Distribution and populations ofDistribution and populations ofArmillariaArmillaria in the regionin the region

R. Rippy

JOHN W. HANNAJOHN W. HANNA1,21,2, James A. Moore, James A. Moore11, Terry M. Shaw, Terry M. Shaw11,,Geral I. McDonaldGeral I. McDonald22, Mee, Mee--Sook KimSook Kim22, Ned B. Klopfenstein, Ned B. Klopfenstein22

11Department of Forest Resources, University of Idaho, Moscow, ID Department of Forest Resources, University of Idaho, Moscow, ID 838448384422USDA Forest Service, RMRS, 1221 S. Main St. Moscow, ID 83843USDA Forest Service, RMRS, 1221 S. Main St. Moscow, ID 83843

Hanna et al. 2004

ArmillariaArmillaria Species and Relative PathogenicitySpecies and Relative Pathogenicity

NABSa Species Pathogenicity Primary Host

I

VI

A. ostoyaeA. melleaA. tabescens

High

High

High?

Conifers

Hardwoods

Hardwoods

II

IX

A. geminaA. nabsnona

Moderate?

Moderate?

Hardwoods

Hardwoods

III

V

VII

XI

A. calvescensA. sinapinaA. gallicaA. cepistipes

Low

Low

Low

Low

Mixed

Mixed

Mixed

Mixed

X Unnamed Rarely pathogenic Mixed

aaNABSNABS = North American Biological Species= North American Biological Species

Hanna et al. 2004

IFTNC IFTNC ArmillariaArmillaria species summaryspecies summary

During the 1999 and 2000 field seasons over 8,500 During the 1999 and 2000 field seasons over 8,500 trees were sampled for trees were sampled for ArmillariaArmillaria, 1,310 isolates of , 1,310 isolates of ArmillariaArmillaria were collected from 23 IFTNC sites. were collected from 23 IFTNC sites. Based on over 8,000 somaticBased on over 8,000 somatic--incompatibility tests, incompatibility tests, these isolates were found to consist of 221 genets these isolates were found to consist of 221 genets (individual clones).(individual clones).

Of the 10 North American Biological Species 6 were Of the 10 North American Biological Species 6 were associated with the IFTNC sites.associated with the IFTNC sites.

Hanna et al. 2004

Huckleberry

Soldier Creek

Enterprise

Stanton

Hanna et al. 2004

NABS I NABS I –– Armillaria ostoyaeArmillaria ostoyae62 genets collected from 17 IFTNC sites62 genets collected from 17 IFTNC sites

Serious plant pathogen found throughout the Northern Serious plant pathogen found throughout the Northern Hemisphere that causes root and butt rot on diverse Hemisphere that causes root and butt rot on diverse woody plant hosts. woody plant hosts.

Adversely impacts commercial timber production by Adversely impacts commercial timber production by causing tree mortality and growth reduction. causing tree mortality and growth reduction.

Genetically diverse and may be comprised of several Genetically diverse and may be comprised of several races. Intraspecific hybrids are common.races. Intraspecific hybrids are common.

Hanna et al. 2004

NABS X NABS X –– Unnamed Unnamed ArmillariaArmillaria speciesspecies69 genets collected from 16 IFTNC sites69 genets collected from 16 IFTNC sites

Unnamed Unnamed ArmillariaArmillaria species found in the northwestern species found in the northwestern United States. United States.

Mainly saprophytic, living on dead organic matter. Mainly saprophytic, living on dead organic matter.

Hanna et al. 2004

NABS III, V, VII complex NABS III, V, VII complex ––A. calvescensA. calvescens, , A. sinapinaA. sinapina, , A. gallicaA. gallica58 genets collected from 14 IFTNC sites58 genets collected from 14 IFTNC sites

Group of similar species of Group of similar species of Armillaria Armillaria found throughout found throughout North America. North America.

Primarily saprophytic with low pathogenicity (less than Primarily saprophytic with low pathogenicity (less than 5%). 5%).

Hybridization is common between species.Hybridization is common between species.

Hybrids are difficult to identify.Hybrids are difficult to identify.

Hanna et al. 2004

NABS X / NABS III, V, VII NABS X / NABS III, V, VII complex hybridscomplex hybrids31 genets collected from 31 genets collected from 12 IFTNC sites12 IFTNC sites

Common within the Inland Common within the Inland Northwest. Northwest.

Previously grouped with NABS X Previously grouped with NABS X until their discovery using molecular until their discovery using molecular techniques.techniques.

Mainly saprophytic (less that 5% Mainly saprophytic (less that 5% pathogenic).

Hanna et al. 2004

pathogenic).

NABS IX NABS IX –– Armillaria nabsnonaArmillaria nabsnona1 genet collected from 1 IFTNC site1 genet collected from 1 IFTNC site

A rare species found primarily on hardwoods, such as A rare species found primarily on hardwoods, such as Alnus Alnus species, in riparian areas of western North America. species, in riparian areas of western North America.

IFTNCIFTNC’’s isolation of this species from s isolation of this species from Abies grandis Abies grandis is is one of the first reports of this species being isolated from a one of the first reports of this species being isolated from a conifer.conifer.

Thomas J. VolkThomas J. VolkHanna et al. 2004

Types of information collected from Types of information collected from Nutrition CoNutrition Co--op sitesop sites

Isolate IDIsolate ID

LocationLocation

Host SpeciesHost Species

Collection Type (Bark Fan, Rhizomorph, or Wood)Collection Type (Bark Fan, Rhizomorph, or Wood)

Host Collection (Dead or Alive)Host Collection (Dead or Alive)

Habitat TypeHabitat Type

Fertilization TreatmentFertilization Treatment

Species IDSpecies ID

Rock TypeRock Type

Hanna et al. 2004

Bark fan collection typeBark fan collection type

Bark fans from live trees indicate high pathogenicityBark fans from live trees indicate high pathogenicityof an of an ArmillariaArmillaria individual individual

Dead trees with bark fans and Dead trees with bark fans and resinosusresinosus and or wound and or wound periderm may indicate pathogenicity periderm may indicate pathogenicity

Dead trees with bark fans without Dead trees with bark fans without resinosusresinosus indicate indicate saprophytic behaviorsaprophytic behavior

Hanna et al. 2004

Wood samples are usually taken from trees that haveWood samples are usually taken from trees that haverecently been killed or are infected with a recently been killed or are infected with a pathogenic pathogenic ArmillariaArmillaria individual. individual.

Often dark lines known as zone lines can be seen in Often dark lines known as zone lines can be seen in infected wood.infected wood.

Wood collection typeWood collection type

Hanna et al. 2004

All species of All species of ArmillariaArmillaria form rhizomorphs. form rhizomorphs.

Most rhizomorphs are found on the surface of a root, some grow Most rhizomorphs are found on the surface of a root, some grow freely through the soil, and others can be found under the bark freely through the soil, and others can be found under the bark of a of a highly infected tree. highly infected tree.

Presence of rhizomorphs does not necessarily indicate pathogenicPresence of rhizomorphs does not necessarily indicate pathogenicity ity of an of an Armillaria Armillaria individual.individual.

Hanna et al. 2004

Rhizomorph collection typeRhizomorph collection type

DNA SequencingDNA Sequencing

Nuclear SmallNuclear Small rDNArDNA

ITS-1IGS-2 ITS-2

Nuclear Large rDNANuclear Large rDNA

IGS-1

5S5S

IGS-2

NuclearNuclearSmallSmallrDNArDNA

5.8S5.8SrDNArDNA

Ribosomal DNA regions currently being investigated forRibosomal DNA regions currently being investigated forphylogenetic signal include IGS, ITS, and LSU.phylogenetic signal include IGS, ITS, and LSU.

Hanna et al. 2004

Variable nucleotide positions of aligned nLSU sequences.

GENET 126bp 129bp 165bp 172-173bp 227bp 407bp 470bp 477bp 481bp 500bp 514bp 518bp 524bp 602bp 656bp 750bp 787bp 812bpT

T

AT C

A

A

A T

T

T

T

T

T C

C

C

C

T

GA

GA

GA

GA

GA T

T

T

T

C

C

C

C

C

C

C

T

T

C

G

G

G

A A

C

C

C

C

C

C

C

G

C

C

C

C

A

A

A

G

C

C

T

T

G

G

G

G

A

A

A

T

TA

T

NC1126, SSF4-B

NC765-B, P1404, R1348-B, R1389-B

837-A***, D82-A***, POR100***, R1299, ST1, ST8*, ST11-A*, U5, U16

T

T

T

T

A

A

C21**, M90** A

C G

G

G

G

T

A

A

A

837-B*** A

ST16** G

G

G

G

G

C

C

C

G

G

C C

C

A

GA T C

R

C G

C

C C

C

ST11-B* A T C

CTC C GA T

CT

C C

T

T

T

D82-B*** A T C GA T C

Y M C

T

T

A

T

T

OR10 A T C GA T C T A T

A

T

T

A

C

A

A

A

A

A

R A C

A

YOR1 A T C GA T C T Y G A A T

R A A TC G A CGA Y C TR1329 A T C

G A A TC G A CT C T TBC18F-B, NC1070-B A T Y

G A A TC G A C

A T TNM115, NM120, TS7, U73 A T C GA T C

A

T TA G

C T

BC18F-A, NC1070-A A T Y

A T T

A A T T

T

T

T

A Y G AY C M GGA T Y YR1374 A T C

T T

T C C G A C G

T

Y G A AC C G A

T

NC837, R1330 A T C GA T Y Y Y

G A A TC G A Y

T

T Y C

T C

C

T C

M

GA T C

GA T C

GA T C

~~

PC514 A T C

MNF4, NC94, NC540, NC765-A, P4352, R1348-A, R1389, SSF4-A, ST2 A T C

Hanna et al. 2004

Radial 50% majority rule consensus trees based on 24,000 trees from the Bayesian inference analysis of 850 base-pairs in the nLSU rDNA region. Scale bar = 0.001 substitutions per site.

0.98

0.98

0.001

0.64

** NABS IX*** NABS X

* NABS II

PC514

D82-B***

837-B***

BC18F-B,NC1070-B

BC18F-A,NC1070-A

NC765-B, P1404,R1348-B, R1389-B

R1329

NC1126,SSF4-B

MNF4, NC94, NC540,NC765-A, P4352, R1348-A,R1389, SSF4-A, ST2

ST11-B*

37-A***, D82-A***, POR100***,1299, ST1, ST8*,T11-A*, U5, U16

OR10

OR1R1374

NC837, R1330

NM115, NM120,TS7, U73

C21**, M90**

ST16**

0.001

NM115, NM120,TS7, U73

C21**, M90**

ST16**

837-A***, D82-A***, POR100***,R1299, ST1, ST8*, ST11-A*, U5, U16

D82-B***

837-B***

ST11-B*

MNF4, NC94, NC540,NC765-A, P4352, SSF4-A,ST2, R1389, R1348-A

NC765-B, P1404,R1348-B, R1389-B

NC1126,SSF4-B

PacificNorthwest

Rockies

Circumboreal

Hanna et al. 2004

Phylogenetic tree showing genetic diversity of Phylogenetic tree showing genetic diversity of Armillaria ostoyae Armillaria ostoyae based on the IGSbased on the IGS--1 DNA region1 DNA region

NC588 N. Idaho NC588 N. Idaho -- Soldier CreekSoldier Creek

NC542A N.E. Oregon NC542A N.E. Oregon -- EnterpriseEnterprise

NC435A N. Idaho NC435A N. Idaho -- StantonStanton

P416A N. Idaho P416A N. Idaho -- StantonStanton

NC403 N. Idaho NC403 N. Idaho -- StantonStanton

NC666A N. Idaho NC666A N. Idaho -- Dick’s CreekDick’s Creek

P2399 N. Idaho P2399 N. Idaho -- Priest RiverPriest River

NC436A N. Idaho NC436A N. Idaho -- StantonStanton

NC94 N. Idaho NC94 N. Idaho -- HuckleberryHuckleberry

NC435B N. Idaho NC435B N. Idaho -- StantonStanton

P4352 N. Idaho P4352 N. Idaho -- Priest RiverPriest River

P416B N. Idaho P416B N. Idaho -- StantonStanton

NC1126 N. Idaho NC1126 N. Idaho -- WhiskeyWhiskey

NC436B N. Idaho NC436B N. Idaho -- StantonStanton

NA142 S.W. Washington NA142 S.W. Washington -- T.T. MungerT.T. Munger

NC542B N.E. Oregon NC542B N.E. Oregon -- EnterpriseEnterprise

NC666B N. Idaho NC666B N. Idaho -- Dick’s CreekDick’s Creek

NM120A New.Mexico NM120A New.Mexico -- JemezJemez

NM115 New.Mexico NM115 New.Mexico -- JemezJemez

NM120B New.Mexico NM120B New.Mexico -- JemezJemez

HUA9242 Japan HUA9242 Japan -- Terashima et al. 1998Terashima et al. 1998

P255A N. Idaho P255A N. Idaho -- Priest RiverPriest River

P255B N. Idaho P255B N. Idaho -- Priest RiverPriest River

Pacific Pacific Northwest ANorthwest A

PacificPacificNorthwest BNorthwest B

RockiesRockies

Hanna et al. 2004

Factors associated with genetic races and Factors associated with genetic races and intraspecific hybridization within intraspecific hybridization within A. ostoyaeA. ostoyae

Adaptation to a variety of environmentsAdaptation to a variety of environments

May be responsible for differing levels of May be responsible for differing levels of pathogenicity and virulencepathogenicity and virulence

“Hybrid vigor” may contribute to enhanced “Hybrid vigor” may contribute to enhanced growth, survival, and/or pathogenicitygrowth, survival, and/or pathogenicity

Hybrids may adapt better to environmental Hybrids may adapt better to environmental changechange

Hanna et al. 2004

Does Does ArmillariaArmillaria affect tree growth?affect tree growth?

Hanna et al. 2004

Impact of Armillaria Root Disease on Tree GrowthImpact of Armillaria Root Disease on Tree Growth

Volume loss has been shown as high as 40% over 4Volume loss has been shown as high as 40% over 4--8 years8 yearsin eighteenin eighteen--yearyear--old Douglasold Douglas--fir (Cruickshank, 2000).fir (Cruickshank, 2000).

Hanna et al. 2004

Average mortality rates in a 10Average mortality rates in a 10--year period for sites year period for sites covering northern Idaho, northeastern Oregon, covering northern Idaho, northeastern Oregon, and eastern Washington vary from 8.8% for and eastern Washington vary from 8.8% for DouglasDouglas--fir, 4.2% for grand fir, and 1.5% fir, 4.2% for grand fir, and 1.5% for redcedar habitat types based on 8 sitesfor redcedar habitat types based on 8 sites(Moore et al., 2000).(Moore et al., 2000).

Growth loss and mortality due to Growth loss and mortality due to ArmillariaArmillariain British Columbia have been cited as in British Columbia have been cited as between 1.4 and 3.8 million mbetween 1.4 and 3.8 million m3 3 annuallyannually(White and Morrison, 1999).(White and Morrison, 1999).

Hanna et al. 2004

Does Does ArmillariaArmillaria affect tree growth?affect tree growth?

Influence of Armillaria ostoyae collected adjacent to growth plots on six-year gross-basal-area growth (sq.ft/a)

0

10

20

30

40

50

60

Spirit Lake Whiskey Huckleberry Soldier Creek Skookum Lake Dick's Creek

IFTNC Site

Ave

rage

incr

ease

in b

asal

are

a (s

q.ft/

a) Without A.ostoyae

With A.ostoyae

0 7752486777 10

*** *

# of plotsaveraged

* **α < .05 α < .10Hanna et al. 2004

!(!(

!(

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!( !(!(!(

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!( !(!(

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D

D

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D

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D

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D

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D

D

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D

D

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D

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D

D

Soldier Creek - Study SiteIntermountain Forest Tree Nutrition Co-op

D Plot corners

[ Bark fans live trees

!( NABS X

!( NABS X - III-V-VII Hybrid

!( NABS I

Plot 1300N & 200K

Plot 2300N & 200K2SO4

Plot 3100N & 0K

Plot 4100N & 0K

Plot 5200N & 0K

Plot 14300N & 0K

Plot 13200N & 0K

Plot 12600N & 0K

Plot 6200N & 0K

Plot 7Control

Plot 8100N & 0K

Plot 9200N & 0K

Plot 10300N & 0K

Plot 11300N & 0K

Tsuga heterophylla / Asarum caudatum

Tsuga heterophylla / Clintonia uniflora

western hemlock / queen cup beadlily

western hemlock / wild ginger

0 140 28070 Meters

´6-year Net- and Gross-Basal-Area Growth (sq.ft/a) in GREEN

59.7G57.5N

35.6G35.6N

55.0G55.0N

32.3G23.1N

32.4G32.2N

49.4G45.8N

41.8G41.8N

45.5G44.4N

51.3G51.3N

60.5G58.2N

42.5G42.0N

44.4G42.8N

49.8G49.8N

53.1G53.0N

1

2

Hanna et al. 2004

Influence of Armillaria ostoyae collected adjacent to growth plots on six-year net-basal-area growth (sq.ft/a)

05

101520253035404550

Spirit Lake Whiskey Huckleberry Soldier Creek Skookum Lake Dick's Creek

IFTNC Site

Ave

rage

incr

ease

in b

asal

are

a(s

q.ft

/a)

Without A.ostoyae

With A.ostoyae

4 72 50 7 7 7 6 8 10 7

*

*

**

* **

# of plotsaveraged

α < .05 α < .10Hanna et al. 2004

D

DD

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Plot 1512.1N & 290.2K

Plot 2300N & 170K

Plot 3300N & 170K

Plot 4300N & 170K

Plot 5600N & 170K

Plot 6512.1N & 49.8K

Plot 7300N & 0K

Plot 8300N & 340K

Plot 987.9N & 290.2KPlot 10

0N & 170K

Plot 11300N & 170K

Plot 1287.9N & 290.2K

Plot 13Control

Plot 14300N & 200K2SO4

Abies Grandis / Clintonia uniflora - Physocarpus malvaceus phaseGrand fir / queen cup beadlily - ninebark phase

Abies Grandis / Physocarpus malvaceus - Physocarpus malvaceus phase

Abies Grandis / Spiraea betulifolia

Grand fir / ninebark phase - ninebark phase

Grand fir / spiraea

LegendD plot corners

D plot centers

!( Genet 1 NABS I A.ostoyae

!( Genet 2 NABS X

!( Genet 3 NABS III,V,VII complex

!( Genet 4 NABS III,V,VII complex

[_ Barkfan on dead tree

[ Barkfan on live tree (active pathogen)

1 inch equals 61.886223 meters

0 280 560140 Meters

Huckleberry Creek - Study SiteIntermountain Forest Tree Nutrition Co-op´

19.3G1.1N

24.0G24.0N

26.6G18.0N

29.2G27.5N

20.2G15.9N

27.9G27.9N

29.3G21.8N

37.2G31.5N

31.9G21.9N

21.3G9.3N

27.4G9.9N23.9G

18.8N

31.6G30.9N

28.1G26.2N

6-year Net- and Gross-Basal-Area Growth (sq.ft/a) in GREEN

Hanna et al. 2004

D

D

D

D

D

D

D

D

D

D

D

DD

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[_

[_

[

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Spirit Lake - Study SiteIntermountain Forest Tree Nutrition Co-op

Plot 4300N & 0K

Plot 5300N & 170K

Plot 1600N & 0K

Plot 20N & 170K

Plot 3200N & 0K

Plot 6100N & 0K

Plot 7Control

0 130 26065 Meters

LegendD plot corners

D plot centers

!( Genet 1 NABS I A.ostoyae

!( Genet 2 NABS I A.ostoyae

!( Genet 3 NABS I A.ostoyae

!( Genet 4 NABS X

[_ Barkfan on dead tree

[ Barkfan on live tree (active pathogen)

[_ Wood Sample (Note: from Holodiscus discolor)

´

western hemlock Series

Thuja plicata

Tsuga heterophylla

western redcedar Series

Thuja plicata

western redcedar Series

Series

Series

Series 6-year Net- and Gross-Basal-Area Growth (sq.ft/a) in GREEN

27.7G24.2N

17.4G-12.9N

21.3G20.7N

28.7G24.7N

31.4G29.5N

19.4G14.7N

18.6G16.4N

1

2

Hanna et al. 2004

Influence of Armillaria species collected adjacent to growth plots on six-year net- basal-area growth (sq.ft/a)

0

10

20

30

40

50

60

Spirit Lake Whiskey Huckleberry SoldierCreek

SkookumLake

Dick's Creek

IFTNC Site

Ave

rage

incr

ease

in b

asal

ar

ea (s

q.ft

/a)

Without Armillaria

NABS X Only

X and I

NABS I Only

* ***

* **

# of plotsaveraged

α < .05 α < .10

0 2 50 0 7 07 4 2 1 7 4 0 4 6 2 2 30 0 7 6 1

Hanna et al. 2004

Factors that interact with Factors that interact with ArmillariaArmillaria totoinfluence tree growthinfluence tree growth

Interacting FactorsInteracting Factors ::

Habitat TypeHabitat TypeSoil CharacteristicsSoil CharacteristicsParent MaterialParent MaterialFertilizationFertilizationOther PestsOther Pests

Hanna et al. 2004

Future StudiesFuture Studies

Better characterize Better characterize ArmillariaArmillaria species with molecular species with molecular genetic techniques to better understand pathogenicity genetic techniques to better understand pathogenicity and virulenceand virulence

Assess direct effects of Assess direct effects of Armillaria ostoyaeArmillaria ostoyae on tree on tree growthgrowth

Evaluate effect of Evaluate effect of ArmillariaArmillaria species interactions on species interactions on tree growthtree growth

Investigate saprophytic Investigate saprophytic ArmillariaArmillaria species as a possible species as a possible biological control of biological control of Armillaria ostoyaeArmillaria ostoyae

Hanna et al. 2004

Tree growth is affected by a multitude of interacting Tree growth is affected by a multitude of interacting factors that include factors that include ArmillariaArmillaria. .

It is difficult to assess how nutritional treatments affect It is difficult to assess how nutritional treatments affect ArmillariaArmillaria impacts on growth without knowledge of preimpacts on growth without knowledge of pre--existing existing ArmillariaArmillaria influences.influences.

Pretreatment studies of Pretreatment studies of ArmillariaArmillaria distribution and distribution and impacts across a range of environments are needed as impacts across a range of environments are needed as baseline information. This baseline would help forest baseline information. This baseline would help forest managers better understand how treatments affect managers better understand how treatments affect ArmillariaArmillaria and overall growth of a stand.and overall growth of a stand.

Hanna et al. 2004

AcknowledgementsAcknowledgements

Intermountain Forest Tree Intermountain Forest Tree Nutrition CoNutrition Co--opop

and theand theUniversity of IdahoUniversity of Idaho

USDA Forest ServiceUSDA Forest Service

Mariann GarrisonMariann Garrison--JohnstonJohnstonMark KimseyMark KimseyHoward JenningsHoward JenningsBob AtwoodBob AtwoodPeter MikaPeter MikaLeonard JohnsonLeonard Johnson

James B. DonleyJames B. DonleyRaini C. RippyRaini C. RippyJane E. StewartJane E. StewartBryce A. RichardsonBryce A. RichardsonTroy HensiekTroy HensiekJoanne TirockeJoanne TirockeKwanKwan--Soo WooSoo WooJonalea R. TonnJonalea R. TonnPaul J. ZambinoPaul J. ZambinoDebbie S. PageDebbie S. Page--DumroeseDumroese

Hanna et al. 2004