A series of studies by Patrick J. Engelken, M. Eric Benbow, Deborah G. McCullough, Nate Siegert, Randall Kolka, Melissa Youngquist and others examine the status of ash (Fraxinus spp.) in the aftermath of the emerald ash borer (EAB) invasion. Initial studies documented the crash of biomass supporting EAB numbers when the large ash trees died (Siegert, Engelken, McCullough. 2021; full citation at end of blog.) More recent studies have focused on bogs and forests in the riparian areas where ash were especially numerous and arguably ecologically most important. I posted a blog about black ash bogs earlier.
I will focus here on the studies in riparian areas of southern and northwest Michigan. Information about impacts in forests of southern Michigan are from Engelken, Benbow and McCullough (2020); information about impacts in northwest Michigan are from Engelken and McCullough (2020). Full citations for both are at the end of the blog.
All study areas had high ash densities before EAB’s arrival. One study (Engelken and McCullough 2020) found ash densities high in the immediate riparian areas (in one case, a strip reaching 100 meters from the streambank) but scattered in surrounding forests.
In all these study areas, populations of mature (reproductive age) ash crashed within 10-15 years after EAB invasion:
- In northeast Michigan, EAB carrying capacity was reduced by 94% – 99%; total ash basal area was reduced by 87 – 97.7% (Siegert, Engelken, McCullough. 2021);
- In southern Michigan, more than 85% of the basal area of green (F. pennsylvanica) and black ash (F. nigra) had been killed by 2020. An estimated 96% of the overstory ash phloem area had died, thus radically reducing EAB carrying capacity (Engelken, Benbow and McCullough 2020);
- In northwest Michigan, more than 95% of the overstory ash have been killed. (Engelken and McCullough 2020).
The worst impact has been on black ash– which plays such an important ecological role in riparian areas and wetlands and has enormous importance in Native American cultures. In all these study areas, there is no stump sprouting by black ash (Siegert, Engelken, McCullough. 2021; Engelken, Benbow and McCullough 2020; Engelken and McCullough 2020). In three watersheds of northwest Michigan where black ash constituted up to a quarter of the overstory species before the EAB invasion, scientists found no black ash recruits, only eight saplings, and a single seedling.
Green ash (F. pennsylvanica) has survived in much higher numbers – so far. However, this species’ ability to grow into reproductive size is still uncertain. In northwest Michigan, green ash saplings are abundant in canopy gaps created by EAB-caused mortality of mature ash. These saplings had established before the EAB invasion so some call them the “orphaned cohort”. However, there are few seedlings of any woody plant species in these gaps because sedges form such dense mats.
Green ash reproduction faces many challenges before persistence of the species can be considered assured. First, populations of EAB – now reduced by the lack of mature ash to support them – might resurge when young ash grow to larger sizes. It is not yet clear the extent to which introduced biocontrol agents and native predators, e.g., woodpeckers, will protect these trees as they grow to reproductive size. Here, again, green ash has an advantage over black ash. While green ash produce seed at a relatively young age, black ash don’t produce seed until they reach 30–40 years. Even then, they produce seeds only sporadically, with intervals of five or more years.
A second challenge is the lack of seed sources – at least until and unless young trees are able to reach reproductive size.
A third challenge is competition for resources from other plants. The canopy gaps eliminate competition for light for the taller plants, i.e., the existing ash saplings. However, the sapling cohort is not supported by a seedling cohort. There are very few seedlings of all woody plant species (including invasive species!). Seed germination is suppressed by the dense mats of wetland-adapted sedges and possibly the higher water tables (which resulted from reduced evapotranspiration following mortality of the mature trees).
Competition for resources is also a factor in the forests outside the immediate riparian zone. There, ash seedlings sprout, but shade created by lateral ingrowth suppresses their growth. In southern Michigan, Engelken, Benbow and McCullough (2020) note that the forests are apparently transitioning from red oak dominated forests to red maple and black cherry dominated forests. This transition is apparently intensified by forest mesophication resulting from reduced fire frequency, decreasing light availability in forest understories and increasing soil moisture content.
Fourth, while stump sprouting of green ash was noted in southern Michigan, in the northwestern forests all the sprouts died. I have already noted the absence of stump sprouting by black ash at all sites.
Beaver & Green Ash in Northern Virginia
In spring 2022 I noticed along one stream in northern Virginia that beavers had cut down green ash saplings; McCullough and Siegert report that this does not appear to be a problem in their study areas.
By December 2022, the beaver-cut trees tried to recover: see the sprouts from a stump [below]. (I think deer or rabbits ate the tips of the sprouts.)
The beavers also continued feeding on the ash — the tree photographed in the spring when it was half-chewed through has now been felled and its branches removed [see below].
Ecosystem Impacts, Especially on Streams
Across much of the upper Midwest, massive ash mortality is causing widespread changes in forest systems.
Riparian forests, i.e., areas adjacent to waterways where periodic inundation occurs, are functionally linked to the aquatic systems. Loss of such a significant proportion of the overstory changes the transfer of energy to adjacent waterways that takes the form of inputs of nutrients from leaf litter and coarse woody debris. Intact forests also stabilize stream banks and maintain channel depth by preventing erosion. Forests moderate temperature of the water. Finally, forests with “coarse woody debris” increase habitat structure. These impacts might be especially important along first order streams, (defined as perennial streams that have no permanently flowing tributaries). These streams are too small to buffer the impacts of major tree loss. The scientists say they are uncertain whether these changes continue to affect larger streams downstream.
Unshaded streams have higher water temperatures that can affect populations of fish, in particular salmonids, by delaying migration, reducing egg viability and increasing egg mortality. Higher temperatures can also alter primary productivity of aquatic algae, potentially increasing eutrophication (Engelken and McCullough 2020).
The scientists expect increasing abundance of coarse woody debris in the forests and streams of northwest Michigan as the 75% of dead ash that are still standing fall. Such debris provides nutrients and habitat for an array of plants and animals, thereby influencing the abundance, activity and species compositions of several ground dwelling insects and seedling establishment. In streams, coarse woody debris provides complex habitat and refuges. It also retains organic matter. Recreationists do find that debris impedes boating.
Loss of ash specifically
As described by Engelken, Benbow and McCullough (2020), and in my earlier blog, ash leaf litter – particularly black ash leaf litter – is highly nutritious. Ash leaf litter has efficient turnover rates and contributes important soil nutrients such as nitrogen, organic carbon and exchangeable cations. Invertebrate communities in headwater streams feed largely on coarse organic material such as leaf litter (Engelken and McCullough 2020). Consequently, loss of the annual influx of ash leaf litter will likely have adverse effects on nutrient availability in riparian forests and adjacent streams.
SOURCES
Engelken, P.J., M.E. Benbow, D.G. McCullough. 2020. Legacy effects of emerald ash borer on riparian forest vegetation and structure. Forest Ecology and Management 457 (2020) 117684
Engelken, P.J. and D.G. McCullough. 2020. Riparian Forest Conditions Along Three Northern Michigan Rivers Following Emerald Ash Borer Invasion. Canadian Journal of Forest Research.
Siegert, N.W., P.J. Engelken, D.G. McCullough. 2021 Changes in demography and carrying capacity of green ash and black ash ten years after emerald ash borer invasion of two ash-dominant forests. Forest Ecology and Management Vol 494, August 2021
Posted by Faith Campbell
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For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
This is a nice summary of a body of research. Regarding the uniqueness of ash, studies by grad student Liz Studer at Hubbard Brook, NH have shown that some spring wildflowers (notably painted trillium) occur preferentially under white ash compared to neighboring trees like maple, beech and birch. This is probably because ash leafs out later than the other species, providing a longer growing window for these understory plants.
Gary – very interesting. + painted trillium is well worth promoting!