12
forest ecology The Precarious State of a Cultural Keystone Species: Tribal and Biological Assessments of the Role and Future of Black Ash Kara K.L. Costanza, William H. Livingston, Daniel M. Kashian, Robert A. Slesak, Jacques C. Tardif, Jeffrey P. Dech, Allaire K. Diamond, John J. Daigle, Darren J. Ranco, Jennifer S. Neptune, Les Benedict, Shawn R. Fraver, Michael Reinikainen, and Nathan W. Siegert Black ash (Fraxinus nigra Marsh.) plays a central role in several Native American teachings (including a Wabanaki creation story) and has long been used for basketry, yet relatively little is known about the species’ ecology. The recent and ongoing invasion of emerald ash borer (Agrilus planipennis Fairmaire), an invasive beetle killing millions of ash trees in eastern North America, threatens the future of black ash and the centuries-old basketry tradition. In recognition of the precarious state of this cultural keystone species, basketmakers, basket-tree harvesters, and researchers assembled to discuss traditional ecological knowledge and research advancements related to black ash. Here we provide an overview of basket-quality ash, synthesize current knowledge of black ash biology and ecology, and report findings from this successful tribal and scientific collaboration. Management recommendations were developed and future research needs outlined in hopes of sustaining an ecologically important tree species and maintaining a Native American tradition that has cultural and spiritual significance. Keywords: Agrilus planipennis, emerald ash borer, Fraxinus nigra, basketry, invasive forest pest, traditional ecological knowledge B lack ash (Fraxinus nigra Marsh.) is a unique and significant tree species from ecological, economic, and cultural perspectives. The species occupies a particular ecological niche in forested wetlands where few other woody plants thrive. Ecosystems containing black ash stands also play an important role in sup- porting other unique species, including flooded jellyskin (Leptogium rivulare [Ach.] Mont.), a threatened arboreal lichen that preferentially occurs on the basal bark of mature black ash (Lee 2004). In addition, Native American and European American basketmakers derive socioeconomic benefit Received August 5, 2016; accepted December 22, 2016; published online February 9, 2017. Affiliations: Kara K.L. Costanza ([email protected]), School of Forest Resources, University of Maine, Orono, ME. William H. Livingston ([email protected]), School of Forest Resources, University of Maine. Daniel M. Kashian ([email protected]), Wayne State University. Robert A. Slesak ([email protected]), Minnesota Forest Resources Council. Jacques C. Tardif ([email protected]), University of Winnipeg. Jeffrey P. Dech ([email protected]), Nipissing University. Allaire K. Diamond ([email protected]), Vermont Land Trust. John J. Daigle ([email protected]), Member of the Penobscot Indian Nation and School of Forest Resources, University of Maine. Darren J. Ranco ([email protected]), Member of the Penobscot Indian Nation and Department of Anthropology, University of Maine. Jennifer S. Neptune ([email protected]), Member of the Penobscot Indian Nation and Maine Indian Basketmakers Alliance. Les Benedict ([email protected]), Member of the Saint Regis Mohawk Tribe and Akwesasne Task Force on the Environment. Shawn R. Fraver ([email protected]), School of Forest Resources, University of Maine. Michael Reinikainen ([email protected]), University of Minnesota. Nathan W. Siegert ([email protected]), USDA Forest Service. Acknowledgments: We appreciate the traditional ecological knowledge regarding black ash basketry and basket-quality ash that was generously shared by Gabriel Frey (Passamaquoddy Tribe, Indian Township), Jeremy Frey (Passamaquoddy Tribe, Indian Township), Eldon Hanning (Micmac Tribe), Frank Hanning (Micmac Tribe), Gerald “Butch” Jacobs (Passamaquoddy Tribe, Pleasant Point), Peter Neptune (Passamaquoddy Tribe, Pleasant Point), Richard Silliboy (Micmac Tribe), Fred Tomah (Maliseet Tribe), and Richard David (Saint Regis Mohawk Tribe). Photos were taken by Sheridan Adams and Knife Edge Productions, Nathan W. Siegert, and Kara K.L. Costanza. This work was supported by the USDA Forest Service, Northeastern Area State and Private Forestry, Forest Health Protection, and by the National Science Foundation (Award EPS-0904155 to Maine EPSCoR at the University of Maine), and by the USDA National Institute of Food and Agriculture, McIntire–Stennis project number #ME0-M-8-00501-12 through the Maine Agricultural and Forest Experiment Station. Maine Agricultural and Forest Experiment Station Publication Number 3519. REVIEW ARTICLE Journal of Forestry • September 2017 435 J. For. 115(5):435– 446 https://doi.org/10.5849/jof.2016-034R1 Copyright © 2017 Society of American Foresters Downloaded from https://academic.oup.com/jof/article-abstract/115/5/435/4599906 by guest on 18 July 2019

The Precarious State of a Cultural Keystone Species

  • Upload
    others

  • View
    2

  • Download
    0

Embed Size (px)

Citation preview

forest ecology

The Precarious State of a Cultural KeystoneSpecies Tribal and Biological Assessments ofthe Role and Future of Black AshKara KL Costanza William H Livingston Daniel M KashianRobert A Slesak Jacques C Tardif Jeffrey P DechAllaire K Diamond John J Daigle Darren J RancoJennifer S Neptune Les Benedict Shawn R FraverMichael Reinikainen and Nathan W Siegert

Black ash (Fraxinus nigra Marsh) plays a central role in several Native American teachings (including aWabanaki creation story) and has long been used for basketry yet relatively little is known about the speciesrsquoecology The recent and ongoing invasion of emerald ash borer (Agrilus planipennis Fairmaire) an invasivebeetle killing millions of ash trees in eastern North America threatens the future of black ash and thecenturies-old basketry tradition In recognition of the precarious state of this cultural keystone speciesbasketmakers basket-tree harvesters and researchers assembled to discuss traditional ecological knowledge andresearch advancements related to black ash Here we provide an overview of basket-quality ash synthesizecurrent knowledge of black ash biology and ecology and report findings from this successful tribal and scientificcollaboration Management recommendations were developed and future research needs outlined in hopes ofsustaining an ecologically important tree species and maintaining a Native American tradition that has culturaland spiritual significance

Keywords Agrilus planipennis emerald ash borer Fraxinus nigra basketry invasive forest pest traditionalecological knowledge

B lack ash (Fraxinus nigra Marsh) is aunique and significant tree speciesfrom ecological economic and

cultural perspectives The species occupiesa particular ecological niche in forestedwetlands where few other woody plantsthrive Ecosystems containing black ashstands also play an important role in sup-porting other unique species includingflooded jellyskin (Leptogium rivulare [Ach]Mont) a threatened arboreal lichen thatpreferentially occurs on the basal bark ofmature black ash (Lee 2004) In additionNative American and European Americanbasketmakers derive socioeconomic benefit

Received August 5 2016 accepted December 22 2016 published online February 9 2017

Affiliations Kara KL Costanza (KaraLorionmaineedu) School of Forest Resources University of Maine Orono ME William H Livingston(williamlmaineedu) School of Forest Resources University of Maine Daniel M Kashian (dkashwayneedu) Wayne State University Robert A Slesak(raslesakumnedu) Minnesota Forest Resources Council Jacques C Tardif (jtardifuwinnipegca) University of Winnipeg Jeffrey P Dech(jeffreydnipissinguca) Nipissing University Allaire K Diamond (allairediamondgmailcom) Vermont Land Trust John J Daigle (jdaiglemaineedu)Member of the Penobscot Indian Nation and School of Forest Resources University of Maine Darren J Ranco (darren_rancoumitmaineedu) Member of thePenobscot Indian Nation and Department of Anthropology University of Maine Jennifer S Neptune (jennifermibaaolcom) Member of the Penobscot IndianNation and Maine Indian Basketmakers Alliance Les Benedict (lesbenedictsrmt-nsngov) Member of the Saint Regis Mohawk Tribe and Akwesasne Task Forceon the Environment Shawn R Fraver (shawnfravermaineedu) School of Forest Resources University of Maine Michael Reinikainen (rein0331umnedu)University of Minnesota Nathan W Siegert (nwsiegertfsfedus) USDA Forest Service

Acknowledgments We appreciate the traditional ecological knowledge regarding black ash basketry and basket-quality ash that was generously shared by Gabriel Frey(Passamaquoddy Tribe Indian Township) Jeremy Frey (Passamaquoddy Tribe Indian Township) Eldon Hanning (Micmac Tribe) Frank Hanning (MicmacTribe) Gerald ldquoButchrdquo Jacobs (Passamaquoddy Tribe Pleasant Point) Peter Neptune (Passamaquoddy Tribe Pleasant Point) Richard Silliboy (Micmac Tribe)Fred Tomah (Maliseet Tribe) and Richard David (Saint Regis Mohawk Tribe) Photos were taken by Sheridan Adams and Knife Edge Productions Nathan WSiegert and Kara KL Costanza This work was supported by the USDA Forest Service Northeastern Area State and Private Forestry Forest Health Protection andby the National Science Foundation (Award EPS-0904155 to Maine EPSCoR at the University of Maine) and by the USDA National Institute of Foodand Agriculture McIntirendashStennis project number ME0-M-8-00501-12 through the Maine Agricultural and Forest Experiment Station Maine Agricultural andForest Experiment Station Publication Number 3519

REVIEW ARTICLE

Journal of Forestry bull September 2017 435

J For 115(5)435ndash446httpsdoiorg105849jof2016-034R1

Copyright copy 2017 Society of American Foresters

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

from black ash via a variety of basketmakingtraditions

Like many of the Native Americans andFirst Nations people in the native range ofblack ash the Wabanaki tribes in Maine(Aroostook Band of Micmac Indians Houl-ton Band of Maliseet Indians Passamaquo-ddy Tribe and Penobscot Indian Nation)and the Saint Regis Mohawk Tribe in NewYork and Canada have a long history of har-vesting black ash for basketry (Benedict andFrelich 2008 Diamond and Emery 2011)Specific wood qualities such as strength andflexibility make black ash unique and ex-plain in part why it is highly preferred forbasketry (Benedict and David 2004) Yet thespeciesrsquo cultural significance goes beyondweaving baskets black ash is at the spiritualcenter of the tribesrsquo teachings including oneof their creation stories (Leland 1884 Bene-dict and David 2004) Many basketmakersrely on ldquobasket-tree harvestersrdquo these indi-viduals are tribal members who have learnedto identify favorable sites and recognize in-dividual black ash trees that are likely to pro-duce basket-quality materials As such theyare also responsible for maintaining andpassing on tribal knowledge about the spe-cies from generation to generation

Emerald ash borer (EAB) (Agrilus pla-nipennis Fairmaire) an invasive insect nativeto Asia that feeds on ash (Fraxinus spp) hascaused the Wabanakis Mohawks and oth-ers great concern about the future of blackash Initially detected in southeastern Mich-igan and southern Ontario in 2002 the EABinvasion has spread rapidly (Siegert et al2014b) and is responsible for killing mil-lions of North American ash trees (Cappaertet al 2005 Herms and McCullough 2014)As of July 2016 EAB has been detected in27 states and the Canadian provinces of On-tario and Quebec1 Although black ashstands in many locations still have valuablebasket-quality trees including those inMaine where to date no EAB detectionshave occurred the EAB invasion nonethe-less continues to spread and threaten whatremains of this unique resource (Siegert et al2015)

In addition to its significance as a cul-tural resource black ash has commercialvalue and can be used for veneer cabinetsand furniture (Ross 2010) Yet relatively lit-tle is known about this important species(Wright and Rauscher 1990) and the scien-tific literature on black ash biology and ecol-ogy is often contradictory For instance thereported shade tolerance of black ash varies

widely (eg Sims et al 1990 Wright andRauscher 1990 Gustafson and Sturtevant2013 Reyes et al 2013) Because of theminimal information available on black ashand the imminent threat from EAB it is es-sential to compile and distribute up-to-datecomprehensive information on black ash be-fore EAB devastates the resource

To assess current knowledge aboutblack ash a group of basket-tree harvestersfrom Maine (Wabanaki tribes) and upstateNew York (Saint Regis Mohawk Tribe)along with a group of black ash researchersassembled at the University of Mainersquos BlackAsh Symposium in November 2014 Thegroup discussed traditional ecologicalknowledge related to black ash and the bas-ketry tradition recent advancements inblack ash ecological research misconcep-tions about the speciesrsquo biology and ecologyand the threat EAB poses to the black ashresource The group then visited four blackash stands in Maine where they assessed thespeciesrsquo current status and discussed poten-tial responses to EAB infestations Based onthese assessments recommendations weredeveloped to address best basket-tree har-vesting practices and to maintain black ashon the landscape in the presence of EABAdditional research needs were also outlinedto fill knowledge gaps and test initial recom-mendations Here we provide a general over-view of basket-quality ash synthesize cur-rent knowledge on black ash biology andreport findings that resulted from the diversecollaboration of tribal university state andfederal partners

Traditional EcologicalKnowledge on Basket-QualityAsh

Basket-tree harvesters and basketmak-ers are the primary forces driving black ashecological research in Maine and other re-gions Maine tribes formed the Black AshTask Force in the early 1990s to address con-cerns about black ash decline The TaskForce which is a partnership of tribal univer-sity and state and federal personnel remainsactive and is currently addressing the potentialloss of black ash due to EAB One approach isto identify individual traits and site character-istics that produce suitable basket-quality treeswhich in turn can guide priorities for tree andstand management before during and afterthe EAB invasion occurs

Each black ash basket-tree harvester hasa range of knowledge and opinions regard-ing the specific attributes associated withbasket-quality trees After a suitable basket-quality tree is identified and felled the trunkis ldquopoundedrdquo to separate the annual growthrings thus creating thin splints that can bemanually smoothed and gauged Differentwood qualities may be sought for differenttypes of baskets from sturdy pack baskets orpotato baskets (Figure 1) to delicate decora-tive fancy baskets As a result each harvesterwill vary in his or her selection of the ldquoidealrdquotree What follows is an attempt to characterizeattributes that can provide an initial indicationof basket-quality trees in the field based on theobservations of one or more basket-tree har-vesters Whether a tree is ultimately selectedfor basketmaking depends on the needs andpreferences of each harvester

Management and Policy Implications

Biological invasions of nonnative organisms in forested ecosystems can have profound economic andecological impacts Black ash is a cultural keystone species in the United States and Canada and is currentlythreatened by the nonnative emerald ash borer (EAB) A better understanding of black ash biology andecology in light of this threat will have significant implications for its future management survival andcontinued use as a source of cultural heritage and basketmaking traditions To address gaps in black ashknowledge that influence basket-quality ash researchers are encouraged to concentrate efforts onstudying the following stand dynamics across a range of hydrologic regimes the role of geneticson basket-quality black ash and EAB population dynamics in infested black ash stands and the effect onregeneration of basket-quality ash Researchers are also encouraged to collaborate with tribal partners andbasketmakers during these studies Diverse collaboration toward a common cause can result in a synergyof traditional ecological knowledge and scientific knowledge that leads to effective and informedmanagement and policy decisionmaking To that effect resource managers and policymakers are alsoencouraged to cooperate with one another to locate maintain and manage existing black ash populationswhile simultaneously considering accessibility to high-quality black ash trees for cultural and basketmakingtraditions before basket-quality black ash in North America are decimated by the EAB invasion

436 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

Site HydrologyWhereas black ash is commonly associ-

ated with poorly drained wetland sites bas-ket-quality black ash is frequently found onsites where water flows or seeps through thesite it is not stagnant (Diamond and Emery2011 Costanza 2015 Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) Black ashoften grows more slowly with smaller an-nual growth rings when there is ponded wa-ter at the surface However soils that aresaturated from a nearby source via capillarityprovide ideal basket-quality conditions soilsare wet enough to allow black ash to out-compete other species but not overly satu-rated to inhibit radial growth (Telander et al2015 Robert Slesak Minnesota Forest Re-sources Council pers comm June 52015) Because of this sites that are wet be-cause of proximity to water or topography(eg swales) produce better-quality ash treesthan sites where hydrology is driven by aconstraining soil horizon or property whichis why some harvesters associate basket-qual-ity black ash with river terraces and flood-plains (Basket-tree harvesters Maine IndianBasketmakers Alliance pers comm MayndashAugust 2013)

Rooting DepthSites that are saturated later into the

growing season and are nutrient poor typi-cally do not produce basket-quality blackash If there are swollen root buttresses orroots that are exposed and easily visible itindicates that the tree and site are probablyinadequate for basketry These attributes

typically represent shallow rooting depth be-cause of impenetrable soils or high subsur-face water tables both of which producepoor-quality black ash with annual ringsthat are too narrow for basketry (Basket-treeharvesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013Symposium field observations November2014)

Annual Ring-WidthFor many harvesters annual ring-width

is the most important factor in determiningthe basket quality of black ash Good basket-quality wood contains growth rings 008inyear for 10 or more consecutive years(Benedict and Frelich 2008) but ideally an-nual growth is relatively consistent through-out the treersquos life If growth rings are toonarrow (eg 004 in) they are unusablefor weaving However growth rings that aretoo large (eg 020 in) are also undesir-able because they result in thick splints thatare difficult to work with (Basket-tree har-vesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013) Ahigh-quality basket tree typically produces8ndash12 splints per radial inch (ca 008ndash012in ring-widthyear)

Stand and Species CompositionA typical stand can contain both good-

and poor-quality black ash Whereas thereare specific site characteristics that often leadto the production of basket-quality ash trees(see above) microsite and stand characteris-tics are also important Light availability iscritical basket-quality ash trees tend to be

more open grown receiving intermediate tomoderate light Likewise microtopographyis also critical within a few inches dramaticchanges in soil water table depth and root-ing depth can be observed (Symposium fieldobservations November 2014)

Basket-quality ash trees are often asso-ciated with reduced competition from otherspecies which in turn increases light avail-ability (Symposium field observations No-vember 2014) Good sites frequently con-tain few if any conifers such as northernwhite-cedar (Thuja occidentalis L) and bal-sam fir (Abies balsamea [L] Mill) Whenconifers are present some harvesters haveobserved an increase in the brittleness ofblack ash wood collected from the site mak-ing their splints more likely to crack andbreak Furthermore common conifer asso-ciates of black ash (eg northern white-ce-dar balsam fir and red spruce [Picea rubensSarg]) can be indicative of poorly drainedsoil conditions which result in poor-qualityblack ash trees (Basket-tree harvestersMaine Indian Basketmakers Alliance perscomm MayndashAugust 2013)

CanopyTrees in the overstory (dominant or

codominant) with full deep crowns thatcover more than 50 of the height of thebole are preferred for basket-quality trees Ifdead limbs or epicormic branching are pres-ent the tree is considered unhealthy and of-ten avoided However dying trees are occa-sionally used if the annual ring-width(described earlier) looks acceptable (Basket-tree harvesters Maine Indian BasketmakersAlliance pers comm MayndashAugust 2013)

BarkTrees with increased bark texture and

bark depth are preferred however black ashrarely displays the distinct furrows seen ongreen or white ash (Diamond and Emery2011) If the bark takes on a ldquocorkyrdquo appear-ance (slightly flexible moist and sticks outfrom the bole) (Figure 2A) it is frequentlyassociated with annual growth rings ideal forbasketry (Basket-tree harvesters Maine In-dian Basketmakers Alliance pers commMayndashAugust 2013) One possible explana-tion for this association is that thicker xylemincrements imply good growth conditionsthat would also produce thicker phloem andcork from lateral meristems which wouldresult in corky bark Faster growth in lateralmeristems probably results from increasedlight availability and more favorable site hy-

Figure 1 Black ash basket Black ash baskets take many shapes and forms from sturdypack baskets to decorative fancy baskets Other baskets mix utility with decoration InMaine potato baskets are common including this basket by Richard Silliboy of the HoultonBand of Micmacs

Journal of Forestry bull September 2017 437

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

drology (Symposium field observations No-vember 2014) Bark characteristics were akey indicator for many basket-tree harvesterstaking part in the Symposium Further-more participants in the Symposium ob-served that increment borer holes healedfaster in trees with ldquocorkyrdquo bark (Figure 2A)than in those with ldquononcorkyrdquo bark (Figure2B) Bark color can also indicate tree qualityA yellow-tinted bark was often associatedwith increased annual growth and improvedwood quality whereas a gray-tinted barkwas associated with poorer quality and brit-tle wood

Tree AttributesSpecific features within the tree can af-

fect its usability The sapwood (aka ldquowhitewoodrdquo) is typically used for fancy basketswhereas the heartwood (aka ldquobrown woodrdquo)is commonly used for pack and potato bas-kets Stem defects such as cracks branchstubs or holes can make the wood unusablefor splints In addition if the tree bole is toocurved it cannot be processed (ldquopoundedrdquo)for splints A harvester can work with a mod-erate sweep in the bole but it takes moretime and effort to pound (Basket-tree har-vesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013)

A good-quality tree will have a straightbole for 45ndash6 ft (or more) and that sectionwill be free of defects Tree size can range

anywhere from 5 to 25 in dbh (breastheight 45 ft) but selection is highly de-pendent on how the tree will be transported(eg on the harvesterrsquos shoulder four-wheeler or truck) (Basket-tree harvestersMaine Indian Basketmakers Alliance perscomm MayndashAugust 2013)

General Black Ash BiologyBlack ash grows on a range of soil types

from peat bogs and mineral soil wetlands towell-drained stream edges and ridges (Simset al 1990 Wright and Rauscher 1990 Eh-renfeld 2012) The species is most fre-quently found on flooded or seasonally wetsites probably due to reduced competitionfrom other species incapable of establishingin hydric conditions Tardif et al (1994) re-ported that young black ash stems can pro-duce hypertrophied lenticels to increase gasexchange as well as use adventitious roots(Sena Gomes and Kozlowski 1980 Tardifet al 1994 Mendelssohn et al 2014) Bothof these adaptations allow young stems topersist on wet sites

When black ash becomes established onbetter-drained mesic sites the species fre-quently exhibits increased annual growthand larger dbh (Costanza 2015 pers observof authors 2015) Conversely in stands sat-urated throughout the growing seasonblack ash trees typically have narrower an-

nual growth rings reduced dbh exposedroots and numerous sprouts One explana-tion is that black ash is often classified ashaving a shallow root system (Wright andRauscher 1990) which is more affected bywater table fluctuations in turn affecting an-nual growth rates Although long-term satu-ration produces poor-quality ash sites withspring flooding followed by surface dryingin the summer favor black ash growth (Simset al 1990 Lenhart et al 2012 Slesak et al2014) These better-drained mesic sites aretherefore locations where basket-tree har-vesters frequently search for basket-qualitytrees Further these differences in site hy-drology that influence black ash growth andquality may be captured in the NaturalCommunity classifications that many statesand provinces have developed to describevegetation soils and site conditions of com-mon communities (see Supplemental TableS1 )

Black ash regeneration also varieswidely The species is capable of both pro-ducing seed and sprouting when cut orstressed yet sprouts from the root collartend to have a higher growth rate as they aregrowing from an already developed root sys-tem (Tardif and Bergeron 1999) Thesesprouts also tend to be more tolerant offlooding and droughts (Sims et al 1990Tardif and Bergeron 1999) Further siteconditions (eg flooding and drought) mayheavily influence seedling survival and estab-lishment Two consecutive dryreducedflood years may be needed for successful es-tablishment the first year allows the seed tobreak dormancy and germinate whereas thesecond year helps establishment and survivalof future flooding (Symposium field obser-vations November 2014) Ash can also pro-duce persistent seeds which is a benefit insites with fluctuating hydrologic regimesHowever several biotic or abiotic site factorsmay influence the survivorship of this seedbank including insects moisture fungi andothers (Klooster et al 2014)

Based on four recent studies averagetree age in black ash dominated stands typi-cally ranges from 35 to 100 years old with afew black ash trees greater than 200 years old(Table 1) (Tardif and Bergeron 1999 Bene-dict and Frelich 2008 Townshend 2011Costanza 2015) Although large old-growthblack ash forests (200 years old) exist inNorth America they are not common

Figure 2 Comparison of black ash bark quality Black ash bark takes on a variety ofappearances Basket-tree harvesters frequently associate spongy corky thicker bark (A)with high-quality basket wood Conversely bark with long thin brittle flakes (B) isfrequently associated with poor-quality black ash

Supplementary data are available with this article at httpdxdoiorg105849jof2016-034R1

438 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

In more disturbed areas (both naturallyand anthropogenically) average tree age isyounger Given basketmakersrsquo preferencefor 008ndash012 in ring-widths and trees5ndash25 in dbh this typical black ash age rangematches the typical expectations of basket-quality trees

Reports on black ash shade tolerancediffer One perspective describes the speciesas shade intolerant (Wright and Rauscher1990 Gustafson and Sturtevant 2013)whereas another describes the species asmoderately shade tolerant becoming moreintolerant with age (Sims et al 1990 Barnesand Wagner 2004) Tolerance may also de-pend on origin seedlings and sprouts mayhave different levels of shade toleranceBased on several recent observations (Reyeset al 2013 Fisichelli et al 2014 Looneyet al 2016 Symposium field observationsNovember 2014) the species appears to bemoderately shade tolerant Black ash is capa-ble of germinating and surviving in the un-derstory for several years although low tomoderate light is required for survival aclosed canopy will result in little to no seed-ling survivorship In Ontario Townshend(2011) found black ash trees of substantialage (eg up to 80 years old) under the mainforest canopy suggesting that the species iscapable of withstanding long periods of sup-pression If black ash can persist in the un-derstory the surviving seedlings could be animportant source of future black ash stemsduring the EAB invasion

Site characteristics regeneration strate-gies and stand dynamics all play a key role inthe quality of black ash trees and heavily in-fluence basketmaking traditions By im-proving our understanding of black ashbiologyecology in Maine and other blackash habitats improved management strate-

gies that simultaneously target EAB controland black ash preservation which will serveto maintain this cultural keystone speciescan be developed

Case Study Four Black AshStands in Maine

In November 2014 basket-tree har-vesters and researchers visited four black ashsites in Maine as part of the Black Ash Sym-posium These sites were representative oftwo common black ash stand types found ina statewide study black ash swamps andhardwood river terraces (Gawler and Cutko2010 Costanza 2015 see also SupplementalTable S1) two sites from each stand typewere visited (Figures 3ndash6) The site visitsallowed Symposium participants to sharetheir knowledge of black ash while also as-sessing the potential impact of EAB on thesetypes of stands and on basket-quality ashWhat follows is a summary of the discus-sions and observations from the Symposiumsite visits

The four black ash stands visited are ex-amples of old-field early-successional forests(100 years old) common in central andsouthern Maine which regenerate on aban-doned agricultural land The two black ashswamps were characterized by standing wa-ter ponding and relatively flat topographywith limited pits and mounds (Figures 4 and6) The two hardwood river terraces in con-trast had little to no standing water slightdownstream slopes and small streams drain-ing the sites (Figures 3 and 5) Based on in-crement cores collected in 2012 (Costanza2015) the highest basket-quality black ashtrees were found on the two river terracesites Harvesters attending the Symposiumsupported this finding noting that they of-

ten associate high-quality black ash withriver banks and floodplains with low tomoderate stand densities

The four Maine stands had significantlylower black ash densities than documentedstands found in Minnesota Ontario andQuebec (Table 2) Stands visited rangedfrom 90ndash162 black ash stemsac1 with theswamps producing higher black ash densi-ties than the river terraces One possible ex-planation for these reduced stand densities isthat most Maine stands are early-succes-sional forests growing in previously har-vested stands or old fields Studies outside ofMaine have focused on well-established for-ests where black ash maintains high densitiesand canopy dominance It is also possiblethat black ash trees in Maine recruit differ-ently from those in other regions howeverrecruitment strategies and shade toleranceneed to be further studied to draw signifi-cant conclusions During the Symposiumsite visits basket-tree harvesters and re-searchers both noted that basketry is moreprevalent in Maine New York and south-eastern Ontario Lower stand densities anddifferent recruitment patterns along withcultural traditions may influence the pro-portion of basket-quality black ash trees inthese regions compared with others withinthe black ash range

Unlike those in many other regionsblack ash trees in Maine frequently occur inmixed species stands which can includenorthern white-cedar balsam fir Americanelm (Ulmus americana L) red maple (Acerrubrum L) and yellow birch (Betula al-leghaniensis Britton) (Gawler and Cutko2010 Costanza 2015) Pure black ash standsare uncommon unless the site is a swamp oran old-growth black ash stand (Ehrenfeld2012 Costanza 2015) Of the sites visited

Table 1 Black ash average tree age in North America based on four studies that sampled a variety of black ash stands across thenorthern United States and southern Canada

Location StudyMaximum black ash

age (4 in dbh)Average black ashage (4 in dbh) No of black ash stands

(yr)

Maine Costanza 2015 273 59 24 stands (1frasl6 ac)Chippewa National Forest Minnesota Benedict and Frelich 2008 1641311351 9698651 67 stands (164 ft radius) 2424191

Northern New York2 Costanza 2015 146 70 16 stands (1frasl6 ac)Ontario Canada Townshend 2011 1613 61 16 stands (1frasl10 ac)Lake Duparquet Queacutebec Canada Tardif and Bergeron 1999 3204 34 5 stands (composed of 54 plots 1frasl40 ac each)

Research sites ranged from pure black ash to mixed forest stands and from old-growth to anthropogenically disturbed stands1 Values correspond to three types of sites lowlandsuplandswoodland ponds Maximum and average tree ages were taken from stand data that included all species however stands were dominatedby black ash A total of 166 black ash were sampled2 In northern New York samples were collected in St Lawrence and Lewis counties3 Oldest veteran black ash tree throughout sample area was 161 years old Oldest veteran black ash tree in each stand ranged from 63ndash161 years old4 The oldest black ash was sexually reproduced The oldest black ash sprout was 120 yr old

Journal of Forestry bull September 2017 439

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

during the Symposium the black ashswamps had moderate conifer densities (pri-marily balsam fir) whereas the hardwoodriver terraces were mixed hardwood standscomposed of red maple American basswood(Tilia americana L) green ash and whiteash (Fraxinus americana L) Harvesters andresearchers both noted the improved qualityof ash on the river terraces and often associ-ated these basket-quality trees with the lackof conifers innear the stand

At the end of the Symposium severalconclusions about Mainersquos black ash popu-lation emerged Unlike those in other re-gions of North America the four blackash stands we visited were early-successional

stands characterized by low to moderatestand densities mixed species compositionand reduced black ash regeneration In ad-dition harvesters confirmed that these char-acteristics (site hydrology species composi-tion and stand structure) play a key role inthe quality of black ash trees produced Withregard to management implications thesites visited were similar to stands in Ontariowhere black ash is a secondary or minorcomponent (Jeffrey Dech Nipissing Uni-versity pers comm Feb 2 2016) andmany green ash stands in southeasternMichigan (Kashian 2016) Therefore re-search from these stands may provide valu-able insight into how similar stands in

Maine and other regions are likely to re-spond to EAB infestations

The EAB ImpactThe potential economic and ecological

impacts of the EAB invasion are staggering(Kovacs et al 2010 Aukema et al 2011) andwill drastically alter the ash components ofNorth American forests (Cappaert et al2005 Herms and McCullough 2014)Black ash in particular is highly susceptibleto the insect (Anulewicz et al 2006 Pure-swaran and Poland 2009 Siegert et al2014a) and dies within 2ndash4 years of beinginfested (Siegert et al 2005 2014b) (Figure7) In EAB-infested areas research has yet tofully address the ecological impacts to blackash Several studies report ash mortality ratesas a pooled statistic including white greenand black ash (eg Kashian and Witter2011 Klooster et al 2014) whereas otherstudies only address early infestation periods(2004ndash2008) and focus on Michigan andOhio (eg Siegert et al 2005 Smith et al2015 Kashian 2016) Researchers in Min-nesota have focused on black ash standssimulating EAB mortality by girdling blackash trees (Slesak et al 2014 Looney et al2015) However additional research isneeded to more critically evaluate black ashstand dynamics through the course of anEAB invasion and in the different habitatsand hydrologic regimes where black ash oc-curs

Based on green ash studies conductedin Michigan after EAB infestation (Kashian2016) it is possible to make a few inferencesabout the potential response of black ashstands to EAB In Michigan green ash treesare persisting with sprouts released ad-vanced regeneration and seedlings pro-duced from sprout-borne seeds on siteswhere green ash densities were relativelyhigh Conversely preliminary work suggestssprouting from EAB-killed black ash may beextremely limited (NW Siegert USDAForest Service unpubl data 2010) Re-duced densities of black ash will probablyleave stands depleted with fewer and smallerstems following the initial wave of the EABinvasion Any surviving black ash trees willprobably be small (3 in dbh) unless estab-lished EAB infestations are radically sup-pressed One study in southeastern Michi-gan conducted when EAB populations wererelatively high found that sprouts from har-vested trees were recolonized by EAB within1 year (Petrice and Haack 2011) Furtherwork by Tardif and Bergeron (1999) indi-

Figure 3 Site 1 hardwood river terrace Site 1 is a sparse to moderately dense black ashmixed hardwood stand The stand is separated from a neighboring conifer stand by a smallstream (A) The majority of black ash gt4 in dbh at this site are high quality (B)

440 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

cates a greater abundance of black ashsprouts than seedlings in young age classesin addition to higher mortality rates insprouts To fully understand how black ashsprouts may persist in EAB-infested standsfurther investigation is needed on sproutingprocesses the survivorship of black ashsprouts the relationship to EAB populationcycles and the ability of these sprouts to be-come basket-quality trees

After a few years of EAB infestationonce the overstory has been killed estab-lished black ash seedlings if present may bereleased if the resulting gaps are not quicklyfilled in by neighboring trees However be-

cause EAB is known to infest and kill trees1 in dbh (Herms and McCullough2014) it is likely that most seedlings inmixed or low-density stands will not achievea larger size (remain 3 in dbh) and themean stand diameter will remain lowWhere black ash is a dominant componentof the stand changes in evapotranspirationafter overstory loss may result in water tablerise (Slesak et al 2014) and conversion toa more open-water wetland condition(Erdmann et al 1987) further threaten-ing ash persistence in the stand In addi-tion gap formation will probably result inan increased risk of competition from re-

sidual non-ash understory trees unaffectedby EAB as well as shrubs and herbaceousvegetation (Slesak et al 2014) Based onthe rapid detrimental impact of EAB it iscritical to identify what additional infor-mation and efforts are needed to preparefor the insectrsquos arrival in many black ashstands

RecommendationsThe first step in preparing for and man-

aging EAB is early detection If and when EABis detected efforts should focus on slowing theinsectrsquos spread (Siegert et al 2015) so the avail-ability of existing basket-quality ash trees canbe extended as long as possible Managementstrategies may be implemented to complementthis effort by reducing EAB populationdensities (McCullough et al 2015) In do-ing so guidelines should be developedthat limit anthropogenic spread of EAB-infested material while allowing harvestersto safely transport basket-quality black ashmaterial without risk of transporting EABto non-infested areas For instance tworecent studies reported that submerginginfested black ash logs in water for 8 ndash16weeks killed all EAB while still maintain-ing useable basket-quality wood (Siegertet al 2014a Poland et al 2015)

If harvesters are available basket-qual-ity trees should be harvested pounded andsplit as soon as an EAB infestation is de-tected in the vicinity This harvesting in-crease would have a considerably differentimpact on forests and on black ash in par-ticular than tribal traditions historicallyhad However dried black ash splints can bestored for several years (Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) increas-ing harvesting would allow harvesters to usewood that would otherwise be lost to EABLightly infested black ash trees remain suit-able for basketmaking materials but degraderapidly as EAB density increases reducingyields from a given log to the point where itmay be unusable for splints (Siegert et al2014a Poland et al 2015) In additionsome have recommended increasing blackash harvesting now before EAB arrives Thiswould potentially reduce concerns aboutquarantines accidental insect transport andloss of basket-quality wood after an EAB in-festation

Last we recommend collaboration be-tween state federal and tribal agencies todevelop memorandums of understanding orjoint agreements before the arrival of EAB

Figure 4 Site 2 black ash swamp Site 2 is a moderately dense mixed hardwood-coniferstand with black ash red maple and balsam fir The site is characterized by minimalponding (A) root buttressing (A) and small black ash trees typically lt6 in dbh (B) Themajority of black ash gt4 in dbh are poor quality

Journal of Forestry bull September 2017 441

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

in high-value black ash stands Such agree-ments can alleviate confusion and promoteaction as soon as EAB is detected in or neartribal lands or other traditional black ashharvesting sites Furthermore such agree-ments can help foster discussions on accessi-bility to black ash trees To preserve theblack ash resource and maintain black ashbasketry in the face of EAB discussions re-garding accessibility and response plansneed to occur

Future ResearchIn addition to making preparations

for EABrsquos arrival in black ash stands there

are countless opportunities to learn moreabout this still-overlooked tree species Toimprove our understanding of black ashtreesrsquo potential response to EAB and im-pacts to cultural traditions future researchdirections were discussed during the BlackAsh Symposium What follows is a summaryof the three most pressing research needsinfluence of black ash stand dynamics andsite hydrology on regeneration and qualityof black ash role of black ash genetics in thefuture of the population and EAB popula-tion dynamics in infested black ash standsand the effect on black ash regeneration

Influence of Stand Dynamics and Hy-drology on Regeneration and Quality

Very little information exists on blackash stand histories conditions and naturaldynamics in Maine For example it is un-clear whether Maine black ash stands un-dergo continuous recruitment and remainstocked over an extended period of time or ifblack ash is replaced by other species Natu-ral stand dynamics and succession are poorlyunderstood partially because there are fewold-growth stands in Maine Locating andstudying a few old-growth black ash stands(200 years old) would greatly improve ourcurrent understanding of succession Theinclusion of managed stands would alsoprove beneficial allowing researchers toquantify the response of black ash to silvicul-tural treatments Furthermore describingthe dynamics of ash across a range of distur-bance histories would allow researchers todevelop a much needed successional frame-work for black ash and to develop a descrip-tion of the speciesrsquo regeneration niche In themidwestern United States recent studieshave suggested that the future of black ashstands faced with EAB and climate changewill probably center around non-ash re-placement species (Looney et al 2015Iverson et al 2016) However in regionswhere black ash is a cultural keystone spe-cies replacing black ash is not an optimalmanagement strategy Describing black ashstand dynamics and recruitment in Maineas well as in other black ash habitats such asthose in Nova Scotia southern and centralOntario and Quebec would provide in-sight on how to maintain the species bothfor basketmaking and ecological reasons

Stand dynamics are heavily influencedby stand hydrology which is poorly under-stood in Maine and other parts of the blackash range A few recent studies in Minnesotahave explored black ash stand hydrology(Slesak et al 2014 Telander et al 2015)however the extent to which water tablefluctuations in black ash stands influence re-generation and recruitment across the entirerange should be further explored For exam-ple in situations where water table dynamicsare driven primarily by topography or localsurface water a change in evapotranspira-tion after overstory loss may not result in aconcomitant change in site hydrology (Rob-ert Slesak Minnesota Forest ResourcesCouncil pers comm June 5 2015) Fur-thermore the role of post-disturbancesprouting of black ash in regulating watertable dynamics is also unclear as is the role

Figure 5 Site 3 hardwood river terrace Site 3 is a moderately dense mixed hardwoodstand with occasional conifer (A) A small stream runs northeast to southwest and anunpaved road runs west to east through the stand In better-drained areas large high-quality black ash are produced Where ponding occurs black ash tend to be smaller andpoorer quality The majority of black ash gt4 in dbh are high quality (B)

442 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

from black ash via a variety of basketmakingtraditions

Like many of the Native Americans andFirst Nations people in the native range ofblack ash the Wabanaki tribes in Maine(Aroostook Band of Micmac Indians Houl-ton Band of Maliseet Indians Passamaquo-ddy Tribe and Penobscot Indian Nation)and the Saint Regis Mohawk Tribe in NewYork and Canada have a long history of har-vesting black ash for basketry (Benedict andFrelich 2008 Diamond and Emery 2011)Specific wood qualities such as strength andflexibility make black ash unique and ex-plain in part why it is highly preferred forbasketry (Benedict and David 2004) Yet thespeciesrsquo cultural significance goes beyondweaving baskets black ash is at the spiritualcenter of the tribesrsquo teachings including oneof their creation stories (Leland 1884 Bene-dict and David 2004) Many basketmakersrely on ldquobasket-tree harvestersrdquo these indi-viduals are tribal members who have learnedto identify favorable sites and recognize in-dividual black ash trees that are likely to pro-duce basket-quality materials As such theyare also responsible for maintaining andpassing on tribal knowledge about the spe-cies from generation to generation

Emerald ash borer (EAB) (Agrilus pla-nipennis Fairmaire) an invasive insect nativeto Asia that feeds on ash (Fraxinus spp) hascaused the Wabanakis Mohawks and oth-ers great concern about the future of blackash Initially detected in southeastern Mich-igan and southern Ontario in 2002 the EABinvasion has spread rapidly (Siegert et al2014b) and is responsible for killing mil-lions of North American ash trees (Cappaertet al 2005 Herms and McCullough 2014)As of July 2016 EAB has been detected in27 states and the Canadian provinces of On-tario and Quebec1 Although black ashstands in many locations still have valuablebasket-quality trees including those inMaine where to date no EAB detectionshave occurred the EAB invasion nonethe-less continues to spread and threaten whatremains of this unique resource (Siegert et al2015)

In addition to its significance as a cul-tural resource black ash has commercialvalue and can be used for veneer cabinetsand furniture (Ross 2010) Yet relatively lit-tle is known about this important species(Wright and Rauscher 1990) and the scien-tific literature on black ash biology and ecol-ogy is often contradictory For instance thereported shade tolerance of black ash varies

widely (eg Sims et al 1990 Wright andRauscher 1990 Gustafson and Sturtevant2013 Reyes et al 2013) Because of theminimal information available on black ashand the imminent threat from EAB it is es-sential to compile and distribute up-to-datecomprehensive information on black ash be-fore EAB devastates the resource

To assess current knowledge aboutblack ash a group of basket-tree harvestersfrom Maine (Wabanaki tribes) and upstateNew York (Saint Regis Mohawk Tribe)along with a group of black ash researchersassembled at the University of Mainersquos BlackAsh Symposium in November 2014 Thegroup discussed traditional ecologicalknowledge related to black ash and the bas-ketry tradition recent advancements inblack ash ecological research misconcep-tions about the speciesrsquo biology and ecologyand the threat EAB poses to the black ashresource The group then visited four blackash stands in Maine where they assessed thespeciesrsquo current status and discussed poten-tial responses to EAB infestations Based onthese assessments recommendations weredeveloped to address best basket-tree har-vesting practices and to maintain black ashon the landscape in the presence of EABAdditional research needs were also outlinedto fill knowledge gaps and test initial recom-mendations Here we provide a general over-view of basket-quality ash synthesize cur-rent knowledge on black ash biology andreport findings that resulted from the diversecollaboration of tribal university state andfederal partners

Traditional EcologicalKnowledge on Basket-QualityAsh

Basket-tree harvesters and basketmak-ers are the primary forces driving black ashecological research in Maine and other re-gions Maine tribes formed the Black AshTask Force in the early 1990s to address con-cerns about black ash decline The TaskForce which is a partnership of tribal univer-sity and state and federal personnel remainsactive and is currently addressing the potentialloss of black ash due to EAB One approach isto identify individual traits and site character-istics that produce suitable basket-quality treeswhich in turn can guide priorities for tree andstand management before during and afterthe EAB invasion occurs

Each black ash basket-tree harvester hasa range of knowledge and opinions regard-ing the specific attributes associated withbasket-quality trees After a suitable basket-quality tree is identified and felled the trunkis ldquopoundedrdquo to separate the annual growthrings thus creating thin splints that can bemanually smoothed and gauged Differentwood qualities may be sought for differenttypes of baskets from sturdy pack baskets orpotato baskets (Figure 1) to delicate decora-tive fancy baskets As a result each harvesterwill vary in his or her selection of the ldquoidealrdquotree What follows is an attempt to characterizeattributes that can provide an initial indicationof basket-quality trees in the field based on theobservations of one or more basket-tree har-vesters Whether a tree is ultimately selectedfor basketmaking depends on the needs andpreferences of each harvester

Management and Policy Implications

Biological invasions of nonnative organisms in forested ecosystems can have profound economic andecological impacts Black ash is a cultural keystone species in the United States and Canada and is currentlythreatened by the nonnative emerald ash borer (EAB) A better understanding of black ash biology andecology in light of this threat will have significant implications for its future management survival andcontinued use as a source of cultural heritage and basketmaking traditions To address gaps in black ashknowledge that influence basket-quality ash researchers are encouraged to concentrate efforts onstudying the following stand dynamics across a range of hydrologic regimes the role of geneticson basket-quality black ash and EAB population dynamics in infested black ash stands and the effect onregeneration of basket-quality ash Researchers are also encouraged to collaborate with tribal partners andbasketmakers during these studies Diverse collaboration toward a common cause can result in a synergyof traditional ecological knowledge and scientific knowledge that leads to effective and informedmanagement and policy decisionmaking To that effect resource managers and policymakers are alsoencouraged to cooperate with one another to locate maintain and manage existing black ash populationswhile simultaneously considering accessibility to high-quality black ash trees for cultural and basketmakingtraditions before basket-quality black ash in North America are decimated by the EAB invasion

436 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

Site HydrologyWhereas black ash is commonly associ-

ated with poorly drained wetland sites bas-ket-quality black ash is frequently found onsites where water flows or seeps through thesite it is not stagnant (Diamond and Emery2011 Costanza 2015 Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) Black ashoften grows more slowly with smaller an-nual growth rings when there is ponded wa-ter at the surface However soils that aresaturated from a nearby source via capillarityprovide ideal basket-quality conditions soilsare wet enough to allow black ash to out-compete other species but not overly satu-rated to inhibit radial growth (Telander et al2015 Robert Slesak Minnesota Forest Re-sources Council pers comm June 52015) Because of this sites that are wet be-cause of proximity to water or topography(eg swales) produce better-quality ash treesthan sites where hydrology is driven by aconstraining soil horizon or property whichis why some harvesters associate basket-qual-ity black ash with river terraces and flood-plains (Basket-tree harvesters Maine IndianBasketmakers Alliance pers comm MayndashAugust 2013)

Rooting DepthSites that are saturated later into the

growing season and are nutrient poor typi-cally do not produce basket-quality blackash If there are swollen root buttresses orroots that are exposed and easily visible itindicates that the tree and site are probablyinadequate for basketry These attributes

typically represent shallow rooting depth be-cause of impenetrable soils or high subsur-face water tables both of which producepoor-quality black ash with annual ringsthat are too narrow for basketry (Basket-treeharvesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013Symposium field observations November2014)

Annual Ring-WidthFor many harvesters annual ring-width

is the most important factor in determiningthe basket quality of black ash Good basket-quality wood contains growth rings 008inyear for 10 or more consecutive years(Benedict and Frelich 2008) but ideally an-nual growth is relatively consistent through-out the treersquos life If growth rings are toonarrow (eg 004 in) they are unusablefor weaving However growth rings that aretoo large (eg 020 in) are also undesir-able because they result in thick splints thatare difficult to work with (Basket-tree har-vesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013) Ahigh-quality basket tree typically produces8ndash12 splints per radial inch (ca 008ndash012in ring-widthyear)

Stand and Species CompositionA typical stand can contain both good-

and poor-quality black ash Whereas thereare specific site characteristics that often leadto the production of basket-quality ash trees(see above) microsite and stand characteris-tics are also important Light availability iscritical basket-quality ash trees tend to be

more open grown receiving intermediate tomoderate light Likewise microtopographyis also critical within a few inches dramaticchanges in soil water table depth and root-ing depth can be observed (Symposium fieldobservations November 2014)

Basket-quality ash trees are often asso-ciated with reduced competition from otherspecies which in turn increases light avail-ability (Symposium field observations No-vember 2014) Good sites frequently con-tain few if any conifers such as northernwhite-cedar (Thuja occidentalis L) and bal-sam fir (Abies balsamea [L] Mill) Whenconifers are present some harvesters haveobserved an increase in the brittleness ofblack ash wood collected from the site mak-ing their splints more likely to crack andbreak Furthermore common conifer asso-ciates of black ash (eg northern white-ce-dar balsam fir and red spruce [Picea rubensSarg]) can be indicative of poorly drainedsoil conditions which result in poor-qualityblack ash trees (Basket-tree harvestersMaine Indian Basketmakers Alliance perscomm MayndashAugust 2013)

CanopyTrees in the overstory (dominant or

codominant) with full deep crowns thatcover more than 50 of the height of thebole are preferred for basket-quality trees Ifdead limbs or epicormic branching are pres-ent the tree is considered unhealthy and of-ten avoided However dying trees are occa-sionally used if the annual ring-width(described earlier) looks acceptable (Basket-tree harvesters Maine Indian BasketmakersAlliance pers comm MayndashAugust 2013)

BarkTrees with increased bark texture and

bark depth are preferred however black ashrarely displays the distinct furrows seen ongreen or white ash (Diamond and Emery2011) If the bark takes on a ldquocorkyrdquo appear-ance (slightly flexible moist and sticks outfrom the bole) (Figure 2A) it is frequentlyassociated with annual growth rings ideal forbasketry (Basket-tree harvesters Maine In-dian Basketmakers Alliance pers commMayndashAugust 2013) One possible explana-tion for this association is that thicker xylemincrements imply good growth conditionsthat would also produce thicker phloem andcork from lateral meristems which wouldresult in corky bark Faster growth in lateralmeristems probably results from increasedlight availability and more favorable site hy-

Figure 1 Black ash basket Black ash baskets take many shapes and forms from sturdypack baskets to decorative fancy baskets Other baskets mix utility with decoration InMaine potato baskets are common including this basket by Richard Silliboy of the HoultonBand of Micmacs

Journal of Forestry bull September 2017 437

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

drology (Symposium field observations No-vember 2014) Bark characteristics were akey indicator for many basket-tree harvesterstaking part in the Symposium Further-more participants in the Symposium ob-served that increment borer holes healedfaster in trees with ldquocorkyrdquo bark (Figure 2A)than in those with ldquononcorkyrdquo bark (Figure2B) Bark color can also indicate tree qualityA yellow-tinted bark was often associatedwith increased annual growth and improvedwood quality whereas a gray-tinted barkwas associated with poorer quality and brit-tle wood

Tree AttributesSpecific features within the tree can af-

fect its usability The sapwood (aka ldquowhitewoodrdquo) is typically used for fancy basketswhereas the heartwood (aka ldquobrown woodrdquo)is commonly used for pack and potato bas-kets Stem defects such as cracks branchstubs or holes can make the wood unusablefor splints In addition if the tree bole is toocurved it cannot be processed (ldquopoundedrdquo)for splints A harvester can work with a mod-erate sweep in the bole but it takes moretime and effort to pound (Basket-tree har-vesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013)

A good-quality tree will have a straightbole for 45ndash6 ft (or more) and that sectionwill be free of defects Tree size can range

anywhere from 5 to 25 in dbh (breastheight 45 ft) but selection is highly de-pendent on how the tree will be transported(eg on the harvesterrsquos shoulder four-wheeler or truck) (Basket-tree harvestersMaine Indian Basketmakers Alliance perscomm MayndashAugust 2013)

General Black Ash BiologyBlack ash grows on a range of soil types

from peat bogs and mineral soil wetlands towell-drained stream edges and ridges (Simset al 1990 Wright and Rauscher 1990 Eh-renfeld 2012) The species is most fre-quently found on flooded or seasonally wetsites probably due to reduced competitionfrom other species incapable of establishingin hydric conditions Tardif et al (1994) re-ported that young black ash stems can pro-duce hypertrophied lenticels to increase gasexchange as well as use adventitious roots(Sena Gomes and Kozlowski 1980 Tardifet al 1994 Mendelssohn et al 2014) Bothof these adaptations allow young stems topersist on wet sites

When black ash becomes established onbetter-drained mesic sites the species fre-quently exhibits increased annual growthand larger dbh (Costanza 2015 pers observof authors 2015) Conversely in stands sat-urated throughout the growing seasonblack ash trees typically have narrower an-

nual growth rings reduced dbh exposedroots and numerous sprouts One explana-tion is that black ash is often classified ashaving a shallow root system (Wright andRauscher 1990) which is more affected bywater table fluctuations in turn affecting an-nual growth rates Although long-term satu-ration produces poor-quality ash sites withspring flooding followed by surface dryingin the summer favor black ash growth (Simset al 1990 Lenhart et al 2012 Slesak et al2014) These better-drained mesic sites aretherefore locations where basket-tree har-vesters frequently search for basket-qualitytrees Further these differences in site hy-drology that influence black ash growth andquality may be captured in the NaturalCommunity classifications that many statesand provinces have developed to describevegetation soils and site conditions of com-mon communities (see Supplemental TableS1 )

Black ash regeneration also varieswidely The species is capable of both pro-ducing seed and sprouting when cut orstressed yet sprouts from the root collartend to have a higher growth rate as they aregrowing from an already developed root sys-tem (Tardif and Bergeron 1999) Thesesprouts also tend to be more tolerant offlooding and droughts (Sims et al 1990Tardif and Bergeron 1999) Further siteconditions (eg flooding and drought) mayheavily influence seedling survival and estab-lishment Two consecutive dryreducedflood years may be needed for successful es-tablishment the first year allows the seed tobreak dormancy and germinate whereas thesecond year helps establishment and survivalof future flooding (Symposium field obser-vations November 2014) Ash can also pro-duce persistent seeds which is a benefit insites with fluctuating hydrologic regimesHowever several biotic or abiotic site factorsmay influence the survivorship of this seedbank including insects moisture fungi andothers (Klooster et al 2014)

Based on four recent studies averagetree age in black ash dominated stands typi-cally ranges from 35 to 100 years old with afew black ash trees greater than 200 years old(Table 1) (Tardif and Bergeron 1999 Bene-dict and Frelich 2008 Townshend 2011Costanza 2015) Although large old-growthblack ash forests (200 years old) exist inNorth America they are not common

Figure 2 Comparison of black ash bark quality Black ash bark takes on a variety ofappearances Basket-tree harvesters frequently associate spongy corky thicker bark (A)with high-quality basket wood Conversely bark with long thin brittle flakes (B) isfrequently associated with poor-quality black ash

Supplementary data are available with this article at httpdxdoiorg105849jof2016-034R1

438 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

In more disturbed areas (both naturallyand anthropogenically) average tree age isyounger Given basketmakersrsquo preferencefor 008ndash012 in ring-widths and trees5ndash25 in dbh this typical black ash age rangematches the typical expectations of basket-quality trees

Reports on black ash shade tolerancediffer One perspective describes the speciesas shade intolerant (Wright and Rauscher1990 Gustafson and Sturtevant 2013)whereas another describes the species asmoderately shade tolerant becoming moreintolerant with age (Sims et al 1990 Barnesand Wagner 2004) Tolerance may also de-pend on origin seedlings and sprouts mayhave different levels of shade toleranceBased on several recent observations (Reyeset al 2013 Fisichelli et al 2014 Looneyet al 2016 Symposium field observationsNovember 2014) the species appears to bemoderately shade tolerant Black ash is capa-ble of germinating and surviving in the un-derstory for several years although low tomoderate light is required for survival aclosed canopy will result in little to no seed-ling survivorship In Ontario Townshend(2011) found black ash trees of substantialage (eg up to 80 years old) under the mainforest canopy suggesting that the species iscapable of withstanding long periods of sup-pression If black ash can persist in the un-derstory the surviving seedlings could be animportant source of future black ash stemsduring the EAB invasion

Site characteristics regeneration strate-gies and stand dynamics all play a key role inthe quality of black ash trees and heavily in-fluence basketmaking traditions By im-proving our understanding of black ashbiologyecology in Maine and other blackash habitats improved management strate-

gies that simultaneously target EAB controland black ash preservation which will serveto maintain this cultural keystone speciescan be developed

Case Study Four Black AshStands in Maine

In November 2014 basket-tree har-vesters and researchers visited four black ashsites in Maine as part of the Black Ash Sym-posium These sites were representative oftwo common black ash stand types found ina statewide study black ash swamps andhardwood river terraces (Gawler and Cutko2010 Costanza 2015 see also SupplementalTable S1) two sites from each stand typewere visited (Figures 3ndash6) The site visitsallowed Symposium participants to sharetheir knowledge of black ash while also as-sessing the potential impact of EAB on thesetypes of stands and on basket-quality ashWhat follows is a summary of the discus-sions and observations from the Symposiumsite visits

The four black ash stands visited are ex-amples of old-field early-successional forests(100 years old) common in central andsouthern Maine which regenerate on aban-doned agricultural land The two black ashswamps were characterized by standing wa-ter ponding and relatively flat topographywith limited pits and mounds (Figures 4 and6) The two hardwood river terraces in con-trast had little to no standing water slightdownstream slopes and small streams drain-ing the sites (Figures 3 and 5) Based on in-crement cores collected in 2012 (Costanza2015) the highest basket-quality black ashtrees were found on the two river terracesites Harvesters attending the Symposiumsupported this finding noting that they of-

ten associate high-quality black ash withriver banks and floodplains with low tomoderate stand densities

The four Maine stands had significantlylower black ash densities than documentedstands found in Minnesota Ontario andQuebec (Table 2) Stands visited rangedfrom 90ndash162 black ash stemsac1 with theswamps producing higher black ash densi-ties than the river terraces One possible ex-planation for these reduced stand densities isthat most Maine stands are early-succes-sional forests growing in previously har-vested stands or old fields Studies outside ofMaine have focused on well-established for-ests where black ash maintains high densitiesand canopy dominance It is also possiblethat black ash trees in Maine recruit differ-ently from those in other regions howeverrecruitment strategies and shade toleranceneed to be further studied to draw signifi-cant conclusions During the Symposiumsite visits basket-tree harvesters and re-searchers both noted that basketry is moreprevalent in Maine New York and south-eastern Ontario Lower stand densities anddifferent recruitment patterns along withcultural traditions may influence the pro-portion of basket-quality black ash trees inthese regions compared with others withinthe black ash range

Unlike those in many other regionsblack ash trees in Maine frequently occur inmixed species stands which can includenorthern white-cedar balsam fir Americanelm (Ulmus americana L) red maple (Acerrubrum L) and yellow birch (Betula al-leghaniensis Britton) (Gawler and Cutko2010 Costanza 2015) Pure black ash standsare uncommon unless the site is a swamp oran old-growth black ash stand (Ehrenfeld2012 Costanza 2015) Of the sites visited

Table 1 Black ash average tree age in North America based on four studies that sampled a variety of black ash stands across thenorthern United States and southern Canada

Location StudyMaximum black ash

age (4 in dbh)Average black ashage (4 in dbh) No of black ash stands

(yr)

Maine Costanza 2015 273 59 24 stands (1frasl6 ac)Chippewa National Forest Minnesota Benedict and Frelich 2008 1641311351 9698651 67 stands (164 ft radius) 2424191

Northern New York2 Costanza 2015 146 70 16 stands (1frasl6 ac)Ontario Canada Townshend 2011 1613 61 16 stands (1frasl10 ac)Lake Duparquet Queacutebec Canada Tardif and Bergeron 1999 3204 34 5 stands (composed of 54 plots 1frasl40 ac each)

Research sites ranged from pure black ash to mixed forest stands and from old-growth to anthropogenically disturbed stands1 Values correspond to three types of sites lowlandsuplandswoodland ponds Maximum and average tree ages were taken from stand data that included all species however stands were dominatedby black ash A total of 166 black ash were sampled2 In northern New York samples were collected in St Lawrence and Lewis counties3 Oldest veteran black ash tree throughout sample area was 161 years old Oldest veteran black ash tree in each stand ranged from 63ndash161 years old4 The oldest black ash was sexually reproduced The oldest black ash sprout was 120 yr old

Journal of Forestry bull September 2017 439

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

during the Symposium the black ashswamps had moderate conifer densities (pri-marily balsam fir) whereas the hardwoodriver terraces were mixed hardwood standscomposed of red maple American basswood(Tilia americana L) green ash and whiteash (Fraxinus americana L) Harvesters andresearchers both noted the improved qualityof ash on the river terraces and often associ-ated these basket-quality trees with the lackof conifers innear the stand

At the end of the Symposium severalconclusions about Mainersquos black ash popu-lation emerged Unlike those in other re-gions of North America the four blackash stands we visited were early-successional

stands characterized by low to moderatestand densities mixed species compositionand reduced black ash regeneration In ad-dition harvesters confirmed that these char-acteristics (site hydrology species composi-tion and stand structure) play a key role inthe quality of black ash trees produced Withregard to management implications thesites visited were similar to stands in Ontariowhere black ash is a secondary or minorcomponent (Jeffrey Dech Nipissing Uni-versity pers comm Feb 2 2016) andmany green ash stands in southeasternMichigan (Kashian 2016) Therefore re-search from these stands may provide valu-able insight into how similar stands in

Maine and other regions are likely to re-spond to EAB infestations

The EAB ImpactThe potential economic and ecological

impacts of the EAB invasion are staggering(Kovacs et al 2010 Aukema et al 2011) andwill drastically alter the ash components ofNorth American forests (Cappaert et al2005 Herms and McCullough 2014)Black ash in particular is highly susceptibleto the insect (Anulewicz et al 2006 Pure-swaran and Poland 2009 Siegert et al2014a) and dies within 2ndash4 years of beinginfested (Siegert et al 2005 2014b) (Figure7) In EAB-infested areas research has yet tofully address the ecological impacts to blackash Several studies report ash mortality ratesas a pooled statistic including white greenand black ash (eg Kashian and Witter2011 Klooster et al 2014) whereas otherstudies only address early infestation periods(2004ndash2008) and focus on Michigan andOhio (eg Siegert et al 2005 Smith et al2015 Kashian 2016) Researchers in Min-nesota have focused on black ash standssimulating EAB mortality by girdling blackash trees (Slesak et al 2014 Looney et al2015) However additional research isneeded to more critically evaluate black ashstand dynamics through the course of anEAB invasion and in the different habitatsand hydrologic regimes where black ash oc-curs

Based on green ash studies conductedin Michigan after EAB infestation (Kashian2016) it is possible to make a few inferencesabout the potential response of black ashstands to EAB In Michigan green ash treesare persisting with sprouts released ad-vanced regeneration and seedlings pro-duced from sprout-borne seeds on siteswhere green ash densities were relativelyhigh Conversely preliminary work suggestssprouting from EAB-killed black ash may beextremely limited (NW Siegert USDAForest Service unpubl data 2010) Re-duced densities of black ash will probablyleave stands depleted with fewer and smallerstems following the initial wave of the EABinvasion Any surviving black ash trees willprobably be small (3 in dbh) unless estab-lished EAB infestations are radically sup-pressed One study in southeastern Michi-gan conducted when EAB populations wererelatively high found that sprouts from har-vested trees were recolonized by EAB within1 year (Petrice and Haack 2011) Furtherwork by Tardif and Bergeron (1999) indi-

Figure 3 Site 1 hardwood river terrace Site 1 is a sparse to moderately dense black ashmixed hardwood stand The stand is separated from a neighboring conifer stand by a smallstream (A) The majority of black ash gt4 in dbh at this site are high quality (B)

440 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

cates a greater abundance of black ashsprouts than seedlings in young age classesin addition to higher mortality rates insprouts To fully understand how black ashsprouts may persist in EAB-infested standsfurther investigation is needed on sproutingprocesses the survivorship of black ashsprouts the relationship to EAB populationcycles and the ability of these sprouts to be-come basket-quality trees

After a few years of EAB infestationonce the overstory has been killed estab-lished black ash seedlings if present may bereleased if the resulting gaps are not quicklyfilled in by neighboring trees However be-

cause EAB is known to infest and kill trees1 in dbh (Herms and McCullough2014) it is likely that most seedlings inmixed or low-density stands will not achievea larger size (remain 3 in dbh) and themean stand diameter will remain lowWhere black ash is a dominant componentof the stand changes in evapotranspirationafter overstory loss may result in water tablerise (Slesak et al 2014) and conversion toa more open-water wetland condition(Erdmann et al 1987) further threaten-ing ash persistence in the stand In addi-tion gap formation will probably result inan increased risk of competition from re-

sidual non-ash understory trees unaffectedby EAB as well as shrubs and herbaceousvegetation (Slesak et al 2014) Based onthe rapid detrimental impact of EAB it iscritical to identify what additional infor-mation and efforts are needed to preparefor the insectrsquos arrival in many black ashstands

RecommendationsThe first step in preparing for and man-

aging EAB is early detection If and when EABis detected efforts should focus on slowing theinsectrsquos spread (Siegert et al 2015) so the avail-ability of existing basket-quality ash trees canbe extended as long as possible Managementstrategies may be implemented to complementthis effort by reducing EAB populationdensities (McCullough et al 2015) In do-ing so guidelines should be developedthat limit anthropogenic spread of EAB-infested material while allowing harvestersto safely transport basket-quality black ashmaterial without risk of transporting EABto non-infested areas For instance tworecent studies reported that submerginginfested black ash logs in water for 8 ndash16weeks killed all EAB while still maintain-ing useable basket-quality wood (Siegertet al 2014a Poland et al 2015)

If harvesters are available basket-qual-ity trees should be harvested pounded andsplit as soon as an EAB infestation is de-tected in the vicinity This harvesting in-crease would have a considerably differentimpact on forests and on black ash in par-ticular than tribal traditions historicallyhad However dried black ash splints can bestored for several years (Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) increas-ing harvesting would allow harvesters to usewood that would otherwise be lost to EABLightly infested black ash trees remain suit-able for basketmaking materials but degraderapidly as EAB density increases reducingyields from a given log to the point where itmay be unusable for splints (Siegert et al2014a Poland et al 2015) In additionsome have recommended increasing blackash harvesting now before EAB arrives Thiswould potentially reduce concerns aboutquarantines accidental insect transport andloss of basket-quality wood after an EAB in-festation

Last we recommend collaboration be-tween state federal and tribal agencies todevelop memorandums of understanding orjoint agreements before the arrival of EAB

Figure 4 Site 2 black ash swamp Site 2 is a moderately dense mixed hardwood-coniferstand with black ash red maple and balsam fir The site is characterized by minimalponding (A) root buttressing (A) and small black ash trees typically lt6 in dbh (B) Themajority of black ash gt4 in dbh are poor quality

Journal of Forestry bull September 2017 441

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

in high-value black ash stands Such agree-ments can alleviate confusion and promoteaction as soon as EAB is detected in or neartribal lands or other traditional black ashharvesting sites Furthermore such agree-ments can help foster discussions on accessi-bility to black ash trees To preserve theblack ash resource and maintain black ashbasketry in the face of EAB discussions re-garding accessibility and response plansneed to occur

Future ResearchIn addition to making preparations

for EABrsquos arrival in black ash stands there

are countless opportunities to learn moreabout this still-overlooked tree species Toimprove our understanding of black ashtreesrsquo potential response to EAB and im-pacts to cultural traditions future researchdirections were discussed during the BlackAsh Symposium What follows is a summaryof the three most pressing research needsinfluence of black ash stand dynamics andsite hydrology on regeneration and qualityof black ash role of black ash genetics in thefuture of the population and EAB popula-tion dynamics in infested black ash standsand the effect on black ash regeneration

Influence of Stand Dynamics and Hy-drology on Regeneration and Quality

Very little information exists on blackash stand histories conditions and naturaldynamics in Maine For example it is un-clear whether Maine black ash stands un-dergo continuous recruitment and remainstocked over an extended period of time or ifblack ash is replaced by other species Natu-ral stand dynamics and succession are poorlyunderstood partially because there are fewold-growth stands in Maine Locating andstudying a few old-growth black ash stands(200 years old) would greatly improve ourcurrent understanding of succession Theinclusion of managed stands would alsoprove beneficial allowing researchers toquantify the response of black ash to silvicul-tural treatments Furthermore describingthe dynamics of ash across a range of distur-bance histories would allow researchers todevelop a much needed successional frame-work for black ash and to develop a descrip-tion of the speciesrsquo regeneration niche In themidwestern United States recent studieshave suggested that the future of black ashstands faced with EAB and climate changewill probably center around non-ash re-placement species (Looney et al 2015Iverson et al 2016) However in regionswhere black ash is a cultural keystone spe-cies replacing black ash is not an optimalmanagement strategy Describing black ashstand dynamics and recruitment in Maineas well as in other black ash habitats such asthose in Nova Scotia southern and centralOntario and Quebec would provide in-sight on how to maintain the species bothfor basketmaking and ecological reasons

Stand dynamics are heavily influencedby stand hydrology which is poorly under-stood in Maine and other parts of the blackash range A few recent studies in Minnesotahave explored black ash stand hydrology(Slesak et al 2014 Telander et al 2015)however the extent to which water tablefluctuations in black ash stands influence re-generation and recruitment across the entirerange should be further explored For exam-ple in situations where water table dynamicsare driven primarily by topography or localsurface water a change in evapotranspira-tion after overstory loss may not result in aconcomitant change in site hydrology (Rob-ert Slesak Minnesota Forest ResourcesCouncil pers comm June 5 2015) Fur-thermore the role of post-disturbancesprouting of black ash in regulating watertable dynamics is also unclear as is the role

Figure 5 Site 3 hardwood river terrace Site 3 is a moderately dense mixed hardwoodstand with occasional conifer (A) A small stream runs northeast to southwest and anunpaved road runs west to east through the stand In better-drained areas large high-quality black ash are produced Where ponding occurs black ash tend to be smaller andpoorer quality The majority of black ash gt4 in dbh are high quality (B)

442 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

Site HydrologyWhereas black ash is commonly associ-

ated with poorly drained wetland sites bas-ket-quality black ash is frequently found onsites where water flows or seeps through thesite it is not stagnant (Diamond and Emery2011 Costanza 2015 Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) Black ashoften grows more slowly with smaller an-nual growth rings when there is ponded wa-ter at the surface However soils that aresaturated from a nearby source via capillarityprovide ideal basket-quality conditions soilsare wet enough to allow black ash to out-compete other species but not overly satu-rated to inhibit radial growth (Telander et al2015 Robert Slesak Minnesota Forest Re-sources Council pers comm June 52015) Because of this sites that are wet be-cause of proximity to water or topography(eg swales) produce better-quality ash treesthan sites where hydrology is driven by aconstraining soil horizon or property whichis why some harvesters associate basket-qual-ity black ash with river terraces and flood-plains (Basket-tree harvesters Maine IndianBasketmakers Alliance pers comm MayndashAugust 2013)

Rooting DepthSites that are saturated later into the

growing season and are nutrient poor typi-cally do not produce basket-quality blackash If there are swollen root buttresses orroots that are exposed and easily visible itindicates that the tree and site are probablyinadequate for basketry These attributes

typically represent shallow rooting depth be-cause of impenetrable soils or high subsur-face water tables both of which producepoor-quality black ash with annual ringsthat are too narrow for basketry (Basket-treeharvesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013Symposium field observations November2014)

Annual Ring-WidthFor many harvesters annual ring-width

is the most important factor in determiningthe basket quality of black ash Good basket-quality wood contains growth rings 008inyear for 10 or more consecutive years(Benedict and Frelich 2008) but ideally an-nual growth is relatively consistent through-out the treersquos life If growth rings are toonarrow (eg 004 in) they are unusablefor weaving However growth rings that aretoo large (eg 020 in) are also undesir-able because they result in thick splints thatare difficult to work with (Basket-tree har-vesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013) Ahigh-quality basket tree typically produces8ndash12 splints per radial inch (ca 008ndash012in ring-widthyear)

Stand and Species CompositionA typical stand can contain both good-

and poor-quality black ash Whereas thereare specific site characteristics that often leadto the production of basket-quality ash trees(see above) microsite and stand characteris-tics are also important Light availability iscritical basket-quality ash trees tend to be

more open grown receiving intermediate tomoderate light Likewise microtopographyis also critical within a few inches dramaticchanges in soil water table depth and root-ing depth can be observed (Symposium fieldobservations November 2014)

Basket-quality ash trees are often asso-ciated with reduced competition from otherspecies which in turn increases light avail-ability (Symposium field observations No-vember 2014) Good sites frequently con-tain few if any conifers such as northernwhite-cedar (Thuja occidentalis L) and bal-sam fir (Abies balsamea [L] Mill) Whenconifers are present some harvesters haveobserved an increase in the brittleness ofblack ash wood collected from the site mak-ing their splints more likely to crack andbreak Furthermore common conifer asso-ciates of black ash (eg northern white-ce-dar balsam fir and red spruce [Picea rubensSarg]) can be indicative of poorly drainedsoil conditions which result in poor-qualityblack ash trees (Basket-tree harvestersMaine Indian Basketmakers Alliance perscomm MayndashAugust 2013)

CanopyTrees in the overstory (dominant or

codominant) with full deep crowns thatcover more than 50 of the height of thebole are preferred for basket-quality trees Ifdead limbs or epicormic branching are pres-ent the tree is considered unhealthy and of-ten avoided However dying trees are occa-sionally used if the annual ring-width(described earlier) looks acceptable (Basket-tree harvesters Maine Indian BasketmakersAlliance pers comm MayndashAugust 2013)

BarkTrees with increased bark texture and

bark depth are preferred however black ashrarely displays the distinct furrows seen ongreen or white ash (Diamond and Emery2011) If the bark takes on a ldquocorkyrdquo appear-ance (slightly flexible moist and sticks outfrom the bole) (Figure 2A) it is frequentlyassociated with annual growth rings ideal forbasketry (Basket-tree harvesters Maine In-dian Basketmakers Alliance pers commMayndashAugust 2013) One possible explana-tion for this association is that thicker xylemincrements imply good growth conditionsthat would also produce thicker phloem andcork from lateral meristems which wouldresult in corky bark Faster growth in lateralmeristems probably results from increasedlight availability and more favorable site hy-

Figure 1 Black ash basket Black ash baskets take many shapes and forms from sturdypack baskets to decorative fancy baskets Other baskets mix utility with decoration InMaine potato baskets are common including this basket by Richard Silliboy of the HoultonBand of Micmacs

Journal of Forestry bull September 2017 437

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

drology (Symposium field observations No-vember 2014) Bark characteristics were akey indicator for many basket-tree harvesterstaking part in the Symposium Further-more participants in the Symposium ob-served that increment borer holes healedfaster in trees with ldquocorkyrdquo bark (Figure 2A)than in those with ldquononcorkyrdquo bark (Figure2B) Bark color can also indicate tree qualityA yellow-tinted bark was often associatedwith increased annual growth and improvedwood quality whereas a gray-tinted barkwas associated with poorer quality and brit-tle wood

Tree AttributesSpecific features within the tree can af-

fect its usability The sapwood (aka ldquowhitewoodrdquo) is typically used for fancy basketswhereas the heartwood (aka ldquobrown woodrdquo)is commonly used for pack and potato bas-kets Stem defects such as cracks branchstubs or holes can make the wood unusablefor splints In addition if the tree bole is toocurved it cannot be processed (ldquopoundedrdquo)for splints A harvester can work with a mod-erate sweep in the bole but it takes moretime and effort to pound (Basket-tree har-vesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013)

A good-quality tree will have a straightbole for 45ndash6 ft (or more) and that sectionwill be free of defects Tree size can range

anywhere from 5 to 25 in dbh (breastheight 45 ft) but selection is highly de-pendent on how the tree will be transported(eg on the harvesterrsquos shoulder four-wheeler or truck) (Basket-tree harvestersMaine Indian Basketmakers Alliance perscomm MayndashAugust 2013)

General Black Ash BiologyBlack ash grows on a range of soil types

from peat bogs and mineral soil wetlands towell-drained stream edges and ridges (Simset al 1990 Wright and Rauscher 1990 Eh-renfeld 2012) The species is most fre-quently found on flooded or seasonally wetsites probably due to reduced competitionfrom other species incapable of establishingin hydric conditions Tardif et al (1994) re-ported that young black ash stems can pro-duce hypertrophied lenticels to increase gasexchange as well as use adventitious roots(Sena Gomes and Kozlowski 1980 Tardifet al 1994 Mendelssohn et al 2014) Bothof these adaptations allow young stems topersist on wet sites

When black ash becomes established onbetter-drained mesic sites the species fre-quently exhibits increased annual growthand larger dbh (Costanza 2015 pers observof authors 2015) Conversely in stands sat-urated throughout the growing seasonblack ash trees typically have narrower an-

nual growth rings reduced dbh exposedroots and numerous sprouts One explana-tion is that black ash is often classified ashaving a shallow root system (Wright andRauscher 1990) which is more affected bywater table fluctuations in turn affecting an-nual growth rates Although long-term satu-ration produces poor-quality ash sites withspring flooding followed by surface dryingin the summer favor black ash growth (Simset al 1990 Lenhart et al 2012 Slesak et al2014) These better-drained mesic sites aretherefore locations where basket-tree har-vesters frequently search for basket-qualitytrees Further these differences in site hy-drology that influence black ash growth andquality may be captured in the NaturalCommunity classifications that many statesand provinces have developed to describevegetation soils and site conditions of com-mon communities (see Supplemental TableS1 )

Black ash regeneration also varieswidely The species is capable of both pro-ducing seed and sprouting when cut orstressed yet sprouts from the root collartend to have a higher growth rate as they aregrowing from an already developed root sys-tem (Tardif and Bergeron 1999) Thesesprouts also tend to be more tolerant offlooding and droughts (Sims et al 1990Tardif and Bergeron 1999) Further siteconditions (eg flooding and drought) mayheavily influence seedling survival and estab-lishment Two consecutive dryreducedflood years may be needed for successful es-tablishment the first year allows the seed tobreak dormancy and germinate whereas thesecond year helps establishment and survivalof future flooding (Symposium field obser-vations November 2014) Ash can also pro-duce persistent seeds which is a benefit insites with fluctuating hydrologic regimesHowever several biotic or abiotic site factorsmay influence the survivorship of this seedbank including insects moisture fungi andothers (Klooster et al 2014)

Based on four recent studies averagetree age in black ash dominated stands typi-cally ranges from 35 to 100 years old with afew black ash trees greater than 200 years old(Table 1) (Tardif and Bergeron 1999 Bene-dict and Frelich 2008 Townshend 2011Costanza 2015) Although large old-growthblack ash forests (200 years old) exist inNorth America they are not common

Figure 2 Comparison of black ash bark quality Black ash bark takes on a variety ofappearances Basket-tree harvesters frequently associate spongy corky thicker bark (A)with high-quality basket wood Conversely bark with long thin brittle flakes (B) isfrequently associated with poor-quality black ash

Supplementary data are available with this article at httpdxdoiorg105849jof2016-034R1

438 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

In more disturbed areas (both naturallyand anthropogenically) average tree age isyounger Given basketmakersrsquo preferencefor 008ndash012 in ring-widths and trees5ndash25 in dbh this typical black ash age rangematches the typical expectations of basket-quality trees

Reports on black ash shade tolerancediffer One perspective describes the speciesas shade intolerant (Wright and Rauscher1990 Gustafson and Sturtevant 2013)whereas another describes the species asmoderately shade tolerant becoming moreintolerant with age (Sims et al 1990 Barnesand Wagner 2004) Tolerance may also de-pend on origin seedlings and sprouts mayhave different levels of shade toleranceBased on several recent observations (Reyeset al 2013 Fisichelli et al 2014 Looneyet al 2016 Symposium field observationsNovember 2014) the species appears to bemoderately shade tolerant Black ash is capa-ble of germinating and surviving in the un-derstory for several years although low tomoderate light is required for survival aclosed canopy will result in little to no seed-ling survivorship In Ontario Townshend(2011) found black ash trees of substantialage (eg up to 80 years old) under the mainforest canopy suggesting that the species iscapable of withstanding long periods of sup-pression If black ash can persist in the un-derstory the surviving seedlings could be animportant source of future black ash stemsduring the EAB invasion

Site characteristics regeneration strate-gies and stand dynamics all play a key role inthe quality of black ash trees and heavily in-fluence basketmaking traditions By im-proving our understanding of black ashbiologyecology in Maine and other blackash habitats improved management strate-

gies that simultaneously target EAB controland black ash preservation which will serveto maintain this cultural keystone speciescan be developed

Case Study Four Black AshStands in Maine

In November 2014 basket-tree har-vesters and researchers visited four black ashsites in Maine as part of the Black Ash Sym-posium These sites were representative oftwo common black ash stand types found ina statewide study black ash swamps andhardwood river terraces (Gawler and Cutko2010 Costanza 2015 see also SupplementalTable S1) two sites from each stand typewere visited (Figures 3ndash6) The site visitsallowed Symposium participants to sharetheir knowledge of black ash while also as-sessing the potential impact of EAB on thesetypes of stands and on basket-quality ashWhat follows is a summary of the discus-sions and observations from the Symposiumsite visits

The four black ash stands visited are ex-amples of old-field early-successional forests(100 years old) common in central andsouthern Maine which regenerate on aban-doned agricultural land The two black ashswamps were characterized by standing wa-ter ponding and relatively flat topographywith limited pits and mounds (Figures 4 and6) The two hardwood river terraces in con-trast had little to no standing water slightdownstream slopes and small streams drain-ing the sites (Figures 3 and 5) Based on in-crement cores collected in 2012 (Costanza2015) the highest basket-quality black ashtrees were found on the two river terracesites Harvesters attending the Symposiumsupported this finding noting that they of-

ten associate high-quality black ash withriver banks and floodplains with low tomoderate stand densities

The four Maine stands had significantlylower black ash densities than documentedstands found in Minnesota Ontario andQuebec (Table 2) Stands visited rangedfrom 90ndash162 black ash stemsac1 with theswamps producing higher black ash densi-ties than the river terraces One possible ex-planation for these reduced stand densities isthat most Maine stands are early-succes-sional forests growing in previously har-vested stands or old fields Studies outside ofMaine have focused on well-established for-ests where black ash maintains high densitiesand canopy dominance It is also possiblethat black ash trees in Maine recruit differ-ently from those in other regions howeverrecruitment strategies and shade toleranceneed to be further studied to draw signifi-cant conclusions During the Symposiumsite visits basket-tree harvesters and re-searchers both noted that basketry is moreprevalent in Maine New York and south-eastern Ontario Lower stand densities anddifferent recruitment patterns along withcultural traditions may influence the pro-portion of basket-quality black ash trees inthese regions compared with others withinthe black ash range

Unlike those in many other regionsblack ash trees in Maine frequently occur inmixed species stands which can includenorthern white-cedar balsam fir Americanelm (Ulmus americana L) red maple (Acerrubrum L) and yellow birch (Betula al-leghaniensis Britton) (Gawler and Cutko2010 Costanza 2015) Pure black ash standsare uncommon unless the site is a swamp oran old-growth black ash stand (Ehrenfeld2012 Costanza 2015) Of the sites visited

Table 1 Black ash average tree age in North America based on four studies that sampled a variety of black ash stands across thenorthern United States and southern Canada

Location StudyMaximum black ash

age (4 in dbh)Average black ashage (4 in dbh) No of black ash stands

(yr)

Maine Costanza 2015 273 59 24 stands (1frasl6 ac)Chippewa National Forest Minnesota Benedict and Frelich 2008 1641311351 9698651 67 stands (164 ft radius) 2424191

Northern New York2 Costanza 2015 146 70 16 stands (1frasl6 ac)Ontario Canada Townshend 2011 1613 61 16 stands (1frasl10 ac)Lake Duparquet Queacutebec Canada Tardif and Bergeron 1999 3204 34 5 stands (composed of 54 plots 1frasl40 ac each)

Research sites ranged from pure black ash to mixed forest stands and from old-growth to anthropogenically disturbed stands1 Values correspond to three types of sites lowlandsuplandswoodland ponds Maximum and average tree ages were taken from stand data that included all species however stands were dominatedby black ash A total of 166 black ash were sampled2 In northern New York samples were collected in St Lawrence and Lewis counties3 Oldest veteran black ash tree throughout sample area was 161 years old Oldest veteran black ash tree in each stand ranged from 63ndash161 years old4 The oldest black ash was sexually reproduced The oldest black ash sprout was 120 yr old

Journal of Forestry bull September 2017 439

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

during the Symposium the black ashswamps had moderate conifer densities (pri-marily balsam fir) whereas the hardwoodriver terraces were mixed hardwood standscomposed of red maple American basswood(Tilia americana L) green ash and whiteash (Fraxinus americana L) Harvesters andresearchers both noted the improved qualityof ash on the river terraces and often associ-ated these basket-quality trees with the lackof conifers innear the stand

At the end of the Symposium severalconclusions about Mainersquos black ash popu-lation emerged Unlike those in other re-gions of North America the four blackash stands we visited were early-successional

stands characterized by low to moderatestand densities mixed species compositionand reduced black ash regeneration In ad-dition harvesters confirmed that these char-acteristics (site hydrology species composi-tion and stand structure) play a key role inthe quality of black ash trees produced Withregard to management implications thesites visited were similar to stands in Ontariowhere black ash is a secondary or minorcomponent (Jeffrey Dech Nipissing Uni-versity pers comm Feb 2 2016) andmany green ash stands in southeasternMichigan (Kashian 2016) Therefore re-search from these stands may provide valu-able insight into how similar stands in

Maine and other regions are likely to re-spond to EAB infestations

The EAB ImpactThe potential economic and ecological

impacts of the EAB invasion are staggering(Kovacs et al 2010 Aukema et al 2011) andwill drastically alter the ash components ofNorth American forests (Cappaert et al2005 Herms and McCullough 2014)Black ash in particular is highly susceptibleto the insect (Anulewicz et al 2006 Pure-swaran and Poland 2009 Siegert et al2014a) and dies within 2ndash4 years of beinginfested (Siegert et al 2005 2014b) (Figure7) In EAB-infested areas research has yet tofully address the ecological impacts to blackash Several studies report ash mortality ratesas a pooled statistic including white greenand black ash (eg Kashian and Witter2011 Klooster et al 2014) whereas otherstudies only address early infestation periods(2004ndash2008) and focus on Michigan andOhio (eg Siegert et al 2005 Smith et al2015 Kashian 2016) Researchers in Min-nesota have focused on black ash standssimulating EAB mortality by girdling blackash trees (Slesak et al 2014 Looney et al2015) However additional research isneeded to more critically evaluate black ashstand dynamics through the course of anEAB invasion and in the different habitatsand hydrologic regimes where black ash oc-curs

Based on green ash studies conductedin Michigan after EAB infestation (Kashian2016) it is possible to make a few inferencesabout the potential response of black ashstands to EAB In Michigan green ash treesare persisting with sprouts released ad-vanced regeneration and seedlings pro-duced from sprout-borne seeds on siteswhere green ash densities were relativelyhigh Conversely preliminary work suggestssprouting from EAB-killed black ash may beextremely limited (NW Siegert USDAForest Service unpubl data 2010) Re-duced densities of black ash will probablyleave stands depleted with fewer and smallerstems following the initial wave of the EABinvasion Any surviving black ash trees willprobably be small (3 in dbh) unless estab-lished EAB infestations are radically sup-pressed One study in southeastern Michi-gan conducted when EAB populations wererelatively high found that sprouts from har-vested trees were recolonized by EAB within1 year (Petrice and Haack 2011) Furtherwork by Tardif and Bergeron (1999) indi-

Figure 3 Site 1 hardwood river terrace Site 1 is a sparse to moderately dense black ashmixed hardwood stand The stand is separated from a neighboring conifer stand by a smallstream (A) The majority of black ash gt4 in dbh at this site are high quality (B)

440 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

cates a greater abundance of black ashsprouts than seedlings in young age classesin addition to higher mortality rates insprouts To fully understand how black ashsprouts may persist in EAB-infested standsfurther investigation is needed on sproutingprocesses the survivorship of black ashsprouts the relationship to EAB populationcycles and the ability of these sprouts to be-come basket-quality trees

After a few years of EAB infestationonce the overstory has been killed estab-lished black ash seedlings if present may bereleased if the resulting gaps are not quicklyfilled in by neighboring trees However be-

cause EAB is known to infest and kill trees1 in dbh (Herms and McCullough2014) it is likely that most seedlings inmixed or low-density stands will not achievea larger size (remain 3 in dbh) and themean stand diameter will remain lowWhere black ash is a dominant componentof the stand changes in evapotranspirationafter overstory loss may result in water tablerise (Slesak et al 2014) and conversion toa more open-water wetland condition(Erdmann et al 1987) further threaten-ing ash persistence in the stand In addi-tion gap formation will probably result inan increased risk of competition from re-

sidual non-ash understory trees unaffectedby EAB as well as shrubs and herbaceousvegetation (Slesak et al 2014) Based onthe rapid detrimental impact of EAB it iscritical to identify what additional infor-mation and efforts are needed to preparefor the insectrsquos arrival in many black ashstands

RecommendationsThe first step in preparing for and man-

aging EAB is early detection If and when EABis detected efforts should focus on slowing theinsectrsquos spread (Siegert et al 2015) so the avail-ability of existing basket-quality ash trees canbe extended as long as possible Managementstrategies may be implemented to complementthis effort by reducing EAB populationdensities (McCullough et al 2015) In do-ing so guidelines should be developedthat limit anthropogenic spread of EAB-infested material while allowing harvestersto safely transport basket-quality black ashmaterial without risk of transporting EABto non-infested areas For instance tworecent studies reported that submerginginfested black ash logs in water for 8 ndash16weeks killed all EAB while still maintain-ing useable basket-quality wood (Siegertet al 2014a Poland et al 2015)

If harvesters are available basket-qual-ity trees should be harvested pounded andsplit as soon as an EAB infestation is de-tected in the vicinity This harvesting in-crease would have a considerably differentimpact on forests and on black ash in par-ticular than tribal traditions historicallyhad However dried black ash splints can bestored for several years (Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) increas-ing harvesting would allow harvesters to usewood that would otherwise be lost to EABLightly infested black ash trees remain suit-able for basketmaking materials but degraderapidly as EAB density increases reducingyields from a given log to the point where itmay be unusable for splints (Siegert et al2014a Poland et al 2015) In additionsome have recommended increasing blackash harvesting now before EAB arrives Thiswould potentially reduce concerns aboutquarantines accidental insect transport andloss of basket-quality wood after an EAB in-festation

Last we recommend collaboration be-tween state federal and tribal agencies todevelop memorandums of understanding orjoint agreements before the arrival of EAB

Figure 4 Site 2 black ash swamp Site 2 is a moderately dense mixed hardwood-coniferstand with black ash red maple and balsam fir The site is characterized by minimalponding (A) root buttressing (A) and small black ash trees typically lt6 in dbh (B) Themajority of black ash gt4 in dbh are poor quality

Journal of Forestry bull September 2017 441

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

in high-value black ash stands Such agree-ments can alleviate confusion and promoteaction as soon as EAB is detected in or neartribal lands or other traditional black ashharvesting sites Furthermore such agree-ments can help foster discussions on accessi-bility to black ash trees To preserve theblack ash resource and maintain black ashbasketry in the face of EAB discussions re-garding accessibility and response plansneed to occur

Future ResearchIn addition to making preparations

for EABrsquos arrival in black ash stands there

are countless opportunities to learn moreabout this still-overlooked tree species Toimprove our understanding of black ashtreesrsquo potential response to EAB and im-pacts to cultural traditions future researchdirections were discussed during the BlackAsh Symposium What follows is a summaryof the three most pressing research needsinfluence of black ash stand dynamics andsite hydrology on regeneration and qualityof black ash role of black ash genetics in thefuture of the population and EAB popula-tion dynamics in infested black ash standsand the effect on black ash regeneration

Influence of Stand Dynamics and Hy-drology on Regeneration and Quality

Very little information exists on blackash stand histories conditions and naturaldynamics in Maine For example it is un-clear whether Maine black ash stands un-dergo continuous recruitment and remainstocked over an extended period of time or ifblack ash is replaced by other species Natu-ral stand dynamics and succession are poorlyunderstood partially because there are fewold-growth stands in Maine Locating andstudying a few old-growth black ash stands(200 years old) would greatly improve ourcurrent understanding of succession Theinclusion of managed stands would alsoprove beneficial allowing researchers toquantify the response of black ash to silvicul-tural treatments Furthermore describingthe dynamics of ash across a range of distur-bance histories would allow researchers todevelop a much needed successional frame-work for black ash and to develop a descrip-tion of the speciesrsquo regeneration niche In themidwestern United States recent studieshave suggested that the future of black ashstands faced with EAB and climate changewill probably center around non-ash re-placement species (Looney et al 2015Iverson et al 2016) However in regionswhere black ash is a cultural keystone spe-cies replacing black ash is not an optimalmanagement strategy Describing black ashstand dynamics and recruitment in Maineas well as in other black ash habitats such asthose in Nova Scotia southern and centralOntario and Quebec would provide in-sight on how to maintain the species bothfor basketmaking and ecological reasons

Stand dynamics are heavily influencedby stand hydrology which is poorly under-stood in Maine and other parts of the blackash range A few recent studies in Minnesotahave explored black ash stand hydrology(Slesak et al 2014 Telander et al 2015)however the extent to which water tablefluctuations in black ash stands influence re-generation and recruitment across the entirerange should be further explored For exam-ple in situations where water table dynamicsare driven primarily by topography or localsurface water a change in evapotranspira-tion after overstory loss may not result in aconcomitant change in site hydrology (Rob-ert Slesak Minnesota Forest ResourcesCouncil pers comm June 5 2015) Fur-thermore the role of post-disturbancesprouting of black ash in regulating watertable dynamics is also unclear as is the role

Figure 5 Site 3 hardwood river terrace Site 3 is a moderately dense mixed hardwoodstand with occasional conifer (A) A small stream runs northeast to southwest and anunpaved road runs west to east through the stand In better-drained areas large high-quality black ash are produced Where ponding occurs black ash tend to be smaller andpoorer quality The majority of black ash gt4 in dbh are high quality (B)

442 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

drology (Symposium field observations No-vember 2014) Bark characteristics were akey indicator for many basket-tree harvesterstaking part in the Symposium Further-more participants in the Symposium ob-served that increment borer holes healedfaster in trees with ldquocorkyrdquo bark (Figure 2A)than in those with ldquononcorkyrdquo bark (Figure2B) Bark color can also indicate tree qualityA yellow-tinted bark was often associatedwith increased annual growth and improvedwood quality whereas a gray-tinted barkwas associated with poorer quality and brit-tle wood

Tree AttributesSpecific features within the tree can af-

fect its usability The sapwood (aka ldquowhitewoodrdquo) is typically used for fancy basketswhereas the heartwood (aka ldquobrown woodrdquo)is commonly used for pack and potato bas-kets Stem defects such as cracks branchstubs or holes can make the wood unusablefor splints In addition if the tree bole is toocurved it cannot be processed (ldquopoundedrdquo)for splints A harvester can work with a mod-erate sweep in the bole but it takes moretime and effort to pound (Basket-tree har-vesters Maine Indian Basketmakers Alli-ance pers comm MayndashAugust 2013)

A good-quality tree will have a straightbole for 45ndash6 ft (or more) and that sectionwill be free of defects Tree size can range

anywhere from 5 to 25 in dbh (breastheight 45 ft) but selection is highly de-pendent on how the tree will be transported(eg on the harvesterrsquos shoulder four-wheeler or truck) (Basket-tree harvestersMaine Indian Basketmakers Alliance perscomm MayndashAugust 2013)

General Black Ash BiologyBlack ash grows on a range of soil types

from peat bogs and mineral soil wetlands towell-drained stream edges and ridges (Simset al 1990 Wright and Rauscher 1990 Eh-renfeld 2012) The species is most fre-quently found on flooded or seasonally wetsites probably due to reduced competitionfrom other species incapable of establishingin hydric conditions Tardif et al (1994) re-ported that young black ash stems can pro-duce hypertrophied lenticels to increase gasexchange as well as use adventitious roots(Sena Gomes and Kozlowski 1980 Tardifet al 1994 Mendelssohn et al 2014) Bothof these adaptations allow young stems topersist on wet sites

When black ash becomes established onbetter-drained mesic sites the species fre-quently exhibits increased annual growthand larger dbh (Costanza 2015 pers observof authors 2015) Conversely in stands sat-urated throughout the growing seasonblack ash trees typically have narrower an-

nual growth rings reduced dbh exposedroots and numerous sprouts One explana-tion is that black ash is often classified ashaving a shallow root system (Wright andRauscher 1990) which is more affected bywater table fluctuations in turn affecting an-nual growth rates Although long-term satu-ration produces poor-quality ash sites withspring flooding followed by surface dryingin the summer favor black ash growth (Simset al 1990 Lenhart et al 2012 Slesak et al2014) These better-drained mesic sites aretherefore locations where basket-tree har-vesters frequently search for basket-qualitytrees Further these differences in site hy-drology that influence black ash growth andquality may be captured in the NaturalCommunity classifications that many statesand provinces have developed to describevegetation soils and site conditions of com-mon communities (see Supplemental TableS1 )

Black ash regeneration also varieswidely The species is capable of both pro-ducing seed and sprouting when cut orstressed yet sprouts from the root collartend to have a higher growth rate as they aregrowing from an already developed root sys-tem (Tardif and Bergeron 1999) Thesesprouts also tend to be more tolerant offlooding and droughts (Sims et al 1990Tardif and Bergeron 1999) Further siteconditions (eg flooding and drought) mayheavily influence seedling survival and estab-lishment Two consecutive dryreducedflood years may be needed for successful es-tablishment the first year allows the seed tobreak dormancy and germinate whereas thesecond year helps establishment and survivalof future flooding (Symposium field obser-vations November 2014) Ash can also pro-duce persistent seeds which is a benefit insites with fluctuating hydrologic regimesHowever several biotic or abiotic site factorsmay influence the survivorship of this seedbank including insects moisture fungi andothers (Klooster et al 2014)

Based on four recent studies averagetree age in black ash dominated stands typi-cally ranges from 35 to 100 years old with afew black ash trees greater than 200 years old(Table 1) (Tardif and Bergeron 1999 Bene-dict and Frelich 2008 Townshend 2011Costanza 2015) Although large old-growthblack ash forests (200 years old) exist inNorth America they are not common

Figure 2 Comparison of black ash bark quality Black ash bark takes on a variety ofappearances Basket-tree harvesters frequently associate spongy corky thicker bark (A)with high-quality basket wood Conversely bark with long thin brittle flakes (B) isfrequently associated with poor-quality black ash

Supplementary data are available with this article at httpdxdoiorg105849jof2016-034R1

438 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

In more disturbed areas (both naturallyand anthropogenically) average tree age isyounger Given basketmakersrsquo preferencefor 008ndash012 in ring-widths and trees5ndash25 in dbh this typical black ash age rangematches the typical expectations of basket-quality trees

Reports on black ash shade tolerancediffer One perspective describes the speciesas shade intolerant (Wright and Rauscher1990 Gustafson and Sturtevant 2013)whereas another describes the species asmoderately shade tolerant becoming moreintolerant with age (Sims et al 1990 Barnesand Wagner 2004) Tolerance may also de-pend on origin seedlings and sprouts mayhave different levels of shade toleranceBased on several recent observations (Reyeset al 2013 Fisichelli et al 2014 Looneyet al 2016 Symposium field observationsNovember 2014) the species appears to bemoderately shade tolerant Black ash is capa-ble of germinating and surviving in the un-derstory for several years although low tomoderate light is required for survival aclosed canopy will result in little to no seed-ling survivorship In Ontario Townshend(2011) found black ash trees of substantialage (eg up to 80 years old) under the mainforest canopy suggesting that the species iscapable of withstanding long periods of sup-pression If black ash can persist in the un-derstory the surviving seedlings could be animportant source of future black ash stemsduring the EAB invasion

Site characteristics regeneration strate-gies and stand dynamics all play a key role inthe quality of black ash trees and heavily in-fluence basketmaking traditions By im-proving our understanding of black ashbiologyecology in Maine and other blackash habitats improved management strate-

gies that simultaneously target EAB controland black ash preservation which will serveto maintain this cultural keystone speciescan be developed

Case Study Four Black AshStands in Maine

In November 2014 basket-tree har-vesters and researchers visited four black ashsites in Maine as part of the Black Ash Sym-posium These sites were representative oftwo common black ash stand types found ina statewide study black ash swamps andhardwood river terraces (Gawler and Cutko2010 Costanza 2015 see also SupplementalTable S1) two sites from each stand typewere visited (Figures 3ndash6) The site visitsallowed Symposium participants to sharetheir knowledge of black ash while also as-sessing the potential impact of EAB on thesetypes of stands and on basket-quality ashWhat follows is a summary of the discus-sions and observations from the Symposiumsite visits

The four black ash stands visited are ex-amples of old-field early-successional forests(100 years old) common in central andsouthern Maine which regenerate on aban-doned agricultural land The two black ashswamps were characterized by standing wa-ter ponding and relatively flat topographywith limited pits and mounds (Figures 4 and6) The two hardwood river terraces in con-trast had little to no standing water slightdownstream slopes and small streams drain-ing the sites (Figures 3 and 5) Based on in-crement cores collected in 2012 (Costanza2015) the highest basket-quality black ashtrees were found on the two river terracesites Harvesters attending the Symposiumsupported this finding noting that they of-

ten associate high-quality black ash withriver banks and floodplains with low tomoderate stand densities

The four Maine stands had significantlylower black ash densities than documentedstands found in Minnesota Ontario andQuebec (Table 2) Stands visited rangedfrom 90ndash162 black ash stemsac1 with theswamps producing higher black ash densi-ties than the river terraces One possible ex-planation for these reduced stand densities isthat most Maine stands are early-succes-sional forests growing in previously har-vested stands or old fields Studies outside ofMaine have focused on well-established for-ests where black ash maintains high densitiesand canopy dominance It is also possiblethat black ash trees in Maine recruit differ-ently from those in other regions howeverrecruitment strategies and shade toleranceneed to be further studied to draw signifi-cant conclusions During the Symposiumsite visits basket-tree harvesters and re-searchers both noted that basketry is moreprevalent in Maine New York and south-eastern Ontario Lower stand densities anddifferent recruitment patterns along withcultural traditions may influence the pro-portion of basket-quality black ash trees inthese regions compared with others withinthe black ash range

Unlike those in many other regionsblack ash trees in Maine frequently occur inmixed species stands which can includenorthern white-cedar balsam fir Americanelm (Ulmus americana L) red maple (Acerrubrum L) and yellow birch (Betula al-leghaniensis Britton) (Gawler and Cutko2010 Costanza 2015) Pure black ash standsare uncommon unless the site is a swamp oran old-growth black ash stand (Ehrenfeld2012 Costanza 2015) Of the sites visited

Table 1 Black ash average tree age in North America based on four studies that sampled a variety of black ash stands across thenorthern United States and southern Canada

Location StudyMaximum black ash

age (4 in dbh)Average black ashage (4 in dbh) No of black ash stands

(yr)

Maine Costanza 2015 273 59 24 stands (1frasl6 ac)Chippewa National Forest Minnesota Benedict and Frelich 2008 1641311351 9698651 67 stands (164 ft radius) 2424191

Northern New York2 Costanza 2015 146 70 16 stands (1frasl6 ac)Ontario Canada Townshend 2011 1613 61 16 stands (1frasl10 ac)Lake Duparquet Queacutebec Canada Tardif and Bergeron 1999 3204 34 5 stands (composed of 54 plots 1frasl40 ac each)

Research sites ranged from pure black ash to mixed forest stands and from old-growth to anthropogenically disturbed stands1 Values correspond to three types of sites lowlandsuplandswoodland ponds Maximum and average tree ages were taken from stand data that included all species however stands were dominatedby black ash A total of 166 black ash were sampled2 In northern New York samples were collected in St Lawrence and Lewis counties3 Oldest veteran black ash tree throughout sample area was 161 years old Oldest veteran black ash tree in each stand ranged from 63ndash161 years old4 The oldest black ash was sexually reproduced The oldest black ash sprout was 120 yr old

Journal of Forestry bull September 2017 439

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

during the Symposium the black ashswamps had moderate conifer densities (pri-marily balsam fir) whereas the hardwoodriver terraces were mixed hardwood standscomposed of red maple American basswood(Tilia americana L) green ash and whiteash (Fraxinus americana L) Harvesters andresearchers both noted the improved qualityof ash on the river terraces and often associ-ated these basket-quality trees with the lackof conifers innear the stand

At the end of the Symposium severalconclusions about Mainersquos black ash popu-lation emerged Unlike those in other re-gions of North America the four blackash stands we visited were early-successional

stands characterized by low to moderatestand densities mixed species compositionand reduced black ash regeneration In ad-dition harvesters confirmed that these char-acteristics (site hydrology species composi-tion and stand structure) play a key role inthe quality of black ash trees produced Withregard to management implications thesites visited were similar to stands in Ontariowhere black ash is a secondary or minorcomponent (Jeffrey Dech Nipissing Uni-versity pers comm Feb 2 2016) andmany green ash stands in southeasternMichigan (Kashian 2016) Therefore re-search from these stands may provide valu-able insight into how similar stands in

Maine and other regions are likely to re-spond to EAB infestations

The EAB ImpactThe potential economic and ecological

impacts of the EAB invasion are staggering(Kovacs et al 2010 Aukema et al 2011) andwill drastically alter the ash components ofNorth American forests (Cappaert et al2005 Herms and McCullough 2014)Black ash in particular is highly susceptibleto the insect (Anulewicz et al 2006 Pure-swaran and Poland 2009 Siegert et al2014a) and dies within 2ndash4 years of beinginfested (Siegert et al 2005 2014b) (Figure7) In EAB-infested areas research has yet tofully address the ecological impacts to blackash Several studies report ash mortality ratesas a pooled statistic including white greenand black ash (eg Kashian and Witter2011 Klooster et al 2014) whereas otherstudies only address early infestation periods(2004ndash2008) and focus on Michigan andOhio (eg Siegert et al 2005 Smith et al2015 Kashian 2016) Researchers in Min-nesota have focused on black ash standssimulating EAB mortality by girdling blackash trees (Slesak et al 2014 Looney et al2015) However additional research isneeded to more critically evaluate black ashstand dynamics through the course of anEAB invasion and in the different habitatsand hydrologic regimes where black ash oc-curs

Based on green ash studies conductedin Michigan after EAB infestation (Kashian2016) it is possible to make a few inferencesabout the potential response of black ashstands to EAB In Michigan green ash treesare persisting with sprouts released ad-vanced regeneration and seedlings pro-duced from sprout-borne seeds on siteswhere green ash densities were relativelyhigh Conversely preliminary work suggestssprouting from EAB-killed black ash may beextremely limited (NW Siegert USDAForest Service unpubl data 2010) Re-duced densities of black ash will probablyleave stands depleted with fewer and smallerstems following the initial wave of the EABinvasion Any surviving black ash trees willprobably be small (3 in dbh) unless estab-lished EAB infestations are radically sup-pressed One study in southeastern Michi-gan conducted when EAB populations wererelatively high found that sprouts from har-vested trees were recolonized by EAB within1 year (Petrice and Haack 2011) Furtherwork by Tardif and Bergeron (1999) indi-

Figure 3 Site 1 hardwood river terrace Site 1 is a sparse to moderately dense black ashmixed hardwood stand The stand is separated from a neighboring conifer stand by a smallstream (A) The majority of black ash gt4 in dbh at this site are high quality (B)

440 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

cates a greater abundance of black ashsprouts than seedlings in young age classesin addition to higher mortality rates insprouts To fully understand how black ashsprouts may persist in EAB-infested standsfurther investigation is needed on sproutingprocesses the survivorship of black ashsprouts the relationship to EAB populationcycles and the ability of these sprouts to be-come basket-quality trees

After a few years of EAB infestationonce the overstory has been killed estab-lished black ash seedlings if present may bereleased if the resulting gaps are not quicklyfilled in by neighboring trees However be-

cause EAB is known to infest and kill trees1 in dbh (Herms and McCullough2014) it is likely that most seedlings inmixed or low-density stands will not achievea larger size (remain 3 in dbh) and themean stand diameter will remain lowWhere black ash is a dominant componentof the stand changes in evapotranspirationafter overstory loss may result in water tablerise (Slesak et al 2014) and conversion toa more open-water wetland condition(Erdmann et al 1987) further threaten-ing ash persistence in the stand In addi-tion gap formation will probably result inan increased risk of competition from re-

sidual non-ash understory trees unaffectedby EAB as well as shrubs and herbaceousvegetation (Slesak et al 2014) Based onthe rapid detrimental impact of EAB it iscritical to identify what additional infor-mation and efforts are needed to preparefor the insectrsquos arrival in many black ashstands

RecommendationsThe first step in preparing for and man-

aging EAB is early detection If and when EABis detected efforts should focus on slowing theinsectrsquos spread (Siegert et al 2015) so the avail-ability of existing basket-quality ash trees canbe extended as long as possible Managementstrategies may be implemented to complementthis effort by reducing EAB populationdensities (McCullough et al 2015) In do-ing so guidelines should be developedthat limit anthropogenic spread of EAB-infested material while allowing harvestersto safely transport basket-quality black ashmaterial without risk of transporting EABto non-infested areas For instance tworecent studies reported that submerginginfested black ash logs in water for 8 ndash16weeks killed all EAB while still maintain-ing useable basket-quality wood (Siegertet al 2014a Poland et al 2015)

If harvesters are available basket-qual-ity trees should be harvested pounded andsplit as soon as an EAB infestation is de-tected in the vicinity This harvesting in-crease would have a considerably differentimpact on forests and on black ash in par-ticular than tribal traditions historicallyhad However dried black ash splints can bestored for several years (Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) increas-ing harvesting would allow harvesters to usewood that would otherwise be lost to EABLightly infested black ash trees remain suit-able for basketmaking materials but degraderapidly as EAB density increases reducingyields from a given log to the point where itmay be unusable for splints (Siegert et al2014a Poland et al 2015) In additionsome have recommended increasing blackash harvesting now before EAB arrives Thiswould potentially reduce concerns aboutquarantines accidental insect transport andloss of basket-quality wood after an EAB in-festation

Last we recommend collaboration be-tween state federal and tribal agencies todevelop memorandums of understanding orjoint agreements before the arrival of EAB

Figure 4 Site 2 black ash swamp Site 2 is a moderately dense mixed hardwood-coniferstand with black ash red maple and balsam fir The site is characterized by minimalponding (A) root buttressing (A) and small black ash trees typically lt6 in dbh (B) Themajority of black ash gt4 in dbh are poor quality

Journal of Forestry bull September 2017 441

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

in high-value black ash stands Such agree-ments can alleviate confusion and promoteaction as soon as EAB is detected in or neartribal lands or other traditional black ashharvesting sites Furthermore such agree-ments can help foster discussions on accessi-bility to black ash trees To preserve theblack ash resource and maintain black ashbasketry in the face of EAB discussions re-garding accessibility and response plansneed to occur

Future ResearchIn addition to making preparations

for EABrsquos arrival in black ash stands there

are countless opportunities to learn moreabout this still-overlooked tree species Toimprove our understanding of black ashtreesrsquo potential response to EAB and im-pacts to cultural traditions future researchdirections were discussed during the BlackAsh Symposium What follows is a summaryof the three most pressing research needsinfluence of black ash stand dynamics andsite hydrology on regeneration and qualityof black ash role of black ash genetics in thefuture of the population and EAB popula-tion dynamics in infested black ash standsand the effect on black ash regeneration

Influence of Stand Dynamics and Hy-drology on Regeneration and Quality

Very little information exists on blackash stand histories conditions and naturaldynamics in Maine For example it is un-clear whether Maine black ash stands un-dergo continuous recruitment and remainstocked over an extended period of time or ifblack ash is replaced by other species Natu-ral stand dynamics and succession are poorlyunderstood partially because there are fewold-growth stands in Maine Locating andstudying a few old-growth black ash stands(200 years old) would greatly improve ourcurrent understanding of succession Theinclusion of managed stands would alsoprove beneficial allowing researchers toquantify the response of black ash to silvicul-tural treatments Furthermore describingthe dynamics of ash across a range of distur-bance histories would allow researchers todevelop a much needed successional frame-work for black ash and to develop a descrip-tion of the speciesrsquo regeneration niche In themidwestern United States recent studieshave suggested that the future of black ashstands faced with EAB and climate changewill probably center around non-ash re-placement species (Looney et al 2015Iverson et al 2016) However in regionswhere black ash is a cultural keystone spe-cies replacing black ash is not an optimalmanagement strategy Describing black ashstand dynamics and recruitment in Maineas well as in other black ash habitats such asthose in Nova Scotia southern and centralOntario and Quebec would provide in-sight on how to maintain the species bothfor basketmaking and ecological reasons

Stand dynamics are heavily influencedby stand hydrology which is poorly under-stood in Maine and other parts of the blackash range A few recent studies in Minnesotahave explored black ash stand hydrology(Slesak et al 2014 Telander et al 2015)however the extent to which water tablefluctuations in black ash stands influence re-generation and recruitment across the entirerange should be further explored For exam-ple in situations where water table dynamicsare driven primarily by topography or localsurface water a change in evapotranspira-tion after overstory loss may not result in aconcomitant change in site hydrology (Rob-ert Slesak Minnesota Forest ResourcesCouncil pers comm June 5 2015) Fur-thermore the role of post-disturbancesprouting of black ash in regulating watertable dynamics is also unclear as is the role

Figure 5 Site 3 hardwood river terrace Site 3 is a moderately dense mixed hardwoodstand with occasional conifer (A) A small stream runs northeast to southwest and anunpaved road runs west to east through the stand In better-drained areas large high-quality black ash are produced Where ponding occurs black ash tend to be smaller andpoorer quality The majority of black ash gt4 in dbh are high quality (B)

442 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

In more disturbed areas (both naturallyand anthropogenically) average tree age isyounger Given basketmakersrsquo preferencefor 008ndash012 in ring-widths and trees5ndash25 in dbh this typical black ash age rangematches the typical expectations of basket-quality trees

Reports on black ash shade tolerancediffer One perspective describes the speciesas shade intolerant (Wright and Rauscher1990 Gustafson and Sturtevant 2013)whereas another describes the species asmoderately shade tolerant becoming moreintolerant with age (Sims et al 1990 Barnesand Wagner 2004) Tolerance may also de-pend on origin seedlings and sprouts mayhave different levels of shade toleranceBased on several recent observations (Reyeset al 2013 Fisichelli et al 2014 Looneyet al 2016 Symposium field observationsNovember 2014) the species appears to bemoderately shade tolerant Black ash is capa-ble of germinating and surviving in the un-derstory for several years although low tomoderate light is required for survival aclosed canopy will result in little to no seed-ling survivorship In Ontario Townshend(2011) found black ash trees of substantialage (eg up to 80 years old) under the mainforest canopy suggesting that the species iscapable of withstanding long periods of sup-pression If black ash can persist in the un-derstory the surviving seedlings could be animportant source of future black ash stemsduring the EAB invasion

Site characteristics regeneration strate-gies and stand dynamics all play a key role inthe quality of black ash trees and heavily in-fluence basketmaking traditions By im-proving our understanding of black ashbiologyecology in Maine and other blackash habitats improved management strate-

gies that simultaneously target EAB controland black ash preservation which will serveto maintain this cultural keystone speciescan be developed

Case Study Four Black AshStands in Maine

In November 2014 basket-tree har-vesters and researchers visited four black ashsites in Maine as part of the Black Ash Sym-posium These sites were representative oftwo common black ash stand types found ina statewide study black ash swamps andhardwood river terraces (Gawler and Cutko2010 Costanza 2015 see also SupplementalTable S1) two sites from each stand typewere visited (Figures 3ndash6) The site visitsallowed Symposium participants to sharetheir knowledge of black ash while also as-sessing the potential impact of EAB on thesetypes of stands and on basket-quality ashWhat follows is a summary of the discus-sions and observations from the Symposiumsite visits

The four black ash stands visited are ex-amples of old-field early-successional forests(100 years old) common in central andsouthern Maine which regenerate on aban-doned agricultural land The two black ashswamps were characterized by standing wa-ter ponding and relatively flat topographywith limited pits and mounds (Figures 4 and6) The two hardwood river terraces in con-trast had little to no standing water slightdownstream slopes and small streams drain-ing the sites (Figures 3 and 5) Based on in-crement cores collected in 2012 (Costanza2015) the highest basket-quality black ashtrees were found on the two river terracesites Harvesters attending the Symposiumsupported this finding noting that they of-

ten associate high-quality black ash withriver banks and floodplains with low tomoderate stand densities

The four Maine stands had significantlylower black ash densities than documentedstands found in Minnesota Ontario andQuebec (Table 2) Stands visited rangedfrom 90ndash162 black ash stemsac1 with theswamps producing higher black ash densi-ties than the river terraces One possible ex-planation for these reduced stand densities isthat most Maine stands are early-succes-sional forests growing in previously har-vested stands or old fields Studies outside ofMaine have focused on well-established for-ests where black ash maintains high densitiesand canopy dominance It is also possiblethat black ash trees in Maine recruit differ-ently from those in other regions howeverrecruitment strategies and shade toleranceneed to be further studied to draw signifi-cant conclusions During the Symposiumsite visits basket-tree harvesters and re-searchers both noted that basketry is moreprevalent in Maine New York and south-eastern Ontario Lower stand densities anddifferent recruitment patterns along withcultural traditions may influence the pro-portion of basket-quality black ash trees inthese regions compared with others withinthe black ash range

Unlike those in many other regionsblack ash trees in Maine frequently occur inmixed species stands which can includenorthern white-cedar balsam fir Americanelm (Ulmus americana L) red maple (Acerrubrum L) and yellow birch (Betula al-leghaniensis Britton) (Gawler and Cutko2010 Costanza 2015) Pure black ash standsare uncommon unless the site is a swamp oran old-growth black ash stand (Ehrenfeld2012 Costanza 2015) Of the sites visited

Table 1 Black ash average tree age in North America based on four studies that sampled a variety of black ash stands across thenorthern United States and southern Canada

Location StudyMaximum black ash

age (4 in dbh)Average black ashage (4 in dbh) No of black ash stands

(yr)

Maine Costanza 2015 273 59 24 stands (1frasl6 ac)Chippewa National Forest Minnesota Benedict and Frelich 2008 1641311351 9698651 67 stands (164 ft radius) 2424191

Northern New York2 Costanza 2015 146 70 16 stands (1frasl6 ac)Ontario Canada Townshend 2011 1613 61 16 stands (1frasl10 ac)Lake Duparquet Queacutebec Canada Tardif and Bergeron 1999 3204 34 5 stands (composed of 54 plots 1frasl40 ac each)

Research sites ranged from pure black ash to mixed forest stands and from old-growth to anthropogenically disturbed stands1 Values correspond to three types of sites lowlandsuplandswoodland ponds Maximum and average tree ages were taken from stand data that included all species however stands were dominatedby black ash A total of 166 black ash were sampled2 In northern New York samples were collected in St Lawrence and Lewis counties3 Oldest veteran black ash tree throughout sample area was 161 years old Oldest veteran black ash tree in each stand ranged from 63ndash161 years old4 The oldest black ash was sexually reproduced The oldest black ash sprout was 120 yr old

Journal of Forestry bull September 2017 439

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

during the Symposium the black ashswamps had moderate conifer densities (pri-marily balsam fir) whereas the hardwoodriver terraces were mixed hardwood standscomposed of red maple American basswood(Tilia americana L) green ash and whiteash (Fraxinus americana L) Harvesters andresearchers both noted the improved qualityof ash on the river terraces and often associ-ated these basket-quality trees with the lackof conifers innear the stand

At the end of the Symposium severalconclusions about Mainersquos black ash popu-lation emerged Unlike those in other re-gions of North America the four blackash stands we visited were early-successional

stands characterized by low to moderatestand densities mixed species compositionand reduced black ash regeneration In ad-dition harvesters confirmed that these char-acteristics (site hydrology species composi-tion and stand structure) play a key role inthe quality of black ash trees produced Withregard to management implications thesites visited were similar to stands in Ontariowhere black ash is a secondary or minorcomponent (Jeffrey Dech Nipissing Uni-versity pers comm Feb 2 2016) andmany green ash stands in southeasternMichigan (Kashian 2016) Therefore re-search from these stands may provide valu-able insight into how similar stands in

Maine and other regions are likely to re-spond to EAB infestations

The EAB ImpactThe potential economic and ecological

impacts of the EAB invasion are staggering(Kovacs et al 2010 Aukema et al 2011) andwill drastically alter the ash components ofNorth American forests (Cappaert et al2005 Herms and McCullough 2014)Black ash in particular is highly susceptibleto the insect (Anulewicz et al 2006 Pure-swaran and Poland 2009 Siegert et al2014a) and dies within 2ndash4 years of beinginfested (Siegert et al 2005 2014b) (Figure7) In EAB-infested areas research has yet tofully address the ecological impacts to blackash Several studies report ash mortality ratesas a pooled statistic including white greenand black ash (eg Kashian and Witter2011 Klooster et al 2014) whereas otherstudies only address early infestation periods(2004ndash2008) and focus on Michigan andOhio (eg Siegert et al 2005 Smith et al2015 Kashian 2016) Researchers in Min-nesota have focused on black ash standssimulating EAB mortality by girdling blackash trees (Slesak et al 2014 Looney et al2015) However additional research isneeded to more critically evaluate black ashstand dynamics through the course of anEAB invasion and in the different habitatsand hydrologic regimes where black ash oc-curs

Based on green ash studies conductedin Michigan after EAB infestation (Kashian2016) it is possible to make a few inferencesabout the potential response of black ashstands to EAB In Michigan green ash treesare persisting with sprouts released ad-vanced regeneration and seedlings pro-duced from sprout-borne seeds on siteswhere green ash densities were relativelyhigh Conversely preliminary work suggestssprouting from EAB-killed black ash may beextremely limited (NW Siegert USDAForest Service unpubl data 2010) Re-duced densities of black ash will probablyleave stands depleted with fewer and smallerstems following the initial wave of the EABinvasion Any surviving black ash trees willprobably be small (3 in dbh) unless estab-lished EAB infestations are radically sup-pressed One study in southeastern Michi-gan conducted when EAB populations wererelatively high found that sprouts from har-vested trees were recolonized by EAB within1 year (Petrice and Haack 2011) Furtherwork by Tardif and Bergeron (1999) indi-

Figure 3 Site 1 hardwood river terrace Site 1 is a sparse to moderately dense black ashmixed hardwood stand The stand is separated from a neighboring conifer stand by a smallstream (A) The majority of black ash gt4 in dbh at this site are high quality (B)

440 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

cates a greater abundance of black ashsprouts than seedlings in young age classesin addition to higher mortality rates insprouts To fully understand how black ashsprouts may persist in EAB-infested standsfurther investigation is needed on sproutingprocesses the survivorship of black ashsprouts the relationship to EAB populationcycles and the ability of these sprouts to be-come basket-quality trees

After a few years of EAB infestationonce the overstory has been killed estab-lished black ash seedlings if present may bereleased if the resulting gaps are not quicklyfilled in by neighboring trees However be-

cause EAB is known to infest and kill trees1 in dbh (Herms and McCullough2014) it is likely that most seedlings inmixed or low-density stands will not achievea larger size (remain 3 in dbh) and themean stand diameter will remain lowWhere black ash is a dominant componentof the stand changes in evapotranspirationafter overstory loss may result in water tablerise (Slesak et al 2014) and conversion toa more open-water wetland condition(Erdmann et al 1987) further threaten-ing ash persistence in the stand In addi-tion gap formation will probably result inan increased risk of competition from re-

sidual non-ash understory trees unaffectedby EAB as well as shrubs and herbaceousvegetation (Slesak et al 2014) Based onthe rapid detrimental impact of EAB it iscritical to identify what additional infor-mation and efforts are needed to preparefor the insectrsquos arrival in many black ashstands

RecommendationsThe first step in preparing for and man-

aging EAB is early detection If and when EABis detected efforts should focus on slowing theinsectrsquos spread (Siegert et al 2015) so the avail-ability of existing basket-quality ash trees canbe extended as long as possible Managementstrategies may be implemented to complementthis effort by reducing EAB populationdensities (McCullough et al 2015) In do-ing so guidelines should be developedthat limit anthropogenic spread of EAB-infested material while allowing harvestersto safely transport basket-quality black ashmaterial without risk of transporting EABto non-infested areas For instance tworecent studies reported that submerginginfested black ash logs in water for 8 ndash16weeks killed all EAB while still maintain-ing useable basket-quality wood (Siegertet al 2014a Poland et al 2015)

If harvesters are available basket-qual-ity trees should be harvested pounded andsplit as soon as an EAB infestation is de-tected in the vicinity This harvesting in-crease would have a considerably differentimpact on forests and on black ash in par-ticular than tribal traditions historicallyhad However dried black ash splints can bestored for several years (Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) increas-ing harvesting would allow harvesters to usewood that would otherwise be lost to EABLightly infested black ash trees remain suit-able for basketmaking materials but degraderapidly as EAB density increases reducingyields from a given log to the point where itmay be unusable for splints (Siegert et al2014a Poland et al 2015) In additionsome have recommended increasing blackash harvesting now before EAB arrives Thiswould potentially reduce concerns aboutquarantines accidental insect transport andloss of basket-quality wood after an EAB in-festation

Last we recommend collaboration be-tween state federal and tribal agencies todevelop memorandums of understanding orjoint agreements before the arrival of EAB

Figure 4 Site 2 black ash swamp Site 2 is a moderately dense mixed hardwood-coniferstand with black ash red maple and balsam fir The site is characterized by minimalponding (A) root buttressing (A) and small black ash trees typically lt6 in dbh (B) Themajority of black ash gt4 in dbh are poor quality

Journal of Forestry bull September 2017 441

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

in high-value black ash stands Such agree-ments can alleviate confusion and promoteaction as soon as EAB is detected in or neartribal lands or other traditional black ashharvesting sites Furthermore such agree-ments can help foster discussions on accessi-bility to black ash trees To preserve theblack ash resource and maintain black ashbasketry in the face of EAB discussions re-garding accessibility and response plansneed to occur

Future ResearchIn addition to making preparations

for EABrsquos arrival in black ash stands there

are countless opportunities to learn moreabout this still-overlooked tree species Toimprove our understanding of black ashtreesrsquo potential response to EAB and im-pacts to cultural traditions future researchdirections were discussed during the BlackAsh Symposium What follows is a summaryof the three most pressing research needsinfluence of black ash stand dynamics andsite hydrology on regeneration and qualityof black ash role of black ash genetics in thefuture of the population and EAB popula-tion dynamics in infested black ash standsand the effect on black ash regeneration

Influence of Stand Dynamics and Hy-drology on Regeneration and Quality

Very little information exists on blackash stand histories conditions and naturaldynamics in Maine For example it is un-clear whether Maine black ash stands un-dergo continuous recruitment and remainstocked over an extended period of time or ifblack ash is replaced by other species Natu-ral stand dynamics and succession are poorlyunderstood partially because there are fewold-growth stands in Maine Locating andstudying a few old-growth black ash stands(200 years old) would greatly improve ourcurrent understanding of succession Theinclusion of managed stands would alsoprove beneficial allowing researchers toquantify the response of black ash to silvicul-tural treatments Furthermore describingthe dynamics of ash across a range of distur-bance histories would allow researchers todevelop a much needed successional frame-work for black ash and to develop a descrip-tion of the speciesrsquo regeneration niche In themidwestern United States recent studieshave suggested that the future of black ashstands faced with EAB and climate changewill probably center around non-ash re-placement species (Looney et al 2015Iverson et al 2016) However in regionswhere black ash is a cultural keystone spe-cies replacing black ash is not an optimalmanagement strategy Describing black ashstand dynamics and recruitment in Maineas well as in other black ash habitats such asthose in Nova Scotia southern and centralOntario and Quebec would provide in-sight on how to maintain the species bothfor basketmaking and ecological reasons

Stand dynamics are heavily influencedby stand hydrology which is poorly under-stood in Maine and other parts of the blackash range A few recent studies in Minnesotahave explored black ash stand hydrology(Slesak et al 2014 Telander et al 2015)however the extent to which water tablefluctuations in black ash stands influence re-generation and recruitment across the entirerange should be further explored For exam-ple in situations where water table dynamicsare driven primarily by topography or localsurface water a change in evapotranspira-tion after overstory loss may not result in aconcomitant change in site hydrology (Rob-ert Slesak Minnesota Forest ResourcesCouncil pers comm June 5 2015) Fur-thermore the role of post-disturbancesprouting of black ash in regulating watertable dynamics is also unclear as is the role

Figure 5 Site 3 hardwood river terrace Site 3 is a moderately dense mixed hardwoodstand with occasional conifer (A) A small stream runs northeast to southwest and anunpaved road runs west to east through the stand In better-drained areas large high-quality black ash are produced Where ponding occurs black ash tend to be smaller andpoorer quality The majority of black ash gt4 in dbh are high quality (B)

442 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

during the Symposium the black ashswamps had moderate conifer densities (pri-marily balsam fir) whereas the hardwoodriver terraces were mixed hardwood standscomposed of red maple American basswood(Tilia americana L) green ash and whiteash (Fraxinus americana L) Harvesters andresearchers both noted the improved qualityof ash on the river terraces and often associ-ated these basket-quality trees with the lackof conifers innear the stand

At the end of the Symposium severalconclusions about Mainersquos black ash popu-lation emerged Unlike those in other re-gions of North America the four blackash stands we visited were early-successional

stands characterized by low to moderatestand densities mixed species compositionand reduced black ash regeneration In ad-dition harvesters confirmed that these char-acteristics (site hydrology species composi-tion and stand structure) play a key role inthe quality of black ash trees produced Withregard to management implications thesites visited were similar to stands in Ontariowhere black ash is a secondary or minorcomponent (Jeffrey Dech Nipissing Uni-versity pers comm Feb 2 2016) andmany green ash stands in southeasternMichigan (Kashian 2016) Therefore re-search from these stands may provide valu-able insight into how similar stands in

Maine and other regions are likely to re-spond to EAB infestations

The EAB ImpactThe potential economic and ecological

impacts of the EAB invasion are staggering(Kovacs et al 2010 Aukema et al 2011) andwill drastically alter the ash components ofNorth American forests (Cappaert et al2005 Herms and McCullough 2014)Black ash in particular is highly susceptibleto the insect (Anulewicz et al 2006 Pure-swaran and Poland 2009 Siegert et al2014a) and dies within 2ndash4 years of beinginfested (Siegert et al 2005 2014b) (Figure7) In EAB-infested areas research has yet tofully address the ecological impacts to blackash Several studies report ash mortality ratesas a pooled statistic including white greenand black ash (eg Kashian and Witter2011 Klooster et al 2014) whereas otherstudies only address early infestation periods(2004ndash2008) and focus on Michigan andOhio (eg Siegert et al 2005 Smith et al2015 Kashian 2016) Researchers in Min-nesota have focused on black ash standssimulating EAB mortality by girdling blackash trees (Slesak et al 2014 Looney et al2015) However additional research isneeded to more critically evaluate black ashstand dynamics through the course of anEAB invasion and in the different habitatsand hydrologic regimes where black ash oc-curs

Based on green ash studies conductedin Michigan after EAB infestation (Kashian2016) it is possible to make a few inferencesabout the potential response of black ashstands to EAB In Michigan green ash treesare persisting with sprouts released ad-vanced regeneration and seedlings pro-duced from sprout-borne seeds on siteswhere green ash densities were relativelyhigh Conversely preliminary work suggestssprouting from EAB-killed black ash may beextremely limited (NW Siegert USDAForest Service unpubl data 2010) Re-duced densities of black ash will probablyleave stands depleted with fewer and smallerstems following the initial wave of the EABinvasion Any surviving black ash trees willprobably be small (3 in dbh) unless estab-lished EAB infestations are radically sup-pressed One study in southeastern Michi-gan conducted when EAB populations wererelatively high found that sprouts from har-vested trees were recolonized by EAB within1 year (Petrice and Haack 2011) Furtherwork by Tardif and Bergeron (1999) indi-

Figure 3 Site 1 hardwood river terrace Site 1 is a sparse to moderately dense black ashmixed hardwood stand The stand is separated from a neighboring conifer stand by a smallstream (A) The majority of black ash gt4 in dbh at this site are high quality (B)

440 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

cates a greater abundance of black ashsprouts than seedlings in young age classesin addition to higher mortality rates insprouts To fully understand how black ashsprouts may persist in EAB-infested standsfurther investigation is needed on sproutingprocesses the survivorship of black ashsprouts the relationship to EAB populationcycles and the ability of these sprouts to be-come basket-quality trees

After a few years of EAB infestationonce the overstory has been killed estab-lished black ash seedlings if present may bereleased if the resulting gaps are not quicklyfilled in by neighboring trees However be-

cause EAB is known to infest and kill trees1 in dbh (Herms and McCullough2014) it is likely that most seedlings inmixed or low-density stands will not achievea larger size (remain 3 in dbh) and themean stand diameter will remain lowWhere black ash is a dominant componentof the stand changes in evapotranspirationafter overstory loss may result in water tablerise (Slesak et al 2014) and conversion toa more open-water wetland condition(Erdmann et al 1987) further threaten-ing ash persistence in the stand In addi-tion gap formation will probably result inan increased risk of competition from re-

sidual non-ash understory trees unaffectedby EAB as well as shrubs and herbaceousvegetation (Slesak et al 2014) Based onthe rapid detrimental impact of EAB it iscritical to identify what additional infor-mation and efforts are needed to preparefor the insectrsquos arrival in many black ashstands

RecommendationsThe first step in preparing for and man-

aging EAB is early detection If and when EABis detected efforts should focus on slowing theinsectrsquos spread (Siegert et al 2015) so the avail-ability of existing basket-quality ash trees canbe extended as long as possible Managementstrategies may be implemented to complementthis effort by reducing EAB populationdensities (McCullough et al 2015) In do-ing so guidelines should be developedthat limit anthropogenic spread of EAB-infested material while allowing harvestersto safely transport basket-quality black ashmaterial without risk of transporting EABto non-infested areas For instance tworecent studies reported that submerginginfested black ash logs in water for 8 ndash16weeks killed all EAB while still maintain-ing useable basket-quality wood (Siegertet al 2014a Poland et al 2015)

If harvesters are available basket-qual-ity trees should be harvested pounded andsplit as soon as an EAB infestation is de-tected in the vicinity This harvesting in-crease would have a considerably differentimpact on forests and on black ash in par-ticular than tribal traditions historicallyhad However dried black ash splints can bestored for several years (Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) increas-ing harvesting would allow harvesters to usewood that would otherwise be lost to EABLightly infested black ash trees remain suit-able for basketmaking materials but degraderapidly as EAB density increases reducingyields from a given log to the point where itmay be unusable for splints (Siegert et al2014a Poland et al 2015) In additionsome have recommended increasing blackash harvesting now before EAB arrives Thiswould potentially reduce concerns aboutquarantines accidental insect transport andloss of basket-quality wood after an EAB in-festation

Last we recommend collaboration be-tween state federal and tribal agencies todevelop memorandums of understanding orjoint agreements before the arrival of EAB

Figure 4 Site 2 black ash swamp Site 2 is a moderately dense mixed hardwood-coniferstand with black ash red maple and balsam fir The site is characterized by minimalponding (A) root buttressing (A) and small black ash trees typically lt6 in dbh (B) Themajority of black ash gt4 in dbh are poor quality

Journal of Forestry bull September 2017 441

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

in high-value black ash stands Such agree-ments can alleviate confusion and promoteaction as soon as EAB is detected in or neartribal lands or other traditional black ashharvesting sites Furthermore such agree-ments can help foster discussions on accessi-bility to black ash trees To preserve theblack ash resource and maintain black ashbasketry in the face of EAB discussions re-garding accessibility and response plansneed to occur

Future ResearchIn addition to making preparations

for EABrsquos arrival in black ash stands there

are countless opportunities to learn moreabout this still-overlooked tree species Toimprove our understanding of black ashtreesrsquo potential response to EAB and im-pacts to cultural traditions future researchdirections were discussed during the BlackAsh Symposium What follows is a summaryof the three most pressing research needsinfluence of black ash stand dynamics andsite hydrology on regeneration and qualityof black ash role of black ash genetics in thefuture of the population and EAB popula-tion dynamics in infested black ash standsand the effect on black ash regeneration

Influence of Stand Dynamics and Hy-drology on Regeneration and Quality

Very little information exists on blackash stand histories conditions and naturaldynamics in Maine For example it is un-clear whether Maine black ash stands un-dergo continuous recruitment and remainstocked over an extended period of time or ifblack ash is replaced by other species Natu-ral stand dynamics and succession are poorlyunderstood partially because there are fewold-growth stands in Maine Locating andstudying a few old-growth black ash stands(200 years old) would greatly improve ourcurrent understanding of succession Theinclusion of managed stands would alsoprove beneficial allowing researchers toquantify the response of black ash to silvicul-tural treatments Furthermore describingthe dynamics of ash across a range of distur-bance histories would allow researchers todevelop a much needed successional frame-work for black ash and to develop a descrip-tion of the speciesrsquo regeneration niche In themidwestern United States recent studieshave suggested that the future of black ashstands faced with EAB and climate changewill probably center around non-ash re-placement species (Looney et al 2015Iverson et al 2016) However in regionswhere black ash is a cultural keystone spe-cies replacing black ash is not an optimalmanagement strategy Describing black ashstand dynamics and recruitment in Maineas well as in other black ash habitats such asthose in Nova Scotia southern and centralOntario and Quebec would provide in-sight on how to maintain the species bothfor basketmaking and ecological reasons

Stand dynamics are heavily influencedby stand hydrology which is poorly under-stood in Maine and other parts of the blackash range A few recent studies in Minnesotahave explored black ash stand hydrology(Slesak et al 2014 Telander et al 2015)however the extent to which water tablefluctuations in black ash stands influence re-generation and recruitment across the entirerange should be further explored For exam-ple in situations where water table dynamicsare driven primarily by topography or localsurface water a change in evapotranspira-tion after overstory loss may not result in aconcomitant change in site hydrology (Rob-ert Slesak Minnesota Forest ResourcesCouncil pers comm June 5 2015) Fur-thermore the role of post-disturbancesprouting of black ash in regulating watertable dynamics is also unclear as is the role

Figure 5 Site 3 hardwood river terrace Site 3 is a moderately dense mixed hardwoodstand with occasional conifer (A) A small stream runs northeast to southwest and anunpaved road runs west to east through the stand In better-drained areas large high-quality black ash are produced Where ponding occurs black ash tend to be smaller andpoorer quality The majority of black ash gt4 in dbh are high quality (B)

442 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

cates a greater abundance of black ashsprouts than seedlings in young age classesin addition to higher mortality rates insprouts To fully understand how black ashsprouts may persist in EAB-infested standsfurther investigation is needed on sproutingprocesses the survivorship of black ashsprouts the relationship to EAB populationcycles and the ability of these sprouts to be-come basket-quality trees

After a few years of EAB infestationonce the overstory has been killed estab-lished black ash seedlings if present may bereleased if the resulting gaps are not quicklyfilled in by neighboring trees However be-

cause EAB is known to infest and kill trees1 in dbh (Herms and McCullough2014) it is likely that most seedlings inmixed or low-density stands will not achievea larger size (remain 3 in dbh) and themean stand diameter will remain lowWhere black ash is a dominant componentof the stand changes in evapotranspirationafter overstory loss may result in water tablerise (Slesak et al 2014) and conversion toa more open-water wetland condition(Erdmann et al 1987) further threaten-ing ash persistence in the stand In addi-tion gap formation will probably result inan increased risk of competition from re-

sidual non-ash understory trees unaffectedby EAB as well as shrubs and herbaceousvegetation (Slesak et al 2014) Based onthe rapid detrimental impact of EAB it iscritical to identify what additional infor-mation and efforts are needed to preparefor the insectrsquos arrival in many black ashstands

RecommendationsThe first step in preparing for and man-

aging EAB is early detection If and when EABis detected efforts should focus on slowing theinsectrsquos spread (Siegert et al 2015) so the avail-ability of existing basket-quality ash trees canbe extended as long as possible Managementstrategies may be implemented to complementthis effort by reducing EAB populationdensities (McCullough et al 2015) In do-ing so guidelines should be developedthat limit anthropogenic spread of EAB-infested material while allowing harvestersto safely transport basket-quality black ashmaterial without risk of transporting EABto non-infested areas For instance tworecent studies reported that submerginginfested black ash logs in water for 8 ndash16weeks killed all EAB while still maintain-ing useable basket-quality wood (Siegertet al 2014a Poland et al 2015)

If harvesters are available basket-qual-ity trees should be harvested pounded andsplit as soon as an EAB infestation is de-tected in the vicinity This harvesting in-crease would have a considerably differentimpact on forests and on black ash in par-ticular than tribal traditions historicallyhad However dried black ash splints can bestored for several years (Basket-tree harvest-ers Maine Indian Basketmakers Alliancepers comm MayndashAugust 2013) increas-ing harvesting would allow harvesters to usewood that would otherwise be lost to EABLightly infested black ash trees remain suit-able for basketmaking materials but degraderapidly as EAB density increases reducingyields from a given log to the point where itmay be unusable for splints (Siegert et al2014a Poland et al 2015) In additionsome have recommended increasing blackash harvesting now before EAB arrives Thiswould potentially reduce concerns aboutquarantines accidental insect transport andloss of basket-quality wood after an EAB in-festation

Last we recommend collaboration be-tween state federal and tribal agencies todevelop memorandums of understanding orjoint agreements before the arrival of EAB

Figure 4 Site 2 black ash swamp Site 2 is a moderately dense mixed hardwood-coniferstand with black ash red maple and balsam fir The site is characterized by minimalponding (A) root buttressing (A) and small black ash trees typically lt6 in dbh (B) Themajority of black ash gt4 in dbh are poor quality

Journal of Forestry bull September 2017 441

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

in high-value black ash stands Such agree-ments can alleviate confusion and promoteaction as soon as EAB is detected in or neartribal lands or other traditional black ashharvesting sites Furthermore such agree-ments can help foster discussions on accessi-bility to black ash trees To preserve theblack ash resource and maintain black ashbasketry in the face of EAB discussions re-garding accessibility and response plansneed to occur

Future ResearchIn addition to making preparations

for EABrsquos arrival in black ash stands there

are countless opportunities to learn moreabout this still-overlooked tree species Toimprove our understanding of black ashtreesrsquo potential response to EAB and im-pacts to cultural traditions future researchdirections were discussed during the BlackAsh Symposium What follows is a summaryof the three most pressing research needsinfluence of black ash stand dynamics andsite hydrology on regeneration and qualityof black ash role of black ash genetics in thefuture of the population and EAB popula-tion dynamics in infested black ash standsand the effect on black ash regeneration

Influence of Stand Dynamics and Hy-drology on Regeneration and Quality

Very little information exists on blackash stand histories conditions and naturaldynamics in Maine For example it is un-clear whether Maine black ash stands un-dergo continuous recruitment and remainstocked over an extended period of time or ifblack ash is replaced by other species Natu-ral stand dynamics and succession are poorlyunderstood partially because there are fewold-growth stands in Maine Locating andstudying a few old-growth black ash stands(200 years old) would greatly improve ourcurrent understanding of succession Theinclusion of managed stands would alsoprove beneficial allowing researchers toquantify the response of black ash to silvicul-tural treatments Furthermore describingthe dynamics of ash across a range of distur-bance histories would allow researchers todevelop a much needed successional frame-work for black ash and to develop a descrip-tion of the speciesrsquo regeneration niche In themidwestern United States recent studieshave suggested that the future of black ashstands faced with EAB and climate changewill probably center around non-ash re-placement species (Looney et al 2015Iverson et al 2016) However in regionswhere black ash is a cultural keystone spe-cies replacing black ash is not an optimalmanagement strategy Describing black ashstand dynamics and recruitment in Maineas well as in other black ash habitats such asthose in Nova Scotia southern and centralOntario and Quebec would provide in-sight on how to maintain the species bothfor basketmaking and ecological reasons

Stand dynamics are heavily influencedby stand hydrology which is poorly under-stood in Maine and other parts of the blackash range A few recent studies in Minnesotahave explored black ash stand hydrology(Slesak et al 2014 Telander et al 2015)however the extent to which water tablefluctuations in black ash stands influence re-generation and recruitment across the entirerange should be further explored For exam-ple in situations where water table dynamicsare driven primarily by topography or localsurface water a change in evapotranspira-tion after overstory loss may not result in aconcomitant change in site hydrology (Rob-ert Slesak Minnesota Forest ResourcesCouncil pers comm June 5 2015) Fur-thermore the role of post-disturbancesprouting of black ash in regulating watertable dynamics is also unclear as is the role

Figure 5 Site 3 hardwood river terrace Site 3 is a moderately dense mixed hardwoodstand with occasional conifer (A) A small stream runs northeast to southwest and anunpaved road runs west to east through the stand In better-drained areas large high-quality black ash are produced Where ponding occurs black ash tend to be smaller andpoorer quality The majority of black ash gt4 in dbh are high quality (B)

442 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

in high-value black ash stands Such agree-ments can alleviate confusion and promoteaction as soon as EAB is detected in or neartribal lands or other traditional black ashharvesting sites Furthermore such agree-ments can help foster discussions on accessi-bility to black ash trees To preserve theblack ash resource and maintain black ashbasketry in the face of EAB discussions re-garding accessibility and response plansneed to occur

Future ResearchIn addition to making preparations

for EABrsquos arrival in black ash stands there

are countless opportunities to learn moreabout this still-overlooked tree species Toimprove our understanding of black ashtreesrsquo potential response to EAB and im-pacts to cultural traditions future researchdirections were discussed during the BlackAsh Symposium What follows is a summaryof the three most pressing research needsinfluence of black ash stand dynamics andsite hydrology on regeneration and qualityof black ash role of black ash genetics in thefuture of the population and EAB popula-tion dynamics in infested black ash standsand the effect on black ash regeneration

Influence of Stand Dynamics and Hy-drology on Regeneration and Quality

Very little information exists on blackash stand histories conditions and naturaldynamics in Maine For example it is un-clear whether Maine black ash stands un-dergo continuous recruitment and remainstocked over an extended period of time or ifblack ash is replaced by other species Natu-ral stand dynamics and succession are poorlyunderstood partially because there are fewold-growth stands in Maine Locating andstudying a few old-growth black ash stands(200 years old) would greatly improve ourcurrent understanding of succession Theinclusion of managed stands would alsoprove beneficial allowing researchers toquantify the response of black ash to silvicul-tural treatments Furthermore describingthe dynamics of ash across a range of distur-bance histories would allow researchers todevelop a much needed successional frame-work for black ash and to develop a descrip-tion of the speciesrsquo regeneration niche In themidwestern United States recent studieshave suggested that the future of black ashstands faced with EAB and climate changewill probably center around non-ash re-placement species (Looney et al 2015Iverson et al 2016) However in regionswhere black ash is a cultural keystone spe-cies replacing black ash is not an optimalmanagement strategy Describing black ashstand dynamics and recruitment in Maineas well as in other black ash habitats such asthose in Nova Scotia southern and centralOntario and Quebec would provide in-sight on how to maintain the species bothfor basketmaking and ecological reasons

Stand dynamics are heavily influencedby stand hydrology which is poorly under-stood in Maine and other parts of the blackash range A few recent studies in Minnesotahave explored black ash stand hydrology(Slesak et al 2014 Telander et al 2015)however the extent to which water tablefluctuations in black ash stands influence re-generation and recruitment across the entirerange should be further explored For exam-ple in situations where water table dynamicsare driven primarily by topography or localsurface water a change in evapotranspira-tion after overstory loss may not result in aconcomitant change in site hydrology (Rob-ert Slesak Minnesota Forest ResourcesCouncil pers comm June 5 2015) Fur-thermore the role of post-disturbancesprouting of black ash in regulating watertable dynamics is also unclear as is the role

Figure 5 Site 3 hardwood river terrace Site 3 is a moderately dense mixed hardwoodstand with occasional conifer (A) A small stream runs northeast to southwest and anunpaved road runs west to east through the stand In better-drained areas large high-quality black ash are produced Where ponding occurs black ash tend to be smaller andpoorer quality The majority of black ash gt4 in dbh are high quality (B)

442 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

of increased deadwood inputs from EAB-induced mortality on hydrology and standdevelopment Site hydrology is also a criticalfactor in determining the quality of blackash trees yet the impact of EAB-inducedchanges in hydrology on basket-quality ashhave yet to be determined To address theseknowledge gaps the establishment of long-term experimental and monitoring sitescould help track succession and develop-ment assess changes in the water tablequantify changes in basket-quality ash andmonitor for the arrival of EAB

Role of Black Ash Genetics and Futureof Species

When speaking with basket-tree har-vesters several types of black ash includ-ing ldquoridgerdquo ldquoriverrdquo ldquoyellowrdquo ldquopinkrdquo andldquoswamprdquo ash were mentioned Accordingto harvesters each of these types producesa different quality basket splint This leadsresearchers to ask Is there any genetic dis-tinction between these types If so canspecific trees with better-quality wood betargeted for seed collection and manage-ment To address these questions genetic

testing of black ash from various standtypes is recommended

Tree characteristics such as vessel size(both earlywood and latewood) and barkdepth play a role in black ash wood qualitybut that role is poorly understood Kameset al (2016) noted large annual variability inblack ash latewood width as well as variabil-ity in earlywood vessel lumen but did notdirectly look at wood quality associated withthis variability A few other studies observeda marked decrease in latewood width on sat-urated sites which resulted in small growthrings unsuitable for basketry (Costanza2015 Jacques Tardif Centre for Forest In-terdisciplinary Research pers comm Feb8 2016) However these reports were basedon visual observations To fully understandhow site characteristics impact seasonalgrowth (earlywoodlatewood) and vessel sizeand in turn wood quality for basketry wepropose a study comparing the ratio of ear-lywood to latewood using previously col-lected black ash cores from a range of sites(eg Tardif and Bergeron 1999 Townsh-end 2011 Costanza 2015 Telander et al2015)

Effects of EAB Infestation on Black AshPopulation Dynamics

Much remains to be learned about thelong-term impacts of the EAB invasion inblack ash stands and how it may affect sur-vival recruitment regeneration and qualityof black ash Establishing permanent plotsin uninfested stands to monitor the responseof black ash stands after EAB infestation inMaine and elsewhere would generate valu-able information that is critically neededTimelines could be developed for salvagingEAB-infested black ash that can still be usedfor basketry Permanent plots would also al-low researchers to measure seedling andsprout response for multiple years post-infestation These stands could be assessedby basket-tree harvesters for abundance andquality of black ash regeneration to deter-mine the presence and frequency of basket-quality wood

Black ash vegetative reproduction (iesprouting) after EAB infestation has notbeen evaluated In black ash stands fre-quently harvested by basketmakers harvest-ers have reported regular occurrences ofstump sprouting Vegetative reproductionthrough stump sprouting may contribute tomaintaining black ash populations in Maineand other regions where basketry is com-mon at least where the relative density of

Figure 6 Site 4 black ash swamp Site 4 is a moderate to highly dense black ash stand Theedge of the stand grades into a lake The site is characterized by mucky saturated soils (A)nearly year-round Most black ash are small with very few trees gt6 in dbh Treesfrequently have sweeping stems epicormic branching and root buttressing (B) Black ashgt4 in dbh are poor quality with very narrow annual growth (B)

Journal of Forestry bull September 2017 443

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

black ash is moderate or high Black ash de-cline and mortality caused by EAB how-ever may deplete root starch reserves result-ing in reduced frequency vigor orsurvivorship of vegetative growth Further-more little is known about the quality ofthese sprouts regenerated from the stumps ofbasket-harvested trees More work is neededin EAB-infested black ash stands particu-

larly basket-quality stands to determine thedegree to which stump sprouting may influ-ence stand dynamics and whether basket-quality trees may be produced before be-coming infested by EAB again If thesesprouts can produce basket-quality treesimplementing a coppicing rotation may bean effective management strategy to main-tain a basket-quality black ash populationSuch studies would improve our under-standing of black ash vegetative reproduc-tion while simultaneously exploring whereand how black ash basketry may persist Byknowing where basket-quality ash is beingproduced and which stands can continue toregenerate basket-quality trees post-EAB re-source managers and tribal members canwork to sustain this cultural keystone speciesand associated cultural traditions for years tocome

ConclusionsThroughout this work and via the

Black Ash Symposium harvesters and scien-tists came together to synthesize currentblack ash knowledge in turn developing arecommended action plan for Mainersquos blackash resource Our results explicitly demon-strate how traditional ecological knowledgeand scientific knowledge can support eachother and also how they can be applied toforest health problems to develop collabora-tive solutions The recommendations sug-gested here while focused on Maine can beapplied in locations with similar black ashhabitat Our findings indicate that Mainersquosblack ash population is typically less denseand produces a relatively large number ofbasket-quality trees compared with other re-gions representing a unique situation Tomaintain this cultural keystone species inMaine and other regions further research on

the speciesrsquo response to EAB infestations isneeded along with a better understanding ofthe speciesrsquo ecology and recruitment pat-terns This information not only will assistthose studying and managing black ash pop-ulations but also will benefit basket-tree har-vesters tribal members and foresters inmaintaining basketry and cultural tradi-tions The recommendations and current re-search needs described here can help informblack ash management at community stateand regional levels and are representative ofthe effective collaboration between tribalmembers basket-tree harvesters and scien-tists

Endnote1 For more information see the Emerald

Ash Borer Information Network at wwwemeraldashborerinfo

Literature CitedANULEWICZ AC DG MCCULLOUGH AND

DL MILLER 2006 Oviposition and develop-ment of emerald ash borer (Agrilus planipennis)(Coleoptera Buprestidae) on hosts and poten-tial hosts in no-choice bioassays Great LakesEntomol 39(3ndash4)99ndash112 Available online athttpmichentsocorggle-pdfsvol39no3-4pdf

AUKEMA JE B LEUNG K KOVACS C CHIVERSKO BRITTON J ENGLIN SJ FRANKEL ET AL2011 Economic impacts of non-native forestinsects in the continental United States PLoSOne 6(9)e24587 doi101371journalpone0024587

BARNES BV AND WH WAGNER 2004 Mich-igan trees A guide to the trees of the Great Lakesregion University of Michigan Press Ann Ar-bor MI 447 p

BENEDICT L AND R DAVID 2004 Handbook ofblack ash Preservation reforestation regenera-tion Department of the Environment Mo-hawk Council of Akwesasne HogansburgNY 79 p

BENEDICT MA AND LE FRELICH 2008 Sitefactors affecting black ash ring growth innorthern Minnesota For Ecol Manage255(8ndash9)3489ndash3493 doi101016jforeco200802029

CAPPAERT D DG MCCULLOUGH TM PO-LAND AND NW SIEGERT 2005 Emerald ashborer in North America A research and regu-latory challenge Am Entomol 51(3)152ndash165 doi101093ae513152

COSTANZA KKL 2015 Growth response ofFraxinus nigra Marsh (Lamiales Oleaceae)used to predict high-quality sites in Maine andnorthern New York An approach to prioritizingpreparedness and management of Agrilus plani-pennis Fairmaire (Coleoptera Buprestidae)MSc thesis Univ of Maine Orono ME120 p

DIAMOND AK AND MR EMERY 2011 Blackash (Fraxinus nigra Marsh) Local ecologicalknowledge of site characteristics and morphol-

Figure 7 EAB infestation in a black ash treeEAB can easily be identified by the distinc-tive S-shaped galleries from larvae feedingon xylem tissue This black ash tree is in-fested with EAB with clear galleries under-neath the bark

Table 2 Black ash stand densities in North America based on five studies conducted inthe northern United States and southern Canada that focused on stands with asignificant component of black ash

Location StudyStand density (stemsac

trees 4 in dbh) No of stands

Northeast Minnesota Palik et al 2011 228ndash2621 54Chippewa National Forest Minnesota Slesak et al 2014 228ndash3712 67Ontario JP Dech unpubl data 152ndash6173 16Lake Duparquet Northwest Quebec Tardif and Bergeron 1999 2049ndash137404 5New York Costanza 2015 24ndash1625 16Maine Costanza 2015 54ndash1865 24

1 Measured stand densities of black ash stands primarily composed of black ash2 Black ash comprised 75 to 100 of stand basal area with pure stands commonly occurring3 Measured stand densities where black ash ranged from 39 to 100 of stand basal area4 Measured all black ash 4 in tall5 Density only includes black ash trees

444 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

ogy associated with basket-grade specimens inNew England (USA) Econ Bot 65(4)422ndash426 doi101007s12231-011-9174-z

EHRENFELD J 2012 Northern red maple andblack ash swamps P 149ndash160 in Wetlandhabitats of North America Ecology and conser-vation concerns Batzer DP and AH Bald-win (eds) Univ of California Press OaklandCA 408 p

ERDMANN GG TR CROW RM PETERSONJR AND CD WILSON 1987 Managing blackash in the Lake States USDA Forest ServiceGen Tech Rep NC-115 North Central For-est Experiment Station St Paul MN 12 phttpswwwtreesearchfsfeduspubs10179

FISICHELLI NA LE FRELICH AND PB REICH2014 Temperate tree expansion into adjacentboreal forest patches facilitated by warmertemperatures Ecography 37(2)152ndash161 doi101111j1600-0587201300197x

GAWLER SC AND A CUTKO 2010 Naturallandscapes of Maine Maine Natural Areas Pro-gram Augusta ME 347 p

GUSTAFSON EJ AND BR STURTEVANT 2013Modeling forest mortality caused by droughtstress Implications for climate change Ecosys-tems 16(1)60ndash74 doi101007s10021-012-9596-1

HERMS DA AND DG MCCULLOUGH 2014Emerald ash borer invasion of North AmericaHistory biology ecology impacts and man-agement Annu Rev Entomol 5913ndash30 doi101146annurev-ento-011613-162051

IVERSON L KS KNIGHT A PRASAD DAHERMS S MATTHEWS M PETERS A SMITHDM HARTZLER R LONG AND J ALMEND-INGER 2016 Potential species replacementsfor black ash (Fraxinus nigra) at the confluenceof two threats Emerald ash borer and a chang-ing climate Ecosystems 19(2)248ndash270 doi101007s10021-015-9929-y

KAMES S JC TARDIF AND Y BERGERON 2016Continuous earlywood vessels chronologies infloodplain ring-porous species can improvedendrohydrological reconstructions of springhigh flows and flood levels J Hydrol 534377ndash389 doi101016jjhydrol201601002

KASHIAN DM 2016 Sprouting and seed pro-duction may promote persistence of green ashin the presence of the emerald ash borer Eco-sphere 7(4)e01332 doi101002ecs21332

KASHIAN DM AND JA WITTER 2011 Assess-ing the potential for ash canopy tree replace-ment via current regeneration following emer-ald ash borer-caused mortality on southeasternMichigan landscapes For Ecol Manage261(3)480ndash488 doi101016jforeco201010033

KLOOSTER WS DA HERMS KS KNIGHTCP HERMS DG MCCULLOUGH A SMITHKJK GANDHI AND J CARDINA 2014 Ash(Fraxinus spp) mortality regeneration andseed bank dynamics in mixed hardwood for-ests following invasion by emerald ash borer(Agrilus planipennis) Biol Invas 16(4)859ndash873 doi101007s10530-013-0543-7

KOVACS KF RG HAIGHT DG MC-CULLOUGH RJ MERCADER NW SIEGERT

AND AM LIEBHOLD 2010 Cost of potentialemerald ash borer damage in US communi-ties 2009ndash2019 Ecol Econ 69(3)569ndash578doi101016jecolecon200909004

LEE R 2004 COSEWIC status report on floodedjellyskin (Leptogium rivulare) in Canada Com-mittee on the Status of Endangered Wildlife inCanada (COSEWIC) Gatineau QC Can-ada 30 p

LELAND CG 1884 The Algonquin legends ofNew England Riverside Press CambridgeMA 379 p

LENHART C K BROOKS M DAVIDSON R SLE-SAK AND A DrsquoAMATO 2012 Hydrologicsource characterization of black ash wetlandsImplications for EAB response In Proc ofConference on riparian ecosystems IV Advancingscience economics and policy Lanfear K (ed)American Water Resource Association Middle-burg VA 167 p

LOONEY CE AW DrsquoAMATO S FRAVER BJPALIK AND MR REINIKAINEN 2016 Examin-ing the influences of tree-to-tree competitionand climate on size-growth relationships in hy-dric multi-aged Fraxinus nigra stands ForEcol Manage 375238ndash248 doi101016jforeco201605050

LOONEY CE AW DrsquoAMATO BJ PALIK AND

RA SLESAK 2015 Overstory treatment andplanting season affect survival of replacementtree species in emerald ash borer threatenedFraxinus nigra forests in Minnesota USACan J For Res 45(12)1728ndash1738 doi101139cjfr-2015-0129

MCCULLOUGH DG RJ MERCADER AND

NW SIEGERT 2015 Developing and inte-grating tactics to slow ash (Oleaceae) mortalitycaused by emerald ash borer (Coleoptera Bu-prestidae) Can Entomol 147(3)349ndash358doi104039tce20153

MENDELSSOHN IA DP BATZER CR HOLTAND SA GRAHAM 2014 Abiotic constraintsfor wetland plants and animals P 61ndash85 inEcology of freshwater and estuarine wetlandsBatzer DP and RR Sharitz (eds) Univer-sity of California Press Oakland CA 372 p

PALIK BJ ME OSTRY RC VENETTE AND EABDELA 2011 Fraxinus nigra (black ash) die-back in Minnesota Regional variation and po-tential contributing factors For Ecol Manage261(1)128ndash135 doi101016jforeco201009041

PETRICE TR AND RA HAACK 2011 Effects ofcutting time stump height and herbicide ap-plication on ash (Fraxinus spp) stump sprout-ing and colonization by emerald ash borer(Agrilus planipennis) North J Appl For28(2)79 ndash 83 httpwwwingentaconnectcomcontentonesafnjaf20110000002800000002art00004

POLAND TM TM CIARAMITARO MR EM-ERY DJ CROOK E PIGEON AND A PIGEON2015 Submergence of black ash logs to con-trol emerald ash borer and preserve wood forAmerican Indian basketmaking Agric ForEntomol 17(4)412ndash 420 doi101111afe12122

PURESWARAN DS AND TM POLAND 2009Host selection and feeding preference of Agri-

lus planipennis (Coleoptera Buprestidae) onash (Fraxinus spp) Environ Entomol 38(3)757ndash765 doi1016030220380328

REYES GP D KNEESHAW AND L GRANDPRE2013 The relative importance of natural dis-turbances and local site factors on woody veg-etation regeneration diversity across a largecontiguous forest region Open J For 3(3)88ndash98 doi104236ojf201333015

ROSS RJ 2010 Wood handbook Wood as an en-gineering material USDA Forest Service GenTech Rep FPL-GTR-190 Forest ProductsLaboratory Madison WI 508 p httpswwwtreesearchfsfeduspubs37440

SENA GOMES AR AND TT KOZLOWSKI 1980Growth responses and adaptations of Fraxinuspennsylvanica seedlings to flooding PlantPhysiol 66(2)267ndash271 doi101104pp662267

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2005 Recon-structing the temporal and spatial dynamics ofemerald ash borer in black ash A case study ofMichiganrsquos Roscommon County outlier siteIn Proc of the Emerald ash borer research andtechnology development meeting 2004 October5ndash6 Romulus MI FHTET-2004-15 andFHTET-2008-08 USDA Forest HealthTechnology Expertise Team MorgantownWV pp 21ndash22

SIEGERT NW T SECORD AND DG MC-CULLOUGH 2014a Submersion as a tactic toprevent emergence of emerald ash borer Agri-lus planipennis from black ash logs Agric ForEntomol 16(3)321ndash325 doi101111afe12057

SIEGERT NW DG MCCULLOUGH AM LIEB-HOLD AND FW TELEWSKI 2014b Dendro-chronological reconstruction of the epicentreand early spread of emerald ash borer in NorthAmerica Divers Distrib 20(7)847ndash858 doi101111ddi12212

SIEGERT NW RJ MERCADER AND DG MC-CULLOUGH 2015 Spread and dispersal ofemerald ash borer (Coleoptera Buprestidae)Estimating the spatial dynamics of a difficult-to-detect invasive forest pest Can Entomol147(3)338ndash348 doi104039tce201511

SIMS RA HM KERSHAW AND GM WICK-WARE 1990 The autecology of major tree speciesin the North Central Region of OntarioCOFRDA Rep 3302 NWO-FTDU TechRep 48 Forestry Canada Ontario RegionSault Ste Marie ON Canada 126 p

SLESAK RA CF LENHART KN BROOKSAW DrsquoAMATO AND BJ PALIK 2014 Watertable response to harvesting and simulated em-erald ash borer mortality in black ash wetlandsin Minnesota USA Can J For Res 44(8)961ndash968 doi101139cjfr-2014-0111

SMITH A DA HERMS RP LONG AND KJKGANDHI 2015 Community composition andstructure had no effect on forest susceptibilityto invasion by the emerald ash borer (Co-leoptera Buprestidae) Can Entomol 147(3)318ndash328 doi104039tce20158

TARDIF JC S DERY AND Y BERGERON 1994Sexual regeneration of black ash (Fraxinus

Journal of Forestry bull September 2017 445

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019

nigra Marsh) in a boreal floodplain AmMidl Nat 132(1)124 ndash135 doi1023072426207

TARDIF JC AND Y BERGERON 1999 Popula-tion dynamics of Fraxinus nigra in response toflood-level variations in northwestern Que-bec Ecol Monogr 69(1)107ndash125 doi1018900012-9615(1999)069[0107PDOFNI]20CO2

TELANDER AC RA SLESAK AW DrsquoAMATOBJ PALIK KN BROOKS AND CF LENHART2015 Sap flow of black ash in wetland forestsof northern Minnesota USA Hydrologic im-plications of tree mortality due to emerald ashborer Agric For Meteorol 2064ndash11 doi101016jagrformet201502019

TOWNSHEND E 2011 Black ash stand dynamicsand flooded jellyskin potential habitat quality in

rich hardwood swamps of the Great Lakes StLawrence forest region BSc thesis NipissingUniv North Bay ON Canada 43 p

WRIGHT JW AND HM RAUSCHER 1990Black ash Fraxinus nigra P 344ndash347 in Silv-ics of North America Vol 2 Hardwoods BurnsRM and BH Honkala (eds) USDA ForestService Agri Handbk 654 Washington DChttpswwwtreesearchfsfeduspubs1548

446 Journal of Forestry bull September 2017

Dow

nloaded from httpsacadem

icoupcomjofarticle-abstract11554354599906 by guest on 18 July 2019