Biological Conservation 117 (2004) 359–373
www.elsevier.com/locate/biocon
Lichens and bryophytes on Eucalyptus obliqua in Tasmania:
management implications in production forests
G. Kantvilasa,*, S.J. Jarmanb
a
Tasmanian Herbarium, Private Bag 4, Hobart, Australia 7001
b
Forestry Tasmania, GPO Box 207, Hobart, Australia 7001
Received 7 March 2003; received in revised form 25 July 2003; accepted 1 August 2003
Abstract
Thirty lichen and 25 bryophyte species have been recorded from the buttresses of Eucalyptus obliqua, the dominant tree in wet
sclerophyll forest at the Warra Long-Term Ecological Research Site in southern Tasmania. The flora, characterised by four major
associations, is very distinctive, containing a relatively high number of eucalypt specialists, particularly among the lichens. A general trend towards increasing species richness with increasing tree diameter is apparent and is attributed mainly to increasing habitat
diversity on the buttresses. The increase in species occurs without the loss of pioneers; thus succession involves addition rather than
replacement of species. The relationships between epiphytes, tree age and forest age are complicated by the periodic occurrence of
fires in the forest. Nevertheless, potential oldgrowth indicators are identified and the possible effects of current silvicultural practices
on the conservation of the species are discussed.
Crown Copyright # 2003 Published by Elsevier Ltd. All rights reserved.
Keywords: Tasmania; Australia; Lichens; Bryophytes; Biodiversity; Eucalyptus obliqua; Oldgrowth forest; Species richness; Tree age; Diameter;
Oldgrowth indicators; Epiphyte succession
1. Introduction
The Warra Long-Term Ecological Research Site is an
area of 15 900 ha in southern Tasmania, about 60 km
west south-west of Hobart (Fig. 1) (see Brown et al.,
2001, for an overview). The Site includes extensive forests administered by Forestry Tasmania and the Tasmanian Department of Primary Industries, Water and
Environment. A silvicultural systems trial is being conducted at the Site (Hickey et al., 2001), one of the aims
of which is to investigate the impact of several alternative harvesting and regeneration techniques on the
forest biota. The results presented here are part of the
trial and are a continuation of our earlier work in the
area on lichens and bryophytes (Jarman and Kantvilas,
2001a, b).
A total of 134 lichen species and 144 bryophyte species has been recorded from unlogged wet sclerophyll
forest in the silvicultural systems trial (Jarman and
Kantvilas, 2001a). Details of the distribution of these
* Corresponding author. Fax: +613-6226-7865.
E-mail address: [email protected] (G. Kantvilas).
species within the forest and their habitat ecology are
given in Jarman and Kantvilas (2001b). In the present
paper we focus on the epiphytic flora of Eucalyptus
obliqua L’Hérit., the forest dominant, and explore the
relationships between epiphyte diversity and the size
and age of the trees.
2. Methods
2.1. Nomenclature
In this study, the term oldgrowth refers to trees and/
or forest over the age of 110 years (Forests and Forest
Industry Council, 1990; Hickey, 1994; Forestry Tasmania, 1996). The term regrowth is applied to younger
trees or forests.
Species nomenclature follows that given in Jarman
and Kantvilas (2001a). Lichens encompassed within the
entity Chaenothecopsis spp. represent five taxa (C. cf.
nana, C. pusilla, C. savonica, C. tasmanica and C. sp.)
that have been grouped because of practical difficulties
in their identification in the field.
0006-3207/$ - see front matter Crown Copyright # 2003 Published by Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2003.08.001
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G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
Fig. 1. Location of the silvicultural systems trial within the Warra Long-Term Ecological Research Site in southern Tasmania.
2.2. Study area and vegetation
The Warra silvicultural systems trial comprises an
area of about 200 ha located towards the southern
boundary of the Long-Term Ecological Research Site
(Fig. 1). Environmental characteristics of the trial area
have been described by a number of authors (e.g.
Hickey and Neyland, 2000; Hickey et al., 2001; Laffan,
2001) and are summarised in Table 1.
Vegetation at the trial site has been classified by
Neyland (2001) into three types (Table 1). All plots in
the present study were located in wet sclerophyll forest
dominated by mixed-age Eucalyptus obliqua which
forms an open canopy, typically at heights of 40–65 m,
above a dense layer of understorey trees. The largest
eucalypts can exceed 250 cm diameter (Alcorn et al.,
2001; this study). The main understorey trees, up to 20
m tall, are Leptospermum lanigerum (Aiton) Smith,
Nematolepis squamea (Labill.) Paul G. Wilson, Melaleuca squarrosa Donn ex Smith and Acacia verticillata
(L’Hérit.) Willd. Other trees such as Pomaderris apetala
Labill., Leptospermum scoparium Forst. & Forst. f. and
Banksia marginata Cav. occur at some sites. The largest
of the understorey trees are generally less than 20 cm in
diameter.
The medium to low understorey vegetation (typically
1–3 m tall but sometimes taller) is dominated by the
scrambling shrub Bauera rubioides Andrews and a tall,
spreading, rosette sedge Gahnia grandis (Labill.) S.T.
Blake. These two species occur separately or intermixed
and can form a very dense, almost impenetrable layer
when they are well developed. More typically, the layer
is broken by large fallen logs or by small gaps that form
a maze of narrow passages through the tangled undergrowth. The forest is relatively open between the shrub/
sedge layer and the canopy of the understorey trees,
with the pole-like trunks of the latter having few low
branches and little foliage.
A thick blanket of eucalypt debris formed of leaves,
bark and branches covers much of the soil surface in
understorey gaps. Substrates raised above the debris
such as rocks, logs and mounds of inorganic soil are
colonised mainly by bryophytes. Ground ferns are
uncommon.
2.3. Fire history and eucalypt age
Natural broadscale regeneration of eucalypts in wet
forest depends on catastrophic disturbance, usually
caused by fire, to provide sufficient light for germination
and seedling establishment (e.g. Gilbert, 1959; Cremer,
1960; Jackson, 1968; Mount, 1979; Wells and Hickey,
1999). Dense regeneration of eucalypts usually occurs
within months of a major disturbance. However, some
regeneration possibly continues for 1 or 2 years while a
seed source, seedbed and sufficient light are available
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
361
Table 1
Summary details for the silvicultural systems trial at the Warra Long-Term Ecological Research Site (modified from Jarman and Kantvilas 2001a)
Locationa
Areaa
Elevationa
Aspecta
Soilsb
Drainageb
Climatec
Climate variables from
the nearest weather
station (Geeveston)c
Temperature
Rainfall
Vegetationd
a
b
c
d
e
43 040 S, 146 410 E, southern Tasmania
200 ha
80–240 m
Southerly
Mostly derived from Jurassic dolerite
More than half the soils are well- or moderately well-drained, about a third are imperfectly drained,
just over a tenth are poorly drained.
Temperate maritime
February max. 16.2 C
July min. 7.6 C
July max. 115 mm
February min. 56.2 mm
(a) Eucalyptus obliqua wet forest with a wet sclerophyll understorey
(b) Eucalyptus obliqua mixed forest with a thamnic rainforeste understorey
(c) Eucalyptus obliqua mixed forest with a callidendrous rainforeste understorey
Information from: Hickey and Neyland (2000).
Laffan (2001).
Packham (1995).
Neyland (2001).
Nomenclature after Jarman et al. (1994).
(J.E. Hickey, personal communication). Multi-aged
stands develop if eucalypts already present before a fire
are able to survive.
In the silvicultural systems trial, wildfires in 1934,
1914 and 1898 are responsible for most of the regrowth
trees (Alcorn et al., 2001). Fire history before 1898 is
less certain but there may have been up to six fires
between 1500 and 1900 that account for the oldgrowth
trees (Alcorn et al., 2001).
2.4. Sampling
Eucalypts were surveyed for their epiphytes on seven
plots (each of 500 m2) distributed across four coupes
(harvesting units). Each plot was subdivided into five
1010 m subplots, and the largest and smallest eucalypt
in each subplot selected for study. Not all subplots
contained two eucalypts, with a total of 55 trees being
sampled in the study. The diameter at breast height
(dbh) of each tree was measured and the trunk then
systematically searched for lichens and bryophytes up to
a height of about 2 m from the ground. Species were
collected and identified as described in Jarman and
Kantvilas (2001a).
For size/diversity analyses, eucalypts were divided
into five diameter (dbh) classes: < 25 cm, 25–49 cm, 50–
99 cm, 100–149 cm and > 150 cm.
3. Results
3.1. Composition of the epiphytic flora
Fifty-five species (30 lichens, 25 bryophytes) were
recorded from the eucalypts sampled in this study.
Despite the prominence of bryophytes on the buttress,
only a few species are common. One liverwort (Bazzania
involuta) and one moss (Rhizogonium novae-hollandiae)
account for most of the biomass, with two smaller, fineleafed mosses, Orthodontium lineare and O. pallens,
being common where charcoal scars are present. Several
other characteristic species (see below) are present in
much smaller quantities.
Species of Cladia, Cladonia, Micarea and the Caliciales (Chaenotheca, Chaenothecopsis, Microcalicium)
dominate the lichen flora. More than two-thirds of the
lichens recorded are crustose species, a group that is
generally poorly known in Tasmania and difficult to
identify. The collections made during the study require
additional taxonomic work which will almost certainly
lead to an increase in the number of species recorded.
Appendices A (lichens) and B (bryophytes) list the
eucalypt epiphytes recorded in this survey (species codes
given in Appendix C). The species are grouped into
generalists and specialists, with the latter subdivided to
show species with a distinct preference for eucalypts but
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also found occasionally on other substrates, and species
found only on eucalypts (see Jarman and Kantvilas,
2001b). It should be noted that the designation here of a
species as a generalist or specialist applies only to the
vegetation examined in this study—in other community
types across Tasmania, the same species may display a
different level of ecological specialisation.
The proportion of eucalypt specialists is high in the
lichen flora but low amongst the bryophytes. Twenty of
the 30 lichens have been found only on eucalypts in this
forest community (Jarman and Kantvilas, 2001b), and a
further four show a clear preference for them. In contrast, only five of the 25 bryophytes are confined to
eucalypts or show a preference for them (including a
preference for eucalypt wood in the form of large logs).
3.2. Phytosociology
Four main communities are discernible in the basal
trunk flora of the eucalypts. The distinction between
these is best observed on the largest trees where the differentiation of microhabitats is sharpest.
3.2.1. Bazzania involuta – Cladia aggregata community
This association is the predominant vegetation on
moist ‘mossy’ buttresses of the oldest trees. It forms a
dense, lumpy band around the base of the tree, irregular
in height but reaching to 2 m or more under suitable
conditions. The dominant species is the liverwort Bazzania involuta, but patches of the moss Rhizogonium
novae-hollandiae are usually common. Other bryophytes
present but covering little area include Lepidozia ulothrix, Lepidozia sp. C, Kurzia hippuroides and, on the
lowest levels of the buttress, Zoopsis argentea. Several
distinctive, pale-coloured macrolichens occur amongst
the bryophytes. These are mostly fruticose or squamulose species that project through the layer of bryophytes. The most prominent and common lichen species
are Cladia aggregata and species of Cladonia. The tiny
crustose lichens, Lecidea cf. botryosa and Trapeliopsis
granulosa, may overgrow the bryophytes, or occur on
the soft, spongy, moist patches of bark that occur in
gaps in the Bazzania layer. Plant cover decreases up the
trunk and the upper edge of the community is often
ill-defined as the basal species gradually diminish in
abundance.
Development of this community can be extremely
variable and, with unfavourable conditions or on small
diameter trees, it may be absent or reduced to small,
discontinuous bryophyte/lichen clumps very close to the
ground.
3.2.2. Orthodontium lineare–O. pallens community
These two moss species occur separately or intermixed
to form a fine-leafed layer with a closely cropped
appearance. It is a basal community that occurs with
the Bazzania–Cladia association but is quite distinct
from it. It is relatively pure in composition and seems to
be best developed on the charcoal surfaces that often
occur between the buttress ridges of the largest trees.
The community is absent from small diameter trees,
although occasionally a few shoots of either species may
be present with negligible cover.
3.2.3. Cladonia rigida – Micarea prasina community
This community occurs where the substrate is somewhat drier, either because it is higher on the trunk than
the Bazzania-dominated community, because the bark is
thinner (e.g. on younger trees), or because physical
effects such as the inclination of the trunk cause local
dryness. This part of the trunk can be richly colonised,
especially on larger trees, but may seem bare because of
the inconspicuous nature of the lichens. For example,
Cladonia rigida and Micarea prasina may form a continuous cover but, because of their morphology, could
be easily overlooked. Where the bark is wettest, ?Icmadophila sp. and Neophyllis melacarpa are common,
whereas marginally drier spots will support Cladia schizopora. Charred bark is frequent within this association
and provides the favoured habitat for Placynthiella
icmalea, Hypocenomyce foveata and the minute
Melaspilea sp. B. Bryophytes are virtually absent.
3.2.4. Caliciales ‘dry’ community
This community is confined to the largest trees and is
dominated by minute crustose lichens from the order
Caliciales. In the lower levels of the forest, the Caliciales
are restricted to dry, overhanging sides of the trunk or
to dry cracks and burnt-out hollows. However, further
up the trunk and above the level of the understorey, this
is the dominant community and forms a continuous
cover for many square metres. Preliminary studies of
recently felled oldgrowth trees elsewhere in the Warra
Long-Term Ecological Research Site indicate that rich,
Caliciales-dominated epiphytic communities can extend
high into the forest, often to the first major limbs on the
eucalypts at a height of about 20 m.
The dominant taxa in the community include Chaenotheca hygrophila, species of Chaenothecopsis and, on
some trees, Microcalicium disseminatum. These species
are frequently associated with other crustose lichens,
including species of Arthonia. The leprose form of
Cladia schizopora forms extensive circular thalli within
this association.
The Caliciales as a group are potentially the most
important group of lichens in the study area in terms of
ecological indicators of forest continuity and integrity.
Numerous studies in the Northern Hemisphere relating
to forest ecology consistently identify the Caliciales as
oldgrowth indicators (e.g. Tibell (1992) and Holien
(1996) in Scandinavia; Goward (1997) and Selva (1994,
1998) in North America; Rose (1976) and Rose and
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
363
lichens, one bryophyte) from a tree of 16 cm dbh. There
is a general increase in species richness with increasing
tree diameter, with the trend stronger in the lichens
(linear regression R2=0.6884, P< 0.0001, Fig. 2) than in
the bryophytes (R2=0.4043, P < 0.0001, Fig. 2). A
similar trend is apparent between age and species richness, although the data are much fewer, with the age of
only six trees being known (by cross-referencing to
individuals in a study by Alcorn et al., 2001). The diameter, estimated age and species richness for the six
trees are shown in Table 2 (also see Fig. 2).
4. Discussion
4.1. Eucalyptus obliqua as a substrate
Fig. 2. The relationship between species richness and diameter (dbh)
in the epiphytic flora of Eucalyptus obliqua. A. Lichens (linear regression: R2=0.6884; P <0.0001). B. Bryophytes (linear regression:
R2=0.4043; P <0.0001). Encircled dots represent trees for which ages
are known.
Table 2
Age–diameter–species richness relationships for six trees in the silvicultural systems trial
Tree
Dbh (cm)
Estimated age
(years)a
No. of cryptogam
species
T1S
T2B
T3B
T4B
T1B
T5B
25
59
63
69
118
122
84
101
100
128
308
316
8
17
13
16
21
29
a
Unpublished data, Forestry Tasmania.
Wolseley (1984) in the United Kingdom). Indeed, several taxa involved in those studies have been recorded in
our work in the silvicultural systems trial.
3.3. Diameter–age–richness relationships
Species richness and diameter for each of the sampled
trees are shown in Appendices A (lichens) and B (bryophytes). The highest number of epiphytes recorded on a
single tree was 29 (17 lichens, 12 bryophytes) from a
trunk of 122 cm dbh. The lowest number was three (two
Eucalyptus obliqua supports a rather meagre epiphytic
flora when compared to other Australian wet forest tree
species for which data are available. In the present
study, 17 lichens and 12 bryophytes were recorded from
the richest eucalypt tree and, although only the buttress
was studied, the figures seem rather indicative of
the total tree flora as seen in a separate study nearby
(outside the silvicultural systems trial). In that study
(Jarman and Kantvilas unpublished data), the richest of
four, large, felled E.obliqua trees yielded 18 lichen species and 13 bryophyte species in an inventory incorporating the buttress and most of the trunk. All four felled
trees were in wet forest (i.e. in conditions considered
favourable for cryptogams in Tasmania) but none
approached the richness reported for rainforest host
trees. For example, a Huon pine tree (Lagarostrobos
franklinii (Hook.f.) Quinn) in rainforest in western Tasmania yielded 55 lichens and 75 bryophytes (Jarman
and Kantvilas, 1995a), and Milne and Louwhoff (1999)
recorded 36 lichens and 28 bryophytes from a windthrown Nothofagus cunninghamii (Hook.) Oersted in
Victoria.
In spite of the relatively small suite of species present,
Eucalyptus obliqua is an important substrate because of
its very distinctive epiphytic flora (Jarman and Kantvilas, 2001b). This is manifest in the lichens by a very high
proportion of eucalypt specialists, resulting in few species shared with other host trees or habitats in the forest. In the bryophyte flora, there are only a few eucalypt
specialists. However, three of these and one other more
widespread species overwhelmingly dominate the bryophyte flora of eucalypts. At the same time, many of the
bryophyte generalists in the forest seem to display an
aversion for eucalypts, so the composition and appearance of the eucalypt flora shows little similarity to that
of other habitats in the forest.
The distinctive character of the eucalypt flora is most
likely to be a response to some specific (but hitherto
unstudied) chemical or physical properties of eucalypt
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bark and wood. The massive size of the trunks also
contributes to producing a range of substrates and
microhabitats found nowhere else in the forest.
4.2. Succession
Succession in epiphytic communities typically
involves a gradual replacement of species, from those
regarded as pioneers to those that characterise climax
associations. It also frequently involves vertical shifts in
species distributions as epiphytes that favour young
bark are increasingly restricted to the upper, younger
parts of trees. This type of succession is exemplified by
the lichen flora on the Tasmanian rainforest tree,
Nothofagus cunninghamii (Kantvilas, 1990), where
increasing girth generally means a turnover of taxa, with
pioneers gradually being replaced as the trunk ages, or
progressively displaced up the tree to the new portions
of young branches and twigs.
Floristic changes associated with increasing girth in E.
obliqua do not follow the replacement sequences that
have been observed generally for species such as Nothofagus (Kantvilas, 1988, 1990). For example, species on
the small eucalypts are also found on the large ones
(Fig. 3) (as well as an additional suite of species on the
latter). Thus, the flora of small trees is dominated by
pioneer lichens such as Micarea prasina, Cladonia rigida
and Cladia schizopora, and bryophytes such as Rhizogonium novae-hollandiae, Bazzania involuta, Kurzia hippuroides and Zoopsis argentea. All of these species also
flourish on the largest trees, along with ‘old trunk colonisers’, including species of the Caliciales.
The displacement upwards of pioneer species on large
eucalypts is not in evidence either. From our cursory
observations of whole mature trees, it appears that no
basal species are found in the canopy. Small thalli of
crustose lichens from genera such as Ramboldia, Caloplaca, Lecanora s.lat. and Buellia may occur on dead
twigs but, overall, canopy limbs of eucalypts are very
poorly colonised.
4.3. Old trees, ancient forests and epiphytes
The relationship between the diversity of cryptogamic
epiphytes and forest age is well-recognised and supported by an extensive body of literature from various
parts of the Northern Hemisphere: for example, Rose
(1976, 1992), Rose and James (1974) and Rose and
Wolseley (1984) in temperate woodlands of Britain;
Goward (1993), Rosentretter (1995), McCune et al.
(2000), Sillet et al. (2000) and Cooper-Ellis (1998) in
North American forests; and Kuusinen and Siitonen
(1998), Tibell (1992) and Hyvärinen et al. (1992) in
Scandinavia. All authors consistently report strong differences in the composition of epiphytic floras between
oldgrowth or ancient forests, and younger stands. Key
forest factors that influence epiphyte diversity are the
ecological continuity in time of the forest cover, especially insofar as it affects microclimate and epiphyte
dispersal and establishment; structural diversity which
provides habitat diversity for epiphytes; and the presence of old trees, which also provide microhabitat
diversity (Gustaffson et al., 1992; Rose, 1976; Neitlich
and McCune, 1997). On this basis, regional assemblages
of cryptogams have been defined which serve as oldgrowth indicators (e.g. see Coppins and Coppins, 2002),
and these notably include, amongst other species, cyanolichens and members of the Order Caliciales
(Goward, 1994; Selva, 1994; Rose, 1992). Two types of
old forest indicator species were identified by Kuusinen
and Siitonen (1998): those that prefer old trees (e.g.
members of the Caliciales) and those that require
specific habitats that develop in very old stands (e.g.
cyanolichens).
There is insufficient information available at present
to classify the epiphytes of oldgrowth E. obliqua in
terms of indicators of forest continuity or age. Such an
outcome depends on very extensive ecological and floristic studies in eucalypt forest of various ages; in this
regard, work in Tasmania is still in its infancy. However, the results suggest that there are late-successional
species in forest that contains oldgrowth trees and such
species would be worthwhile investigating further for
their potential as oldgrowth indicators (Table 3). Some
of these species were recorded occasionally on small
diameter trees (see Appendices A and B) but, in view of
the variability demonstrated in age/diameter relationships by Alcorn et al. (2001) in the silvicultural systems
trial area, it is quite possible that these small trees are
actually old suppressed ones. If this were found to be
the case, then epiphytes may well provide insights into
forest development in cases where tree size or forest
structure is ambiguous.
Among the potential oldgrowth indicators found in E.
obliqua forest (Table 3), potential old tree indicators
(e.g. Caliciales taxa) are well represented but old forest
indicators (e.g. cyanolichens) are particularly scarce. In
comparison, Tasmania’s cool temperate rainforest, a
climax vegetation type, supports both types of species
(e.g. see Jarman and Kantvilas, 1995b), including some
that are either identical or closely related to old forest
indicator species in Northern Hemisphere forests (see
Kantvilas, 1988). The scarcity of old forest indicators in
the study area is not surprising. Because of the regeneration strategy of the dominant tree, wet eucalypt forest does not attain the status of an ancient forest in the
sense of one without disruption to the forest cover over
many hundreds of years and over more than one generation of a long-lived forest dominant. Ecological continuity is limited, at its longest, to the lifespan of the
oldest trees (350–450 years) and is often much less. The
paradox is that at the scale of a local stand, the enormous
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
365
Fig. 3. The occurrence (% frequency) of lichens and bryophytes on trees in five diameter classes, excluding rare species (those occurring on less than
five trees). Species codes are given in Appendix C.
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Table 3
Potential oldgrowth indicators in Eucalyptus obliqua forest
Lichens
Arthonia sp. B
Chaenotheca hygrophila
Chaenothecopsis spp.
Cladonia cf. murrayia
Cladonia ustulataa
Cladonia weymouthiia
Hypocenomyce scalaris
?Icmadophila sp.
Lecide cf. botryosa
Melaspilea sp. Bb
Micarea cf. adnata
Micarea cf. mutabilis
Micarea prasina aggr. form C
Microcalicium disseminatum
Placynthiella icmaleaa,b
Species C
Trapeliopsis granulosaa
Bryophytes
Orthodontium lineareb
Orthodontium pallensb
Lepidozia sp. C
Riccardia crassa (as an epiphyte)c
a
These lichens occur in other vegetation types in Tasmania where
they are not epiphytes. Their occurrence at the site as epiphytes rather
than ground colonisers may be indicative of oldgrowth conditions.
b
These species show an association with charcoal on the trunks.
c
Although this liverwort occurs on non-epiphytic habitats in the
forest, it is epiphytic only on large eucalypts.
eucalypt trees are fireweeds (cf. Cremer, 1960) and their
long-term presence in a particular location hinges on
their periodic catastrophic destruction.
4.3.1. Old trees
Eucalypts can be massive (over 90 m tall; e.g. see
Hickey et al., 2000) and potentially over four hundred
years old. Most eucalypt species, including E. obliqua,
have an ability to survive fires and may be burnt several
times during their lifetime. Each time, the surface of the
tree may be partially or completely ‘wiped clean’ of its
epiphytic flora. Similarly, the surrounding vegetation
may be partially or totally destroyed, depending on the
intensity of the fire. As a result, the age of an oldgrowth
dominant is not necessarily indicative of the age of its
epiphytic flora nor that of the standing forest around it.
The question then arises of whether the epiphytic flora
of these oldgrowth trees has any special characteristics
that can be attributed to the great age and size of the
host (oldgrowth indicators) or whether the flora is
young and more in keeping with one originating from
the most recent fire.
In the forest examined, there are obvious floristic differences between large diameter and small diameter
trees, suggesting that size or age (or a related factor)
does confer some special characteristics on the epiphytic
flora. Not only is species richness higher on the larger
trees, but 11 of the 16 eucalypt specialists in the lichen
flora were recorded only on trees greater than 80 cm
diameter.
Various reasons may account for the floristic differences seen on large and small trees. Increasing girth is
accompanied by an increase in surface area and this
probably accounts for many of the additional bryophytes found on the larger trees. However, for lichens
and perhaps a few of the bryophytes, the increase in
species richness with diameter is attributed mainly to an
associated increase in habitat diversity, in much the
same way that landscape variability is associated with
increased species richness in vascular plants. As the trees
become larger, they develop moisture gradients around
the trunk, moisture and light gradients up the trunk,
and other variations such as hollows, fluting, charcoal
and dead, bleached lignum. Dry furrows and crevices in
the bark provide particularly specialised niches colonised chiefly by lichens. In addition, because of their
very large size and variable shape, larger diameter trees
are more likely than smaller ones to experience some
variation in fire intensity around their trunk. It may be
that some patches of bark on the trunk escaped burning
in the last fire or even the one before that, enabling
oldgrowth colonisers to establish/survive alongside the
younger flora in the more recently burnt patches.
The relative contribution of tree age (versus tree size)
to epiphytic diversity is not clear. Very large trees are
obviously old, but suppression, genetic variability and
site factors result in considerable variation in girth to
the extent that age classes of the smaller diameter trees
remain equivocal in the absence of actual age measurements (e.g. see Alcorn et al., 2001). A better understanding of the relationship between age alone and
epiphyte richness in Eucalyptus obliqua might be gained
by comparing trees of similar diameter but different age
(i.e. old suppressed trees and younger unsuppressed
trees). It has been demonstrated in Northern Hemisphere studies that age, in so far as it leads to changes
in the physical or chemical properties of the bark, can
have an important effect on epiphyte colonisation (e.g.
Barkman, 1958; Wolseley and O’Dare, 1990; Hyvärinen
et al., 1992).
The results from this study indicate that large oldgrowth eucalypts are important for cryptogamic diversity even though they occur in a landscape that has
experienced catastrophic disturbance in the recent past
(in this case, about 70 years ago). Almost certainly, their
epiphytic flora would contribute towards recolonisation
of young eucalypts around them, once the latter develop
the appropriate microhabitats and environmental conditions are suitable. This interaction between young and
old trees was shown by Sillet and Goslin (1999) to occur
in multiple-aged Douglas fir forest in the Northern
Hemisphere.
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
4.4. Management implications
4.4.1. Dispersal and recolonisation
Successful recolonisation by cryptogams after a major
ecological disruption depends on several factors,
including a source of propagules, the dispersal ability of
the species and the development of a suitable substrate
in a suitable environment. At present, all of the coupes
(harvesting units) in the silvicultural systems trial are
surrounded by uncut forest similar to that previously
occurring in the coupes. Although little detailed sampling has been undertaken in these forests, it is likely
that a very high percentage of cryptogamic species
found in the logged coupes presently occurs in the surrounding vegetation. Thus, a potential source of propagules for the new forest exists in the immediate vicinity
of the coupes.
The dispersal abilities of the species involved are
unknown. Circumstantial evidence suggests that at least
some species are able to spread very quickly, but no
direct studies on this aspect of cryptogam ecology have
been undertaken in Tasmania. To have been recorded at
all, the species in this study have demonstrated an ability to establish within about 70 years of a wildfire
(assuming all plots were burnt by the most recent wildfire in 1934), although the severity and uniformity of
that fire are unknown. What is also unclear is how many
species previously present have not managed to recolonise the forest within 70 years. It is possible that only
the most successful recolonisers have been recorded and
others with poorer dispersal abilities or narrower habitat
requirements need longer times to establish.
Regardless of dispersal ability, colonisation by cryptogams is unlikely to take place until suitable habitats
establish within the regrowth forest. This involves the
whole forest environment, including microclimatic conditions such as shade, moisture and temperature, and
the presence of appropriate surfaces for colonisation in
suitable microhabitats. Given the difference in environmental conditions between the pre- and post-logged
areas, suitable habitats are likely to take many years to
develop. This is clearly the case for epiphytes that will
only colonise mature trees or trees that have reached a
certain girth (see below). If, as part of landscape planning in the area, the adjacent forest is targeted for logging when the young trees reach a height of five metres
(Forest Practices Board, 2000), then the adjacent source
of propagules could be removed before suitable habitats
have had time to redevelop.
4.4.2. Rotation times
The routine silvicultural regime prescribed for lowland wet eucalypt forests in Tasmania is based on a
clearfell, burn and sow system, with a rotation time of
80–100 years (Forestry Tasmania, 1998). At Warra, this
rotation time is likely to result in smaller, more uniform
367
trees and a narrower range of sizes compared to those
seen in old, natural stands. This is indicated by the work
of Alcorn et al. (2001) which shows that the growth rate
of eucalypts at the Warra silvicultural systems trial is
quite slow. For example, the largest of 19 trees aged
between 80 and 105 years (the approximate length of the
rotation period) had a diameter of 83 cm dbh; all others
were 70 cm or less, many considerably less.
Results from the present study indicate that the longterm effects of current prescriptions on forest biodiversity, specifically on the epiphytic flora of the eucalypts, are likely to be severe because the regime, if
unmodified, does not allow for the replacement of very
large, very old trees. It can be seen from Appendix A
that 11 of the 16 eucalypt specialists (lichens) were
found only on trees with a diameter of more than 80 cm,
and nine of these were found only on trees greater than
100 cm dbh. On present information, the needs of these
species are unlikely to be met by regrowth trees.
It might well be argued that with improved technology, forest managers could produce faster growing trees
that attain larger diameters within the prescribed rotation time. However, our results suggest that it is not size
as such that is the major driver of species diversity, at
least with respect to the lichens, but rather the increase
in habitat diversity that develops on the trunk as the
tree ages. Analagous age/size-related changes to eucalypts have been found to be important for a range of
biota, including, for example, arboreal marsupials
(Smith and Lindenmayer, 1988; Lindenmayer et al.,
1991), bats (Taylor and Savva, 1988) and birds (Saunders et al., 1982; Nelson and Morris, 1994). In accord
with findings from these animal studies, we would
expect that rotation times longer than those prescribed
routinely, and retention within the coupe of some large
eucalypts, especially deformed ones with maximum
substrate variability on the trunk, would contribute significantly to maintaining cryptogamic diversity in production forests. Similar conclusions have been drawn
from cryptogamic studies in the Northern Hemisphere
(e.g. Kuusinen and Siitonen, 1998; Sillet and Goslin,
1999; Sillet et al., 2000) where loss of old trees is seen as
contributing to a reduction in biodiversity.
4.5. Impact of large trees on non-epiphytic habitats
utilised by cryptogams
As well as their value as live habitat, there are other
benefits of retaining large eucalypts in the coupes. The
trees provide a source of logs for the forest floor, and
large decaying logs have been shown to have high
conservation value in Northern Hemisphere studies
(e.g. Söderström, 1988; Gustafsson and Hallingbäck,
1988; Andersson and Hytteborn, 1991; Hallingbäck,
1994). They are expected to be similarly important in
Tasmania.
368
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
In addition, large trees, when windthrown, can contribute towards providing a special habitat for cryptogams that grow only on inorganic soil. Specialist species
of this nature colonise the freshly exposed soil in and
around the hollows left by the upturned tree; others
colonise the soil-covered root mass which is raised well
above the moist, shaded conditions of the forest floor
(see Jarman and Kantvilas, 2001b), enabling species
that would be outcompeted at ground level to survive.
The larger the tree, the more effective it is in creating the
habitat. In some cases, the face of inorganic soil may
rise 3 m or more above ground level.
If eucalypts remain standing after death, they provide
an important habitat for colonisers of brightly lit, dry
wood (certain lichen species). This habitat occurs naturally in undisturbed forests and in forests logged in past
times when very high stumps were left after trees were
felled. Such habitats are likely to be scarce in Tasmania
in landscapes where there are large areas of even-aged
regrowth arising from clearfelling. One of the many
challenges for forest management and silviculture is to
be able to maintain such oldgrowth values in production forests in order to ensure the conservation of particular lichen and bryophyte species.
Acknowledgements
We thank H.J. Elliott, M.J. Brown, J.E. Hickey and
M.G. Neyland (Forestry Tasmania) for critical reading
of the manuscript, and R. Taylor (formerly Forestry
Tasmania) for his interest and support for the cryptogamic study since its inception in 1997. Work by GK at the
Tasmanian Herbarium was supported by funding from
Forestry Tasmania which is gratefully acknowledged.
APPENDIX ON NEXT PAGE
369
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
Appendix A. Lichen flora of 55 trees, in order of increasing tree diameter. Dotted lines indicate inferred maximum
diameter of trees at 80 years and 100 years, harvested on a 80–100 year rotation. (Species codes are given in
Appendix C)
+
+
+
+
+
+
+
+
+
+
+
+
+ +
+ +
+
+
+
+ +
+ +
Plac ic
+
+
+
+ +
+
+
+
+ +
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+ +
+
+
+
+
+
+
+
+
+
+ +
+
+
+ +
+
+
Trap gr
?Icmad
Sp C
Lec bot
Clado m
Chaen hy
Chaen spp
Mic pr C
Melasp
Microcal
Hypocen
Clado we
Mic cf. m
Clado ust
105
111
118
120
122
133
140
146
147
150
150
155
180
200
230
255
+
+
Arth sp. B
4B
5B
1B
T8
5B
2B
2B
5B
2B
5B
2B
1B
4B
2B
5B
3B
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Confined
to eucalypts
Cladia sc
Bn418
Bn418
T
C
T
C
Bn418
M
B518
B519
B106
M
M
B372
B372
Bn418
+
Neo mel
86
96
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Clado sub
3B
2S
+
+
+
+
+
Mic cf. A
C
C
Mic pr A
7
10
12
12
14
14
15
16
17
19
20
20
22
25
25
27
33
35
40
40
40
40
40
46
48
50
54
55
58
58
59
60
63
63
66
69
78
Clado rig
4S
3S
5S
2S
4S
3S
4S
2S
4S
5S
1S
5S
4B
4S
1S
2S
5S
3S
1S
3S
4S
4B
3S
3B
1B
5S
3B
2B
1B
3S
2B
3B
T7
3B
5B
4B
3B
Prefer
eucalypts
Cladia ag
B518
B519
M
T
M
T
T
B518
B372
B373
M
T
B372
C
T
M
C
M
Bn418
Bn418
Bn418
B518
B106
B106
B106
Bn418
B518
M
Bn418
B372
T
B372
C
T
C
T
M
Various
habitats
Clado ra
Dbh
(cm)
Mic pr D
Tree
no.
Arth sp. A
Plot
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+ 80 cm dbh
+
No.
of
spp.
per
tree
4
4
3
3
3
4
5
2
4
4
5
4
4
3
3
2
4
4
2
3
3
3
6
3
6
5
3
4
5
6
5
6
5
7
7
7
7
——————————————————————————————————
+ + +
+ + +
+ +
+ +
+
100 cm dbh
+
+ + +
+
9
7
+
5
8
14
8
17
15
9
14
6
8
12
12
9
11
15
14
——————————————————————————————————
Frequency (n=55)
+
+
+
+
+ + +
+ + +
+
+
+
+ +
+
+
+ +
+ +
+
+
1
4
5
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+ +
+
+
+ +
+
+ +
+
+ +
+
+ + +
+
+
+ + +
+
+ + +
+ +
+
+
+ +
+
+ + +
+
+ + +
+
+ +
+
+ + +
+
+ + +
+
+
+ + +
31 54 54
2
9
36 49
1
1
1
2
+
+ +
+
+
+
+ +
+ +
+ +
+
+
+
+
+ +
+
2
2
2
6
7
+
+
+
+ + + +
+
+
+ + + + + +
+
+
+
+
+
+ + +
+
+
+
+ +
+
+ +
+
+
+
+ +
+ + + + +
+ + + + + +
7
7
9
+
+
+
+
+
+
+
+
+
+
+
+
+
10 13 15 21
370
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
Appendix B. Bryophyte flora of 55 trees, in order of increasing tree diameter. Dotted lines indicate inferred maximum
diameter of trees at 80 years and 100 years, harvested on a 80–100 year rotation. (Species codes are given in
Appendix C)
+ +
+ +
+
+
+ +
+ + + + +
+
+ + +
+ + +
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+ +
+
+
+ + +
+
+
+ +
+
+
+ + +
+
+
+
+
+
+ +
+
+ + +
+
+
+ + +
+ +
+
+
80 cm dbh
+ + +
+ + +
9
8
+
+
+
+
+
+
+ + + +
+ +
+ +
+
+ +
+ +
+ +
+ +
+ +
+
+ +
+ +
+
+ +
+ +
+
+ +
+ +
+
+ +
+ +
+
+ +
+ +
+
+
+ +
+ +
+ +
+ +
+
+ +
+
+ +
+ +
+ +
+ +
+
+ +
+
+
+
+
+
+
+
+
+
+
+
+
+
No.
of
spp.
per
tree
2
4
2
3
6
5
4
1
7
4
3
5
5
3
5
3
5
8
2
6
5
3
11
7
4
4
3
7
6
6
8
4
3
10
8
9
10
+
+
Rhiz no
Orth lin
Orth pal
Lep C
Baz m
Zoo arg
Kur hip
Baz inv
Tel her
Tel pat
Lep ulo
Ric coc
Ric cra
Acr col
105
111
118
120
122
133
140
146
147
150
150
155
180
200
230
255
Tel tas
4B
5B
1B
T8
5B
2B
2B
5B
2B
5B
2B
1B
4B
2B
5B
3B
Lep W
Bn418
Bn418
T
C
T
C
Bn418
M
B518
B519
B106
M
M
B372
B372
Bn418
Lep pro
86
96
Kur sex
3B
2S
Podom
C
C
Wijkia
7
10
12
12
14
14
15
16
17
19
20
20
22
25
25
27
33
35
40
40
40
40
40
46
48
50
54
55
58
58
59
60
63
63
66
69
78
Leucob
4S
3S
5S
2S
4S
3S
4S
2S
4S
5S
1S
5S
4B
4S
1S
2S
5S
3S
1S
3S
4S
4B
3S
3B
1B
5S
3B
2B
1B
3S
2B
3B
T7
3B
5B
4B
3B
Confined to
or prefer
eucalypts
Dicr bil
B518
B519
M
T
M
T
T
B518
B372
B373
M
T
B372
C
T
M
C
M
Bn418
Bn418
Bn418
B518
B106
B106
B106
Bn418
B518
M
Bn418
B372
T
B372
C
T
C
T
M
Various
habitats
?Ceph
Dbh
(cm)
Metzg
Tree
no.
Dicr set
Plot
+ +
+ +
+
+ + +
——————————————————————————————————
+
+
100 cm dbh
+
+ +
+ + +
+ + +
+
+ +
+ + + +
+ +
+
+
+
+
+
+ +
+
+ +
+ +
+ +
+
+ +
——————————————————————————————————
Frequency (n=55)
+
+
+
+
+
+
+
+
+ +
+
+ + + +
+ +
+
+ +
+ +
+
+
+
+
+
+
+
1
1
1
2
2
2
2
2
2
2
3
4
7
+
+
+
+
+
+
+ + + 5
+
+ + + 6
+
+ + + + 7
+
+ + + 9
+ + +
+ + 12
+
+ + + 10
+
+ + + + 8
+
+ + + + 10
+
+ + + 4
+
+ +
+ 6
+
+ + + + 9
+
+ + + + 9
+
+ + + + 9
+
+ + + + 9
+
+ + + + 12
+
+ + + + 9
11 11 15 19 30 44 53
1
15 29 31 52
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
371
Appendix C. Species codes for Fig. 3, and Appendices A (lichens) and B (bryophytes)
Species code
Lichens
Arth sp. A
Arth sp. B
Chaen spp
Species
Chaen hy
Cladia ag
Cladia sc
Clado m
Clado ra
Clado rig
Clado sub
Clado ust
Clado we
Hypocen
?Icmad
Lec bot
Melasp
Mic cf. a
Mic cf. m
Mic pr A
Mic pr C
Mic pr D
Microcal
Neo mel
Plac ic
Sp C
Trap gr
Arthonia sp. A
Arthonia sp. B
5 species: Chaenothecopsis cf. nana Tibell, C. pusilla (Ach.) A.F.W. Schmidt, C. savonica (Räsänen) Tibell,
C. tasmanica Tibell, C. sp.
Chaenotheca hygrophila Tibell
Cladia aggregata (Sw.) Nyl.
Cladia schizopora (Nyl.) Nyl.
Cladonia cf. murrayi W. Martin
Cladonia ramulosa (With.) Laundon
Cladonia rigida (Hook.f. & Taylor) Hampe var. rigida
Cladonia subsubulata Nyl.
Cladonia ustulata (Hook.f. & Taylor) Leighton
Cladonia weymouthii F. Wilson ex A.W. Archer
Hypocenomyce scalaris (Ach.) M. Choisy
?Icmadophila sp.
Lecidea cf. botryosa (Fr.) Th. Fr.
Melaspilea sp. B
Micarea cf. adnata Coppins
Micarea cf. mutabilis Coppins & Kantvilas
Micarea prasina Fr. aggr. form A
Micarea prasina Fr. aggr. form C
Micarea prasina Fr. aggr. form D
Microcalicium disseminatum (Ach.) Vainio
Neophyllis melacarpa (F. Wilson) F. Wilson
Placynthiella icmalea (Ach.) Coppins & P. James
Species C
Trapeliopsis granulosa (Hoffm.) Lumbsch
Bryophytes
Acr col
Baz inv
Baz m
?Ceph
Dicr bil
Dicr set
Kur hip
Kur sex
Lep C
Lep pro
Lep ulo
Lep W
Leucob
Metzg
Orth lin
Orth pal
Podom
Rhiz no
Ric coc
Ric cra
Tel her
Tel pat
Tel tas
Wijkia
Zoo arg
Acromastigum colensoanum (Lehm. & Lindenb.) Evans
Bazzania involuta (Mont.) Trevis.
Bazzania monilinervis (Lehm. & Lindenb.) Trevis.
?Cephaloziella sp. A
Dicranoloma billardierei (Brid.) Paris
Dicranoloma setosum (Hook.f. & Wilson) Paris
Kurzia hippuroides (Hook.f. & Taylor) Grolle
Kurzia sexfida (Steph.) Grolle
Lepdozia sp. C
Lepidozia procera Mitt.
Lepidozia ulothrix (Schwaegr.) Lindenb.
Lepidozia sp. W
Leucobryum candidum (P. Beauv.) Wilson
Metzgeria sp.
Orthodontium lineare Schwaegr.
Orthodontium pallens (Hook.f. & Wilson) Broth.
Podomitrium phyllanthus (Hook.) Mitt.
Rhizogonium novae-hollandiae (Brid.) Brid.
Riccardia cochleata (Hook.f. & Taylor) Kuntze
Riccardia crassa (Schwaegr.) Carring. & Pears.
Telaranea herzogii (E.A. Hodgs.) E.A. Hodgs.
Telaranea patentissima (Hook.f. & Taylor) Hodgs.
Telaranea tasmanica (Steph.) Engel & Merrill
Wijkia extenuata (Brid.) Crum
Zoopsis argentea (Hook.f. & Taylor) Hook.f. ex Gottsche et al.
372
G. Kantvilas, S.J. Jarman / Biological Conservation 117 (2004) 359–373
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