Heart-rot and associated fungi in Alnus glutinosa stands in Latvia

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  • This article was downloaded by: [Colorado State University]On: 30 September 2013, At: 06:42Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: MortimerHouse, 37-41 Mortimer Street, London W1T 3JH, UK

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    Heart-rot and associated fungi in Alnus glutinosastands in LatviaNatalija Arhipova a b , Talis Gaitnieks b , Janis Donis b , Jan Stenlid a & Rimvydas Vasaitisa

    a Department of Forest Mycology and Pathology, Swedish University of AgriculturalSciences, SE-75007, Uppsala, Swedenb Latvian State Forest Research Institute Silava, LV2169, Salaspils, LatviaAccepted author version posted online: 01 Mar 2012.Published online: 26 Mar 2012.

    To cite this article: Natalija Arhipova , Talis Gaitnieks , Janis Donis , Jan Stenlid & Rimvydas Vasaitis (2012) Heart-rotand associated fungi in Alnus glutinosa stands in Latvia, Scandinavian Journal of Forest Research, 27:4, 327-336, DOI:10.1080/02827581.2012.670727

    To link to this article: http://dx.doi.org/10.1080/02827581.2012.670727

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  • ORIGINAL ARTICLE

    Heart-rot and associated fungi in Alnus glutinosa stands in Latvia

    NATALIJA ARHIPOVA1,2, TALIS GAITNIEKS2, JANIS DONIS2, JAN STENLID1 &

    RIMVYDAS VASAITIS1

    1Department of Forest Mycology and Pathology, Swedish University of Agricultural Sciences, SE-75007 Uppsala, Sweden,

    and 2Latvian State Forest Research Institute Silava, LV2169 Salaspils, Latvia

    AbstractThe interest in Alnus glutinosa (L.) Gaertn. as plantation species has increased during last years, but its prospects should beevaluated from the perspective of forest health. The aims of the present study were to: (1) estimate the incidence of stemdecay in Latvian A. glutinosa stands, (2) measure the extent of decay within individual stems and on a stand level and (3)identify decay-causing fungi. In four A. glutinosa stands, 450 trees were randomly sampled with an increment borer and thepresence/absence of decay was recorded. As a result, 112 sound-looking and 338 decayed trees were detected, and acorresponding number of wood samples were collected for fungal isolations. A total of 34 stems with decay symptoms werecut to measure the extent of internal decay. The incidence of decayed stems in studied stands was 75.1% on average. Thelength of the decay column was 7.795.4 m on average, and that of spongy rot was 4.294.5 m on average, implying thatyield losses for fully stocked 80-years-old A. glutinosa stand would comprise 49.2% of the total stand volume, and the lossesfrom spongy rot alone 30.5%. In total, 1134 isolates representing 68 fungal taxa were obtained. The most common decay-causing fungi were Inonotus radiatus and Armillaria sp.

    Keywords: Black alder, Inonotus radiatus, Armillaria, stem decay, wood-inhabiting fungi, yield losses.

    Introduction

    Currently, stands of black alder (Alnus glutinosa (L.)

    Gaertn.) comprise 5.1% (161,200 ha) of the total

    forest area of Latvia (Central Statistical Bureau of

    Latvia, 20082010). In Latvia, A. glutinosa typicallygrows on wet peatlands, usually comprising pure

    stands, or stands mixed with Alnus incana (L.)

    Moench., Betula spp., Populus tremula L. and Picea

    abies (L.) Karst. (Kundzins, 1969; Prieditis, 1993).

    Specific characteristics of this tree species are frost

    and waterlogging tolerance, a strong root system that

    penetrates both vertically and horizontally, adapta-

    tion to various soil conditions, and ability to fix

    nitrogen (McVean, 1953; Wheeler et al., 1986).

    Leaves are also nitrogen-rich and, after being shed,

    increase nitrogen concentration in soil (Cote &

    Camire, 1985; Dawson & Funk, 1981;

    Perez-Corona et al., 2006). A. glutinosa grows well

    on marshlands, riverbanks and other kinds of wet

    sites, and is an excellent pioneer species (Claessens

    et al., 2010; Fremstad, 1983; Obidzinski, 2004).

    Under suitable conditions it can be as productive as

    Fraxinus or Acer, yielding wood of high quality

    (Claessens et al., 2010), usable for wide variety of

    purposes, e.g. sawn timber, pulp and others

    (Claessens et al., 2010; Fennessy, 2004; McVean,

    1953).

    The combination of rapid early growth with a

    coppicing ability makes A. glutinosa suitable for a

    short rotation forestry (Wittwer & Immel, 1978;

    Wittwer & Stringer, 1985), while the capacity for

    pioneering and nitrogen fixation makes this species

    suitable for afforestation of former agricultural land

    and non-productive sites, as, e.g. reclaimed mining

    areas (Chodak & Niklinska, 2010; Kuznetsova et al.,

    2010; Pregent & Camire, 1985; Torbert et al., 1985;

    Vares et al., 2004; Wittwer & Immel, 1978). The

    species has also been used in mixed forest planta-

    tions to increase overall wood production (Bohanek

    & Groninger, 2005; Chodak & Niklinska, 2010), as

    interplanting with A. glutinosa was shown to have

    positive effects on growth of adjacent trees of other

    Correspondence: Natalija Arhipova, Department of Forest Mycology and Pathology, Uppsala BioCenter, Swedish University of Agricultural Sciences, PO Box

    7026, SE-75007 Uppsala, Sweden. E-mail: natalija.arhipova@slu.se

    Scandinavian Journal of Forest Research, 2012; 27: 327336

    (Received 29 August 2011; accepted 22 February 2012)

    ISSN 0282-7581 print/ISSN 1651-1891 online # 2012 Taylor & Francishttp://dx.doi.org/10.1080/02827581.2012.670727

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  • species (Cote & Camire, 1984, 1987; Hansen &

    Dawson, 1982; Paschke et al., 1989; Plass, 1977).

    Consequently, nowadays A. glutinosa is becoming

    increasingly important as a plantation species. Aside

    from commercial forestry, this tree species is im-

    portant for riparian ecosystems and their biodiver-

    sity, providing habitats for specific wetland flora and

    fauna and stabilising riverbanks (Brown et al., 1997;

    Claessens et al., 2010; Popovska et al., 2008;

    Prieditis, 1997).

    The potential of different tree species considered

    for afforestation needs to be evaluated from many

    points of view, and the aspect of forest health is an

    important consideration. As in numerous countries

    with intense management of boreal temperateforests, heart-rot of standing trees is a considerable

    problem also in Latvian forestry. Based on observed

    average heart-rot incidence, spread of the decay

    inside a stem, and applying stand growth models it

    was estimated that in fully stocked stands of P. abies

    volumes of wood, degraded by the heart-rot

    comprise about 20 m3 ha1 at the age of 40 years,

    5560 m3 ha1 at the age of 60100 years and about90 m3 ha1 at the age of 120 years, corresponding to

    616% of a total standing volume (Arhipova et al.,2011a). Similar situation was also observed in fully

    stocked pure stands of A. incana, where volumes of

    decayed wood comprised 32 m3 ha1 at the age

    of 4550 years and 60 m3 ha1 at the age of6065 years, corresponding to about 10% and20% of all standing volume (Arhipova et al.,

    2011b). To date, however, no published data in

    this respect are available for A. glutinosa.

    Generally, A. glutinosa is regarded a short-living

    tree species, starting to die out naturally at about

    60 years of age under Central European conditions

    (Vyhldkova et al., 2005). However, depending on

    the region and growth conditions, the species might

    reach age of 100160 years (Claessens et al., 2010).Some authors noted that A. glutinosa is usually

    attacked by a stem rot at the age of 5070 years,especially on wet sites (Claessens, 2005; Claessens

    et al., 2010; Immler, 2004; Kotar, 2000). Moreover,

    there are observations that a large proportion of trees

    with a diameter at breast height over 30 cm are

    attacked by decay fungi and become susceptible to

    stem-breakage (Ilisson et al., 2004), or have stem

    cavities (Remm et al., 2006). In his review McVean

    (1953) named polypore Inonotus radiatus (Sowerby)

    Karst. as the most important cause of heart-rot in

    A. glutinosa, which was later supported by the

    observations from Central Europe (Schumacher

    et al., 2001, Vyhldkova et al., 2005). To date,

    information on fungal communities in living stems of

    A. glutinosa is scarce, comprising only a couple of

    records of endophytic fungi (Fisher & Petrini, 1990;

    Moricca, 2002). In this context, the aims of the

    present study were to: (1) estimate the incidence of

    stem decay in Latvian A. glutinosa stands, (2)

    measure the extent of decay within individual stems

    and on a stand level and (3) identify decay-causing

    and other fungi that inhabit living A. glutinosa stems.

    Materials and methods

    Field work

    The fieldwork included: (1) sampling living stems of

    A. glutinosa for estimating the frequency of decay

    and discoloration and for subsequent fungal isola-

    tion, (2) felling and dissection of decayed stems

    containing decay and measuring the extent of

    decay. Five 51- to 84-year-old A. glutinosa stands

    of 0.53.6 ha in size were investigated (Table I). Thestands were located in central-eastern Latvia,

    Kalsnava forest district (56.6838 N, 25.9678 E). Atotal of 450 trees have been sampled at 1020 cmheight using an increment borer (Table I) and the

    presence/absence of stem decay was recorded follow-

    ing visual examination of each core. Each tree was

    sampled once, extracting 10- to 20-cm-long bore

    cores. All 450 wood samples were placed into sterile

    plastic tubes and transported to the laboratory, and

    full length of the cores was subsequently used for

    fungal isolations. In each stand, trees were sampled

    by random, always choosing the most adjacent tree

    to the one previously sampled. From those sampled

    trees, 34 decayed stems (as determined by presence

    of decay in the extracted cores) were felled and

    dissected. Age, height, stem diameter at breast

    Table I. Characteristics of investigated A. glutinosa stands and number of trees cut for heart-rot examination.

    Age, years Species composition (%)* Forest site type Sampled trees, no. Decayed stems (%) Cut trees, no.

    51 100 A.g. Mercurialiosa** 100 98 74 80 A.g.20 A.i. Filipendulosa 100 82 1476 60 A.i.30 A.g.10 P.a. Oxalidosa turf.** 150 53 2084 80 A. g.20 A. i. Oxalidosa turf.** 100 78

    Average 75

    *A.g. Alnus glutinosa; A.i. Alnus incana; P.a. Picea abies.**Drained (ameliorated).

    328 N. Arhipova et al.

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  • height (d.b.h.) and stump diameter were assessed for

    every tree. Age of the felled trees ranged from 74 to

    81 years (7793 years), stump diameter from 15.3to 43.1 cm (28.297.5 cm), d.b.h. from 14.1 to36.3 cm (23.296.1 cm) and height from 13.2 to27.3 m (21.893.5 m). Two types of decay weredistinguished: (1) discoloured wood without or with

    slight changes in mechanical properties, throughout

    the paper referred as decay and (2) decomposed

    wood squeezable with fingers, throughout the paper

    classed as spongy rot. Columns of the spongy rot

    were always situated inside decay columns, being

    shorter in length and smaller in diameter. Diameters

    at stump level and the total column lengths were

    separately measured both for decay and, when

    present, for spongy rot.

    Isolation and identification of fungi

    In the laboratory, the procedure of fungal isolation

    from wood samples was performed as in our earlier

    studies (Vasiliauskas & Stenlid, 1998; Vasiliauskas

    et al., 1996). Briefly, the isolation was done next day

    after the sampling. Overnight samples were stored in

    refrigerator at 48C. All samples were flame-sterilised,placed on Hagem agar (Modess, 1941) media and

    incubated at 198C in the dark for 12 weeks. Toobtain pure cultures, the emerging mycelia were

    subcultured in separate Petri dishes, containing

    Hagem agar media. After 23 weeks of incubation,all pure cultures were examined under the light

    microscope (Leica DM400B) and grouped accord-

    ing to morphological features of the fungal myce-

    lium. From those, several species and genera

    (possessing distinct mycelial morphotypes and re-

    peatedly sequenced in our previous studies) were

    identified microscopically (Table III).

    One to three representatives from the morpho-

    types that have not been identified microscopically

    were subjected to a molecular identification

    (Vasiliauskas et al., 2004, 2005). DNA extraction

    and PCR amplification followed established proto-

    cols (Karen et al., 1997). After the amplification,

    PCR products were purified using Calf Intestine

    Alkaline Phosphatase (CIAP) (Fermentas GmbH,

    St. Leon-Rot, Germany) and Exonuclease I (Exo I)

    (Fermentas GmbH, St. Leon-Rot, Germany) and

    sent for Sanger sequencing (Sanger et al., 1977) to

    Macrogen Inc. (Seoul, Republic of Korea). For the

    ITS region sequencing, primer ITS4 was used for

    every specimen. All sequences were aligned and

    manually edited using Lasergene software package

    SeqMan (version 5.07, DNASTAR, Madison, WI,

    USA). BLAST (Basic Local Alignment Search Tool)

    searches (Altschul et al., 1997) were performed

    using two reference databases one at the Depart-ment of Forest Mycology and Pathology, Swedish

    University of Agricultural Sciences, and one of the

    GenBank (http://www.ncbi.nlm.nih.gov/blast). The

    ITS sequence homology was set at 98100% forspecies level and 9497% for genus level as in ourprevious studies (Arhipova et al., 2011a, 2011b;

    Bakys et al., 2009a, 2009b, 2011; Vasiliauskas et al.,

    2004, 2005). ITS sequence information for selected

    isolates was deposited in the GenBank (Table III).

    Volume calculations and statistical analyses

    Stand characteristics were obtained from the stand

    inventory data in the Forest State Register

    (Mr. Indulis Brauners, Latvian State Forests Inc.,

    personal communication). Calculations of decayed

    log volume were based on actual lengths of decay

    and spongy rot in the analysed trees (Arhipova...

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