As we all know, the emerald ash borer (EAB) has killed millions of ash trees in its invaded range across eastern North America. However, field studies have detected robust regeneration of ash seedlings and saplings in various invaded areas. Ward et al. 2021 (full citation at end of blog) set out to determine whether this regeneration will result in recovery of mature ashes that can perform their ecological role. They conclude that it will not. Instead, they say, the EAB invasion will probably alter successional patterns and composition of large areas of naturally regenerating forests, causing a cascade of ecological impacts in ash-containing ecosystems
Ward and colleagues used USDA Forest Service Forest Inventory and Analysis (FIA data) to quantify ash recruitment and regeneration across the entire eastern United States. Theirs is the first study to evaluate trends across the region, rather than specific locations or stands. They related the FIA recruitment data to EAB spread, as measured by USDA Animal and Plant Health Inspection Service’ (APHIS) record of the first EAB detection in each county.
FIA inventories in 2002-2007 and 2013-2018 show large numbers of ash seedlings and saplings in counties invaded in the first wave of invasion, 2002–2006. These areas had higher densities of both seedlings and saplings than plots in other counties. The earliest-invaded counties were in areas that had extraordinarily high densities of ash before the EAB invasion, so the numbers of seedlings and saplings probably reflected that abundant seed source.
However, by the 2013-2018 inventory ash trees in the smallest overstory class (12.7 cm dbh) were dying at faster rates than they were recruited from seedlings or saplings in all 362 counties recorded by APHIS as EAB-infested before 2013. Ward and colleagues found these negative population trajectories on plots that have been invaded for more than about 10 years. This trend suggests that ash will continue to decline in abundance and may become functionally extinct across the invaded range.
Some U.S. Forest Service biologists are more optimistic about ash recovery in response to biocontrol of the EAB. See their podcast here.
In the risk of functional extinction, ash trees are unfortunately not unique. The authors note similar impacts from the invasion of the hemlock woolly adelgid and beech bark disease.
Data Reveal History of Invasion (spread)
Ward and colleagues focused on the risk of mortality for young ashes as they developed from seedlings to saplings, and, eventually, to overstory trees. The youngest “overstory” trees are 12.7 to 17 cm dbh. FIA data show that even the largest trees in this class are 3 cm smaller than trees that produce seeds.
Mortality was initially uniformly low – less than 2.1% — as measured by the first FIA inventory (2002–2007). This is not surprising because EAB was detected only in 2002, and then in only few counties. (EAB had probably been present for a decade before it was detected.)
By the 2013-2018 FIA inventory, mortality had quadrupled to 8–11% in counties invaded during the 2002–2006 period. In the counties invaded during the 2007–2012 period, morality also rose to 3-5%. Both measurements included all diameter classes. Annual mortality rates in the FIA 2013-2018 inventory were still highest for the counties invaded during 2002–2006 except for the largest trees (those greater than 40 cm dbh). By the time of the 2013-2018 FIA survey, overstory ash densities near the epicenter had since declined substantially. They had been nearly eliminated in some counties in southeastern Michigan. There were still sufficient numbers of smaller trees in the region to exhibit an elevated mortality rate – more than 10% per year in several counties in Michigan, Indian, and Ohio. By contrast, in the most recently invaded areas – those counties recorded by APHIS as infested after 2013 – there was very little change in ash densities compared to the 2002-2007 period. This is hardly surprising since it takes years for mortality to reach levels observable by the FIA process.
Considering trees just entering the overstory category (those with diameters of 12.7 cm dbh), annual mortality increased substantially across the region. Between the first FIA inventory (conducted in 2002-2007) and the second inventory (conducted in 2013-2018), their average annual mortality rose more than four-fold, from 0.08 trees per ha to 0.37 trees per ha. By 2013-2018, recruitment in the 2002–2006 invasion cohort was about 50% less than tree mortality levels; recruitment and mortality were about equivalent for the counties invaded in the 2007–2012 period. Recruitment was [still] significantly higher than mortality for the counties recorded as invaded in 2013–2018. However, Ward and colleagues expect mortality rates of this cohort to accelerate over the next five to 10 years – even in areas with lower ash densities.
Ward and colleagues note that many of the young ash trees were dying before they could reach reproductive age – which they estimated to be about 20 years with a dbh of about 20 cm.
As the invasion progresses and hosts are depleted, mortality rates could slow, but, for ash to persist, it is critical that sufficient numbers of trees reach reproductive age before succumbing to residual EAB populations.
Other factors that might influence ash include competition with trees in other genera. The biocontrol agents now becoming established in young ash forests might increase the likelihood of ash persistence. Still, seed production and seedling survival will need to be frequent and widespread if they are to offset expected mortality. Resilience might also vary depending on individual species’ vulnerability to changes in the climate and to EAB (green and black ash are more vulnerable than white ash).
SOURCE
Ward, S.F., A.M. Liebhold, R.S. Morin, S. Fei. 2021. Population dynamics of ash across the eastern USA following invasion by emerald ash borer. Forest Ecology and Management 479 (2021) 118574
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