Black Ash Ecology

Black ash grows in a variety of forest types across its range. In the northeastern U.S., black ash is typically found in smaller pockets of riparian forests, ephemerally wet forests, and swamps. In this region, black ash is not always the dominant tree species and may co-occur with hemlock, yellow birch, balsam fir, red maple, American basswood, and balsam poplar

Similar stands can occur in the St. Lawrence River region of New York and in northern portions of the Great Lakes states. In these areas, however, black ash can dominate wetland forests that range in size from a few acres to hundreds of acres. Associated tree species in these regions include red maple, American basswood, yellow birch, balsam fir, balsam poplar, and hemlock.1

A wet black ash site in early spring. St. Lawrence County, NY. Photo by River Mathieu, MSU.

Many tree species are not able to establish, survive, or grow in areas that are frequently or continually flooded or areas where soils remain wet for much of the year. These areas tend to be dominated by herbaceous plants or sedges that can tolerate flooded conditions and low soil oxygen levels.1,2 Black ash has evolved several adaptations that allow the species to occupy these otherwise harsh conditions. 2,3,4,5 For example, soils in forested wetlands are typically shallow and perched above high-water tables. Black ash can establish sturdy root structures that are relatively shallow and closer to the soil surface.1 Hypertrophied lenticles and adventitious roots are adaptations that allow black ash, and other bottomland woody species, to cope with the low levels of oxygen common in soils in swampy or boggy areas.6

A wetland dominated with herbaceous plants, grasses, and sedges on the edge of a black ash stand in northern NY. Note the dead black ash trees present in the image that likely died due to the sites very high-water table. EAB is not yet present here. Photo by River Mathieu, MSU

Growth rates of black ash trees vary, depending on the site conditions and competition with other tree species. Black ash is intolerant of shade and suppressed trees that are less than 10 cm in diameter may be as much as 60-80 years old. 7 In more productive sites, black ash trees exposed to full or nearly full sun can grow relatively quickly.

Black ash seeds from a mature tree in Vermontville, MI. Photo by Tim Harrison, MSU.

Black ash trees may begin producing seeds when they are 30 to 40 years in age.1 Seeds mature by late summer and drop to the forest floor in autumn. Black ash reportedly produces large seed crops at 5-8-year intervals. Some reports indicate viable seeds can persist in the seedbank for up to 3 or 4 years.2,8 Recent research however, found little or no persistence of a black ash seedbank hydric forest sites dominated by black ash.9

Black ash regeneration, which refers to seedlings, saplings, and the larger recruits, often comprises a large component of the understory in black ash forests.10 Recruits, typically 1-4 inches in diameter, are young trees that are likely to eventually become overstory trees. Saplings, generally < 1 inch in diameter, are often moreabundant than recruits. Saplings can become recruits if they survive deer browse, competition, shade, ice, or other damage. Densities of ash seedlings can be high in some black ash stands; particularly where ample amounts of sunlight reach the ground. In other stands that are frequently flooded or deeply shaded, black ash seedlings may be absent or present at relatively low densities.

Dense black ash seedlings growing in a stand with ample black ash regeneration, characteristic of healthy black ash sites. Photo by Patrick Engelken, USFS

Like all ash species, black ash is ring porous, a term that refers to the way that trees transport water and nutrients from the roots up the trunk and into canopy branches and leaves. In ring porous trees, all or nearly all of this transport occurs in the outer ring of growth. Each annual ring is formed by the combination of earlywood and latewood cells in the xylem. In spring, black ash trees begin forming earlywood vessels in the xylem. These earlywood cells have large lumens (the opening in the middle) and are loosely packed together. This facilitates transport of relatively large volumes of water to the growing leaves and shoots. In mid-summer, trees stop producing earlywood and begin producing latewood. Latewood vessels are smaller, have thicker cell walls and cells are more tightly packed together than earlywood. Together, the earlywood and latewood comprise a tree ring, which is one year's worth of radial growth. Black ash trees have notably large vessels, especially in earlywood tissue. This contributes to the pliable nature of the splints that are used for weaving baskets. 8,10,11,12

Early wood and latewood shown on a black ash tree core. Photo by River Mathieu, MSU

The impact of EAB on black ash is likely to be profound and could lead to the functional loss of black ash as a major overstory species in wetland forests. Larvae of EAB feed on phloem in galleries that are often serpentine and that increase in width as the larvae grow. Phloem is the inner bark that transports carbohydrates produced by the leaves down the trunk to the roots. In black ash, these larval galleries usually extend further horizontally than on other ash species. This means that each larva injures more tissue and fewer larvae are required to cause mortality of black ash compared with other ash species.13, 14 This pattern, combined with the harsh site conditions where most black ash tree occur, can lead to complete or nearly complete mortality of overstory black ash trees, sometimes within 4-6 years of invasion. In forests dominated by black ash, ecological effects of this mortality may be substantial. For example, in autumn, black ash leaves provide an input of high-quality leaf litter, which drives nutrient cycles.15 Loss of overstory black ash trees is likely to affect site hydrology,16,17 indirectly impacting other plant species.5 Habitat available for various vertebrate and invertebrate animals in these systems is also affected by mortality of overstory black ash.15

A Michigan State University researcher assessing a black ash site in the southern UP, Michigan. Photo by River Mathieu, MSU

Many questions are yet to be answered about the indirect and cascading effects of EAB invasion on black ash ecosystems. Ongoing research will shed light on how young black ash fare in sites where older trees have been killed. Whether black ash regeneration can co-exist with low densities of EAB and survive long enough to reach the overstory remains unknown. Potential competition between young black ash and co-occurring tree species will also influence the long-term persistence of black ash.

Page References

  1. Gucker, C. L. 2005. Fraxinus nigra. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer).
  2. Wright, J. ., & Rauscher, M. H. (1990). Fraxinus nigra Marsh. Silvics of North America, Hardwoods. USDA For. Serv. Agriculture Handbook 654, vol 2, pp. 344-347.
  3. Erdmann, G. G., Crow, T. R., Peterson, R. M., & Wilson, C. D. 1987. Managing black ash in the Lake States. USDA Forest Service
  4. COSEWIC. 2018. Committee on the Status of Endangered Wildlife in Canada assessment and status report on the black ash Fraxinus nigra in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. xii + 95 pp. (Species at Risk Public Registry).
  5. Shannon, J., Van Grinsven, M., Davis, J., Bolton, N., Noh, N., Pypker, T., et al. 2018. Water level controls on sap flux of canopy species in black ash wetlands. Forests. 9(3). 147. doi:10.3390/f9030147
  6. Havens, K. J., & Virginia Institute of Marine Science, Wetlands Program. (1996) Plant Adaptations to Saturated Soils and the Formation of Hypertrophied Lenticels and Adventitious Roots in Woody Species. Wetlands Program Technical Report no. 96-2. Virginia Institute of Marine Science, College of William and Mary.
  7. Windmuller-Campione, M. A., Russell, M. B., Slesak, R. A., & Lochner, M. (2020). Regeneration responses in black ash (Fraxinus nigra) wetlands: Implications for forest diversification to address emerald ash borer (Agrilus planipennis). New For., 52(4), 537-558. doi:10.1007/s11056-020-09807-0
  8. Benedict, L., & David R. 2004. Handbook for black ash preservation reforestation/regeneration. Mohawk Council of Akwesasne, Department of the Environment.
  9. Klooster, W. S., Herms, D. A., Knight, K. S., Herms, C. P., McCullough, D. G., Smith, A., Gandhi, K. J. K., & Cardina, J. (2014). Ash (Fraxinus spp.) mortality, regeneration, and seed bank dynamics in mixed hardwood forests following invasion by emerald ash borer (Agrilus planipennis). Biol. Invasions, 16(4), 859-873.
  10. Benedict, M. A., & Frelich, L. E. 2008. Site factors affecting black ash ring growth in northern Minnesota. For. Ecol. Manage., 255(8), 3489-3493. doi:10.1016/j.foreco.2008.02.029
  11. Costanza, K. K. L., Livingston, W. H., Kashian, D. M., Slesak, R. A., Tardif, J. C., Dech, J. P., et al. 2017. The precarious state of a cultural keystone species: Tribal and biological assessments of the role and future of black ash. J. For. 115(5). 435-446. doi:10.5849/jof.2016-034R1
  12. Diamond, A. K., & Emery, M. R. 2011. Black ash (Fraxinus nigra Marsh.): Local ecological knowledge of site characteristics and morphology associated with basket-grade specimens in New England (USA). Econ. Bot. 65(4). 422-426. doi:10.1007/s12231-011-9174-z
  13. Herms, D. A., & McCullough, D. G. 2014. Emerald ash borer invasion of North America: History, biology, ecology, impacts, and management. Ann. Rev. of Entomol. 59: 13-30. doi:10.1146/annurev-ento-011613-162051.
  14. McCullough, D. G. 2019. Challenges, tactics, and integrated management of emerald ash borer in North America. Forestry, 93: 197-211. doi:10.1093/forestry/cpz049.
  15. Kolka, R. K., D'Amato, A. W., Wagenbrenner, J. W., Slesak, R. A., Pypker, T. G., Youngquist, M. B., Grinde, A. R., Palik, B. J. 2018. Review of ecosystem level impacts of emerald ash borer on black ash wetlands: What does the future hold? Forests 9: 179. doi:10.3390/f9040179.
  16. Palik, B. J., D'Amato, A. W., & Slesak, R. A. 2021. Wide-spread vulnerability of black ash (Fraxinus nigra marsh.) wetlands in Minnesota USA to loss of tree dominance from invasive emerald ash borer. Forestry. 94(3). 455-463. doi:10.1093/forestry/cpaa047
  17. Slesak, R. A., Lenhart, C. F., Brooks, K. N., D'Amato, A. W., Palik, B. J. 2014. Water table response to harvesting and simulated emerald ash borer mortality in black ash wetlands in Minnesota, USA. Can. J. For. Res. 44: 961-968.

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