region is springy, wooded, calcareous slopes, where limey water percolates through gravelly soils. Associate plant species in this habitat may include Angelica atropurpurea (angelica), Caltha palustris (marsh marigold), Cardamine bulbosa (spring cress), Fraxinus nigra (black ash), Lilium michiganense (Michigan lily), Mitella diphylla (bishop's cap), Ribes americanum (swamp gooseberry), Smilacina stellata (starry false solomon's seal), Solidago patula (swamp goldenrod), and Symplocarpus foetidus (skunk cabbage) (Swink and Wilhelm 1979).

The habitat in Indiana is mesic forests (Homoya 1992).

The typical habitat in Kentucky is mesic wooded ravines along streams, mesic limestone gorges, mesic deciduous forest, second growth in mixed hemlock forest, and upland mesic woods, occurring with Fraxinus americana (white ash), and several other species (KY HP 1992).

In Tennessee, Juglans cinerea occurs along creek bottoms in mesic forests and on lower slopes (TN ESD 1992).

In Mississippi, this species occurs on calcareous bluffs, along river and stream drainages, in mesophytic forests, and calcareous ravines. Populations are often found on north-facing slopes. Associate plant species include Acer spp. (maple), Carya spp. (hickory), Juglans nigra (black walnut), Liriodendron tulipifera (tuliptree), and Quercus macrocarpa (bur oak) (MS NHP 1992).

Habitat in Delaware consists of northwest-facing slopes of pastures, banks of streams running through meadows, and floodplain woods with partial shade (DE NHP 1992).

In Massachusettes this species a generalist in terms of soil moisture (it's found along a continuum from river bottomlands to mesic forests to talus and dry balds) but it does not seem to be a generalist in terms of bedrock substrate-in MA it avoids acid, nutrient poor substrates and appears to be restricted to areas underlain by basalt, marble, or dolomite (M. Dow Cullina, pers. comm., 2006).

Ecology

Juglans cinerea achieves optimal growth on well-drained soils of bottomlands and floodplains, but rarely occurs in pure stands. According to Rink (1990), this species grows best in riparian sites and well-drained soils, but is seldom found on dry, compact, or infertile soils. Rink also notes that butternut is found at much higher elevations (up to 4900 ft) in the Virginias than black walnut. Butternut is shade-intolerant, growing best in full sunlight. Young trees can tolerate some competition from the side, but will not withstand shade from above. This species needs to be in the canopy in order to survive. Reproduction is successful only in areas where shade does not inhibit its growth, such as stand openings or in fields (Skilling 1993, Ostry et al. 1994).

This species is being seriously impacted, if not devastated, by a canker fungus (Sirococcus clavigignenti-juglandacearum) that is spreading rapidly throughout its range, and few stands remain uninfected. The origin of the disease is unknown, although Fogelson and Campbell (1990) believe it may have been introduced about 40 years ago on the east coast of the United States. According to Anderson (1993), 40- year-old cankers have been observed in North Carolina. Long noted as suffering from a canker dieback (frequently called "butternut decline"), scientists from Wisconsin first identified the disease agent as a new species of Sirococcus in 1967, although tests of trees in North Carolina demonstrated its presence there in 1952. This disease has spread quickly (within 30 years) throughout butternut's range, infecting and killing trees in all locations. In the Great Lakes states, all sites examined by researchers had been infected with canker. The disease vectors are poorly known, but the canker is believed to be transmitted at least in part by raindrops splashing spores from infected trees onto healthy ones, and possibly via insects. If the spores become airborne, they are able to be dispersed over great distances. The spores are produced throughout the growing season and can survive when weather conditions are cool and skies are overcast. (Kuntz et al. 1979, Fogelson and Campbell 1990, Forest Service News 1992, Ostry et al. 1994).

Symptoms of the disease include the appearance of lens-shaped cankers on the trunk, limbs, twigs, and immature nut of the tree. In spring, an inky-black, thin fluid is exuded from the cankers. These cankers usually continue to grow in size and girdle the tree, eventually killing it by destroying the cambium. The girdling of the tree often causes a wilting of the leaves, especially noticeable in the crown (Fogelson and Campbell 1990). Trees with reduced vigor become susceptible to insects and secondary disease agents, particularly Melanconis juglandis (E. and E.) Graves, which causes branch dieback (Skilling 1993).

Butternut is an important source of mast for wildlife, especially in the northern part of its range, where black walnut (Juglans nigra) does not occur. Squirrels and other rodents are some of the consumers of the seeds. (Ostry et al. 1994).

Reproduction

Male and female flowers borne seperately on the same tree. Squirrels, rabbits and white tailed deer eat the nuts. Squirrels disperse them.
Terrestrial Habitats
Forest/WoodlandWoodland - Mixed
Other Nations (2)
United StatesN3
ProvinceRankNative
IndianaS2Yes
District of ColumbiaS1Yes
South CarolinaS2Yes
IowaSUYes
WisconsinS2Yes
MaineSUYes
OhioS2Yes
ConnecticutSNRYes
AlabamaS1Yes
DelawareS3Yes
New HampshireS3Yes
MarylandS2Yes
MinnesotaS1Yes
MichiganS3Yes
New JerseyS1Yes
GeorgiaS2Yes
New YorkS4Yes
IllinoisS1Yes
NebraskaS1Yes
PennsylvaniaS4Yes
VermontS3Yes
VirginiaS3Yes
West VirginiaS2Yes
WashingtonSNANo
ArkansasS3Yes
MissouriSUYes
MassachusettsS4Yes
MississippiS2Yes
North CarolinaS2Yes
TennesseeS3Yes
KentuckyS2Yes
CanadaN2
ProvinceRankNative
QuebecS1Yes
ManitobaSNANo
New BrunswickS1Yes
OntarioS2Yes
Prince Edward IslandSNANo
Nova ScotiaSNANo
Threat Assessments
ThreatScopeSeverityTiming
8 - Invasive & other problematic species, genes & diseasesPervasive (71-100%)Serious or 31-70% pop. declineHigh (continuing)
8.5 - Viral/prion-induced diseasesPervasive (71-100%)Serious or 31-70% pop. declineHigh (continuing)
8.5.2 - Named "species" (disease)Pervasive (71-100%)Serious or 31-70% pop. declineHigh (continuing)

Plant Characteristics
DurationPERENNIAL, Long-lived, DECIDUOUS, SPRING-FLOWERING
Economic Value (Genus)No
Roadless Areas (18)
Kentucky (1)
AreaForestAcres
WolfpenDaniel Boone National Forest2,835
Missouri (1)
AreaForestAcres
Spring Creek Rare II Study AreaMark Twain National Forest4,899
North Carolina (3)
AreaForestAcres
Bald MountainPisgah National Forest11,085
Overflow CreekNantahala National Forest3,379
Tusquitee BaldNantahala National Forest13,670
Tennessee (1)
AreaForestAcres
Bald MountainCherokee National Forest11,743
Vermont (3)
AreaForestAcres
Bread LoafGreen Mountain and Finger Lakes National Forests1,768
Devil's Den 09083Green Mountain and Finger Lakes National Forests9,169
Griffith Lake 09084Green Mountain and Finger Lakes National Forests1,833
Virginia (4)
AreaForestAcres
Kelley MountainGeorge Washington National Forest7,590
Peters Mountain Addition A (VA)Jefferson National Forest1,268
Raccoon BranchJefferson National Forest4,388
Seng MountainJefferson National Forest6,428
West Virginia (5)
AreaForestAcres
Dolly Sods Roaring PlainMonongahela National Forest13,392
Middle MountainMonongahela National Forest19,020
North Mountain HopevilleMonongahela National Forest6,525
Peters Mountain Addition A (WV)Jefferson National Forest343
Seneca CreekMonongahela National Forest22,287
References (32)
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  2. Anderson, R. L. and L. A. LaMadeleine. 1978. The distribution of butternut decline in eastern United States. USDA Forest Service. Survey Report S-3-78. 5 pp.
  3. Brown, C.L., and L.K. Kirkman. 1990. Trees of Georgia and adjacent states. Timber Press, Inc., Portland, OR. 291 pp.
  4. Carlson, J. C. 1993. Insects and diseases. Butternut: are There any Healthy trees left? WCTPA Quarterly Journal April: 28-31.
  5. Cusick, A. 1992. Ohio Natural Heritage Program. Personal communication with M. Penskar, MI NFI: ESA questionnaire for Juglans cinerea. 4 pp.
  6. Dodds, J.S. 2024. <i>Juglans cinerea</i> Rare Plant Profile. New Jersey Department of Environmental Protection, State Parks, Forests & Historic Sites, Forests & Natural Lands, Office of Natural Lands Management, New Jersey Natural Heritage Program, Trenton, NJ. 20 pp. [https://www.nj.gov/dep/parksandforests/natural/heritage/docs/juglans-cinerea-butternut.pdf]
  7. Fernald, M. L. 1950. Gray's manual of botany. 8th edition. Corrected printing (1970). D. Van Nostrand Company, New York. 1632 pp.
  8. Gleason, H.A., and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. New York Botanical Garden, Bronx, New York. 910 pp.
  9. Homoya, M. 1992. Ecologist/Botanist, Indiana Natural Heritage Data Center. Personal communication with M. Penskar, MI NFI: ESA questionnaire for Juglans cinerea. 4 pp.
  10. Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
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  12. Little, E.L., Jr. 1979. Checklist of United States trees (native and naturalized). Agriculture Handbook No. 541. U.S. Forest Service, Washington, D.C. 375 pp.
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  15. Morin, R., K. Gottschalk, M. Ostry, and A. Liebhold. 2018. Regional patterns of declining <i> (</i><i>Juglans cinerea </i>L.) suggest site characteristics for restoration. Ecology and Evolution 8: 546-559.
  16. Nicholls, T. H., K. J. Kessler, Jr., and J. E. Kuntz. 1978. How to identify butternut canker. U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station, St. Paul, MN. (Pamphlet).
  17. Ostry, M.E. and K.Woeste. 2004. Spread of butternut canker in North America, host range, evidence of resistance within butternut populations and conservation genetics. <i>In </i>Black Walnut in a New Century: Proc. 6th Walnut Council Research Symposium. Eds. C.H. Michler, P.M. Pijut, J. Van Sambeek, M. Coggeshall, J. Seifert, K. Woeste, R. Overton, F. Ponder. USDA For. Serv. North Central Res. Stn.Gen. Tech. Rep. NC-243, St. Paul, MN.
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  19. Penskar, Mike. Personal communication. Botanist, Michigan Natural Features Inventory. Lansing, MI.
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  22. Rock, J. 1992. Botanist, Great Smoky Mountains National Park. Personal communication with M. Penskar, MI NFI: ESA questionnaire for Juglans cinerea. 4 pp.
  23. Rose, G. A. 1992. Butternut harvest moratorium. Division of Forestry, Minnesota Department of Natural Resources. 1 p.
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  28. Skilling, D. D. North Central Forest Experiment Station.
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