decaying logs as key habitat in tasmania's wet sclerophyll production forests

8/14/2019 Decaying Logs as Key Habitat in Tasmania's wet sclerophyll production forests

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119Tasforests Vol. 13 No. 1 December 2001

Not just w aste wood: decaying logs

as key hab itats in Tasman ias wetsclerophyllEucaly ptus obliqua

production forests: the ecology

oflarge and small logs compared

Abstract

Large diameter decaying logs are a characteristic

feature of unmanaged wet sclerophyll forests in

Tasmania. In production coupes, however, rotation

lengths of around 80 y ears will eventually lead

totheir elimination. This paper describes an

on-going st udy into whether small diameter logs

derived from small trees left at the end of the

rotation are likely to follow the same successional

pathways as large diameter logs, thus supporting

the same range of decay types and fungal and

invertebrate biodiversity. The studys findings

may indicate whether some concerns over the use

of dead wood as industrial fuelwood are justified.

It may also indicate whether there is a need to

cater specifically for the maintenance of large

diameter trees and logs in managed coupes,

orwhether smaller ones will suffice.

Introdu ction: coarse woody d ebris as a k ey

resource in m anaged forests

The importance of coarse wood y d ebris

(CWD) in Australian forests has gone largely

M. Yee1,3*, Z.-Q. Yuan1,3 and C. Mohammed 1,2,3

1School of Agricultu ral Science,

University of Tasman ia, Sand y Bay, Hobart 70052CSIRO Forestry and Forest Products,

GPO Box 252-12, Hobar t 70013CRC for Sustainable Prod uction Forestry,

GPO Box 252-12, Hobar t 7001

un recognised ap art from a few recent

stud ies (Meggs 1996; Grove a nd Stork 1999;

Lindenmayer et al. 1999; Grove 2002). This

dearth of research interest in CWD m ay

beattributed to the cryptic nature of the

organisms th at inhabit it, the short-term

natu re of most research fund ing relative

tothe decay process, and a general attitude

that wood left on the forest floor is waste.

Yet decaying logs on th e forest floor are

anextremely valuable biological resource.

They support a rich diversity of invertebrates

and fungi, as has been documented

repea tedly in the scientific literatu re (Elton

1966; Harmon et al. 1986; Fran klin et al. 1987;

Speight 1989; And ersson and Hytteborn

1991;Parson et al. 1991; Esseen et al. 1992;

Kirby an d Drake 1993; Kaila et al.1994;

Jonsson and Kruys 2001). Coarse woodydebris p rovides habitat for saproxylic (dead-

wood dependent) insects (Key 1993) and for

wood decay fun gi (basidiomycetes), wh ich

are the primary decomposers of dead wood

and hence contribute to nu trient cycling

(Edmonds and Eglitis 1989; Speight 1989;

Edmond s and Marra 1999).

In many Northern Hemisphere forests, the

dynamics of CWD recruitment have been* Corresponding authore-mail: ma [email protected]


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grossly altered across the land scape du e

tocenturies of timber harvesting and the

removal of firewood (Gore and Patterson

1986; Edm ond s and Marra 1999). This hascontributed to the d ecline and in some cases

extinction of a range of insect and fungal

species reliant on CWD (Kirby and Drake

1993; Visnen et al. 1993; kland et al.

1996;Hgv ar a nd k land 1997; Kaila et al.

1997;Martikainen et al. 1999; Sipp ola 1999;

Eriksson 2000; Khler 2000). In these regions,

a d isproportionately large percentage of CWD

dep enden t species are either red-listed or

considered rare and threatened (Kirby

and Drake 1993; Raniu s and Jansso n 2000;

Ehnstrm 2001). Coarse woody debris is nowthe subject of extensive conservation research,

with many recent studies confirming the

particular value of larger diameter logs for

biodiversity conservation (Wood 1978; Araya

1993; Edm ond s and Marra 1999; Kru ys and

Jonsson 1999; Kolstrm and Lumatjrvi

2000; Ranius a nd Jansson 2000). Forest

management agencies in Europe and North

America increasingly recognise the need for

mod ification of forest pr actices (Kaila et al.

1997; Bragg an d Kersh ner 1999; Hagan

and Grove 1999; Grove 2001; Ehn strm

2001;Schiegg 2001). These includ e thedevelopment and implementation of action

plan s for red -listed species (Jonsell 1999;

Keyet al. 2000; Sverdrup-Thygeson 2001), and

active man agement to m aintain or r estore

CWD levels, particularly for larger diameter

material (Hard ing and Alexand er 1994).

Produ ction forestry in Tasman ia has a

relatively short history (c. 150 yrs), meaning

that its long-term impacts may not yet be

apparent. How ever, it is clear that successive

logging of native wet sclerophyll regrowthforests on p lanned rotations will alter CWD

quality and quantity (Meggs 1996). Inevitably

only smaller diameter ( 100 cm) CWD being prod uced w ithin the

forest coup e. If there are su ites of species

inthese forests that are d epend ent on larger

diameter CWD, then the risk that they may

be extingu ished from harv ested areas within

prod uction forests is only too app arent.

The ecology of log decay

The decomposition of CWD is an extremely

complex process, varying amongst differenttree species (Schwarze et al. 2000) an d

according to prevailing environmental

cond itions. As CWD decomposes, it passes

through various stages, from solid w ood

tomaterial devoid of any structure. The

metabolic and ph ysical actions of the wood

decay fungi chemically and structurally

change the wood , resulting in a

characteristic rot type. For examp le, brow n

rotted wood arises when so-called brown-

rotfungi selectively remove cellulose and

hemicellulose from the wood, leaving aresidu e of slightly mod ified lignin. By

contrast, white-rot fungi utilise all

components of the wood cells, removing

lignin, cellulose an d hem icellulose an d

leaving the w ood bleached, with a spon gy,

stringy or laminated structure (Kaarik 1974;

Gilbertson 1984; Schw arze et al. 2000).

Rot types can be d ifficult to define, as they

can vary in colour an d texture dep ending

onthe decompositional stage. Different

types are often d efined by visual and other

subjective characteristics. A log is genera llyinvad ed b y a su ccession of fungi, attacking

it at different positions d epend ent up on

environ men tal cond itions. Therefore, across

time and space, a log can exhibit man y rot

types; for exam ple, a sing le cross-section

canhave a particular heartwood rot,

sapw ood rot and rot pockets. To add to the

complexity, several fungi may bring about

similar typ es of rot, and each fun gus is not

confined to on e type of rot (Parkin 1942),

thus p osing m any challenges in studying

rotted w ood.

In old living trees, a suite of fungi may

bepresent an d may initiate decay of the

heartwood before the tree dies. How ever,

such fungi may not be present in a younger,

smaller d iameter tree, such that w hen a

young tree dies it may follow a different

decomp ositional path way. This potential

difference between large and small logs

maywell influence faunal diversity, as


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saproxylicinsects have specific microhabitat

requ irements. For examp le, some lucanid

and prostomid beetles are only foun d in red-

brown rotted wood in advanced stages ofdecay (Araya 1993; Lawren ce and Britton

1994). By contra st, some tipu lid flies live

only in wood invaded by white rot fungi

and with a very high moisture content,

and some elaterid beetles prey on th ese

tipulids (Dajoz 2000). Therefore,

understanding the relationships between

rottypes and the associated faun a is a

prerequisite for developing meaningful

indicators with which to monitor and

manage CWD dependent communities

inprodu ction forests.

Project objectives

The three-year project described here

wasset up by the Un iversity of Tasmania,

Forestry Tasmania and Gunn s Limited, and

is based at the Cooperative Research Centre

for Sustainable Production Forestry in Hobart.

The four objectives of the project are,

inregard to Eucalyptu s obliqua CWD in

Tasmanias wet sclerophyll forests:

1. To better und erstand the ecology of

decomposing CWD, in terms of the

associated wood decay fungi and

sap roxylic insects.

2. To investigate differences in fun gal and

insect biodiversity between small and

large diameter logs. Small logs range

from 3060 cm diameter, representative of

a tree felled at 6080 years ag e, and large

logs have a d iameter greater than 100 cm,

representative of a 150-year-old tree.

3. To provide a checklist of the fungal and

insect species dep end ent on E. obliqua

CWD in wet sclerophyll forests.

4. To look for direct associations between

decayed wood, wood decay fungi and

sap roxylic insects.

The research is being condu cted in wet

sclerophyllEucalyptus obliqua forest, in

Tasmanias Southern Forests about 60 km

south -west of Hobart (Figure 1). Of the ten

sites selected, six are in the Warra Valley (a

focus for long-term ecological research, asdescribed elsewhere in this issue), and four

in the West Picton Valley. Half the sites are

in regrowth and half in mature-age oldgrow th

forests. Regrowth sites were 2040-year-old

forests regenerated following clearfelling in

the 1960s1970s. Allsites h ave soils derived

from Jurassic dolerite, and all have a similar

slope/ aspect, with elevation ran ging

between 100 m and 150 m above sea level.

In this study, investigations have been

restricted to E. obliqua logs of anintermediate decay stage. The visualcharacteristics of this stage are th e lack of

bark, coupled w ith the p resence of decayed

sapwood, heartwood showing d istinct rot

type p atterns and colour, and some solidwood . At this stage, log shap e is retained.

This stage was chosen because:

1. Insects occurring in this intermediate

decompositional stage have been shown

to be the most vulnerable to logging

effects (Helivaara and Visnen 1984).

2. It is at this intermediate decompositionalstage that the actions of wood decayfungi are m ost prom inent, exhibiting a

characteristic rot type.

3. This choice thereby complements the on-

going Warra LTER log decay study, which

is looking a tE. obliqua logs in an earlydecay stage (Bashford et al. 2001).

The insect compon ent of the research has

also been confined to the beetles (Coleoptera).

Amongst the complex assemblage of

saproxylic insects present inany forest, beetlesare numerically a very important group,

exceeding all other insect group s in terms of

numbers of families, species and individuals.

Functionally, they are also very important,

with m embers in arange of feeding gu ilds,

includ ing w ood feeders (xyloph ages),

fungus feeders (mycophages), detritivores

and predators (Gilbertson 1984; Dajoz 2000;

Speight 1989). Many beetle species shar e an

intimate relationship with wood decay fungi


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and dead wood (Gilbertson 1984; Haa ck and

Slansky 1987; Lawrence 1989; Dajoz 2000),

for instance, carrying sp ores on th eir body

surface or dispersing sp ores via their frass

(Paine et al. 1997; Garcia and Mor rell 1999).In addition, beetles have a relatively well-

un derstood ecology at the family level and

are r elatively easy to iden tify, especially to

morphospecies (Oliver and Beattie 1996).

They are also robust for transport and

han dling. For all these reasons, they have

been the subject of more comprehensive

research elsewhere than have other groups,

thu s allowing this stud y to be more

readilycomparable.

Research objective 1.Developing a better

understanding of the ecology of decomposing

CW D

This research objective is being conducted insix of the ten stu dy sites (3R an d 3O: Figure 1).

At each site, three large and three small

E.obliqua logs have been destructively

sampled , totalling 18 in each size class. Each

log was saw n into th ree (1 m length ) sections

at the apical, middle and basal positions. A

5cm cross-sectioned disc was taken from each

end of the 1 m long section, giving six discs

per log. Each disc was photograp hed and

coded, and the rotted wood described. To

Figure 1. Location of stu dy sit es in Tasmanias Southern Forests. R = regrowth; 0 = oldgrowth forest.

B = location of the Warra LTER log decay study. Destructive sampling was conducted at sites that are

underlined.

HuonRiver

River

Picton

RiveauxRd

Rd

Arve

Rd

Edw

ards

South

WeldRd

WarraRd

Manuka

Rd

Pic

ton

Rd

R

R

B

O O

OR

R

O

R

to Geeveston

[

N

O

PICTON

VALLEY

WARRA


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Table1.TherottypeclassificationforEucalyptusspp.fallenlogsusedinthepresentstudy,comparedwithearlierclassificationsbasedonlogsandmaturetrees.

Rottypes

inthesamerowarethoughttobethesamerottype.

Whereavailable,informationonthelocationoftherottypeisindicatedas(H)attackingheartwood,a

nd(SF)

attackingsurface/sapwood.

Meggs(1996)

Rot

Thisstudy(2001)

variousEucalytusspp.

Mesibov(1988)

Parkin(1942)

Refshuage(1938)

type

E.o

bliqua

includingE.obliqua

E.o

bliqua

E.regnans

E.regnans

1

Red-brownwetblocky

Redblockyrot

Blockyrot

Browncub

icalrot

Browncubicalrot(H)

muddyrot(H)

Redblockyrotwithseams

ofwhitefungalhyphae

Orange/redclayeyrot

Mudgutrot

2

Browndrycrumbly

Orange/red/brown

Friable/crumblyrot

chalkyrot(H)

crumblyrot

Yellowcrumblyrot

3

Yellowstringysurfacerot(SF)

Softyellowfibrousrot

Spongy/fibrousrot

Yellowishstringyrot

4

Whitestringy/spongyrot(H)

Spongywhiterot

Whitespongyrot

Largewhitepocketro

tof

whitestringytype

5

Brownstringyrot(SF&

H)

Orange/red/brown

Yellowbro

wnspongy

Brownstringyrot(H)

fibrousrot

rot(H)

6

Whitejellyrot(SF)

Other(includesbluestain

Skeletalrot

Largewhitepocketro

t(H)

fungiandwet,jelly-likerot)

7

White/brownpocketrot(SF)

Whitepocketrot

Smallwhitepocketro

t(H)

Smallbrownpocketrot(H)

8

Otherdiscoloured/sta

ined

Brownstainassociatedwith

wood(H)

smallwhitepocketrot(H)


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isolate the wood decay fungi (Basidiomycete),

samples of the rotted wood were taken to the

laboratory, excised, and incubated on selective

med ia. The three 1 m sections w ere intensively

examined for beetles, noting w here and in

which rot type any beetles occurred.

A classification scheme for rot types has

been devised, based on general appearance,

colour, texture and position. As men tioned

above, the classification of rot types can be

extremely subjective, partly because there

are so many variables involved. The scheme

developed for this stud y aims to build on

past stu dies in this field w hile reducing the

ambigu ity and confusion that has arisen

from su ch stud ies (Table 1).

So far in this study, eight rot types have been

found , and app roximately 700 basidiomycete

isolates have been obtained from m aterialgrow n on selective med ia. At this

stage,isolates have been identified to

morphospecies based on colony morphology

and biochemical tests. Molecular techniqu es

will be used to assist in id entifying

isolatesto species level. This will involve

comparing the rotted wood isolates with

isolates derived from iden tified basidiocarps

collected from the field, and from reference

collections held at Forestry Tasmania an d

CSIRO Forestry and Forest Products Clayton

Laboratories. The resu lts of this stud y will

be compared with those of two earlier

studies (Refshuage 1938; Parkin 1942) that

looked at the fungal species responsible for

particular types of rot.

So far, about 104 beetle morphospecies have

been hand-collected from the log sections.

Thirty-six per cent have been identified

togenus level, and further efforts will be

madeto identify these to species level. The

mostcommon rot typ e found in the large

diameter logs has been Type 1 (red brown

blocky mud dy heartw ood rot). Of the 15

most commonly found beetles, eight are

primar ily associated w ith this rot type

(Table2), which appears to arise through

theenzymatic activities of two unidentified

basidiomycete species (coded LF213 and

LF485). Early results indicate that themuddy rot sub-type (also known as clayey

rot or mudguts: Meggs 1996; Mesibov 1988)

is the later decay stage of the blocky rot sub-

type (also known as cubical rot: see Table 1).

As this rot type is m ost consistently foun d

inlarge logs, successive short rotations

resulting in sm all logs could poten tially

impact on beetle communities dependent

onthis rot type. This rot type was rarely

present in small diameter logs, so the

Table 2. The 15 beetle species most commonly found in seventy -two 1 m log sections. Species marked

with an asterisk are those particularly associated with the red-brown blocky muddy rot type (Type 1).

Species Family Feeding Guild

Prostomis atkinsoni* Prostomidae xylophagousParablax spp.* Elateridae predatory

Pycnomerus sp. 1* Zopheridae xylophagousDryopthorus sp 1.* Curculionidae xylophagous

Coripera deplanata Tenebrionidae xylophagousCossonus sim soni* Curculionidae xylophagousLissotes cancroides Lucanidae xylophagous

Aleocharinae (STAPH 35) Staphylinidae predatoryToxeutes arcuatus Cerambycidae xylophagous

SCIRT6* Scirtidae detritivorousPhilothermus tasmanicus* Cerylonidae mycophagousChrysophtharta bimaculata Chrysomelidae other

Cryptorhynchinae (CUR3) Curculionidae xylophagousTEN9* Tenebrionidae xylophagous

Syn desus cornutu s Lucanidae xylophagous


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conservation of this particular beetle

community may be difficult unless specific

measures are taken to en sure its survival.

Research objective 2.Investigating

differences in fungal and insect biodiversity

between small and large diameter logs.

This research objective is being conducted

atall ten study sites (Figure 1). The aim

istoassess whether smaller diameter logs

support fungal and beetle assemblages

similar to those of larger diameter logs. For

the beetle component, log emergence traps

are being used to monitor beetle emergence

from d ecaying logs over 16 months.

Trapp ing has advantages over hand

samp ling in that it provides a quan titative

meth od of collecting saproxylic beetles, it

can collect over extended periods (thu s

provid ing information on seasonality),

itisnon-destructive, and it requires less

effortfor data retrieval. It is also relatively

unbiased in terms of operation error.

At each site, six traps have been erected,

three on larger diameter logs and th ree

onsmaller diameter logs (60 traps in total).

Thelog emergence traps used (Figure 2) are

mod ified from the d esign u sed in the Warra

LTER log d ecay study (Bashford et al. 2001).

A two-metre long section of the log is

encased with gauze net material (Broadway

Textiles: Trilobal cloth). A collecting bott le

ispositioned at the top of the trap, to catch

beetles whose behaviour is to move tow ards

the light, and th ere is a container at th e base

to catch those that drop when emerging.

Material was attached to the log using a

staple gun, and wooden stakes were placed

within the trap to keep the m aterial off

thelog. Ethylene glycol was u sed a s the

preserving fluid. Traps were erected in

SeptemberOctober 2000 and will be

mon itored for 18 mon ths, un til April 2002.

Samples are being collected m onthly d uring

summer and bimonthly during winter.

At the end of the trapping period, particular

logs that have specific beetles emerging will

be cut up to identify the wood rot type from

wh ich they emerged . Fung i will be isolated

from the various rot typ es derived from this

destru ctive analysis and g rown in culture

for DNA extraction and profiling. In this

way, the beetle-fungi-rot type associations

iden tified from Research Objective 1 can

bevalidated.

Research objective 3.Providing a checklistof the fungal and insect biodiversity dependent

on Eucalyptus obliqua CWD

A comprehensive checklist of insect and

funga l species associated withE. obliqua

CWD will be compiled from specimens

found and identified du ring this stud y. This

baseline information will prove valuable for

futu re stud ies at the Warra LTER Site. As a

key resource demonstrating the ecological

Figure 2. The log emergence trap design used in this study.


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value of CWD, it shou ld also prove v aluable

in forest planning . This is especially

relevant given the current debate over

theintensification of management of theSouthern Forests and the pr ospect of CWD

(biomass, residue, waste) being used for

power generation.

Research objective 4. To look for direct

associations between decayed wood, wood decay

fungi and saproxy lic insects

Basidiomycete fung i have been isolated from

the frass ofToxeutes arcuatus larvae and other

selected beetles. It is possible that such

species may act as dispersers for fung i, as hasbeen recorded elsewhere (Paine et al. 1997;

Garcia and Morrell 1999). Looking for direct

relationships between beetles and fungi will

provide a deeper understanding of the nature

and ecological significance of this association .

Ou tcomes for the forest indu stry

The ecological information from this

research will contribute to a ssessing w hether

curren t forest p ractices are ecologically

sustainable, developing appropriateindicators for monitoring forest biodiversity,

and proposing prescriptions that ensure the

conservation of these diverse dead wood-

associated comm un ities. Key issues to

wh ich this research relates includ e how best

to conserve CWD at the coupe level and in th eman aged land scape. For instance, wh at is the

best arrangement for retaining habitat trees so

thatthey can continue to fulfil their ecological

function in m anaged coupes long after their

death ? Is a land scape-level app roach to

CWD management (incorporating informal

or formal reserves) sufficient, or w ill

prescriptions also be required w ithin coup es

to ensure the survival of larger diameter

ma terial? Most critical, at presen t, is the

issue of how best to minimise any threat

todead wood-associated biodiversity posedbythe intend ed h arvesting of CWD as a

biofuelfor p ower generation.

Acknowledgements

This research was sup ported by the

Australian Research Council, Forestry

Tasmania and Gun ns Ltd. Marie Yee was

granted a University of Tasmania Strategic

Scholarship and a top-up scholarship from

the CRC for Sustainable Production Forestry.

Simon Grove and Tim Wardlaw of ForestryTasman ia are thanked for comments on

themanuscript.

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