changes in plant species and functional composition with time since fire in two mediterranean...

Journal of Vegetation Science && (2012)

Changes in plant species and functional compositionwith time since fire in twomediterranean climate plantcommunities

Carl R. Gosper, Colin J. Yates & Suzanne M. Prober

Keywords

Fire-return interval; Mallee; Obligate seeder;

Plant functional type; Resprouter; Seed bank;

Senescence; Serotinous; South-western

Australia

Nomenclature

Western Australian Herbarium (1998–2011)

Received 27 October 2011

Accepted 18 April 2012

Co-ordinating Editor: Juli Pausas

Gosper, C.R. (corresponding author,

[email protected]) & Yates, C.J.

([email protected]): Science Division,

Department of Environment and Conservation,

Locked Bag 104, Bentley Delivery Centre,

Kensington, WA, 6983, Australia

Gosper, C.R. & Prober, S.M. (suzanne.

[email protected]): CSIRO Ecosystem Sciences,

Private Bag 5, Wembley, WA, 6913, Australia

Abstract

Question: Do floristic composition and plant functional type (PFT) richness and

dominance change with time since fire, in the directions predicted through

consideration of their fire response traits?

Location: Two vegetation communities in the globally significant biodiversity

hotspot of south-western Australia: mallee, dominated by resprouters, and mal-

lee-heath, dominated by non-resprouters.

Methods: Species richness and cover were sampled in replicated plots across a

time since fire gradient ranging from 2 to >55 yr post-fire, using a space-for-time

approach. Species were allocated to PFTs according to traits relevant to the pro-

cesses of vegetation change underpinning the initial floristic compositionmodel of

vegetation assembly: their capacity to resprout, the location and persistence of the

seed bank, competitive stratum and longevity. Ordination and ANOVAwere used

to test for differences in floristic and PFT composition between young (<10 yr

post-fire),mature (18–35 yr) and old (>40 yr) vegetation in each community.

Results: Plant functional type and floristic analyses were similar, showing sub-

stantial changes in the composition of mallee-heath vegetation with time since

fire, but not in mallee. The direction of change in PFT composition in mallee-

heathwas consistent with predictions, with increasing cover of non-resprouting,

serotinous PFTs, an intermediate peak in cover of PFTs with persistent soil-

stored seed banks, and decreasing cover of post-fire ephemerals and non-respro-

uting, non-serotinous dwarf shrubs, herbs and graminoids with increasing time

since fire. Success in predicting changes in PFT dominance in mallee was lower.

Conclusions: The similarity of floristic and PFT analyses suggest that these

approaches are interchangeable for characterizing vegetation change with

increasing time since fire. PFTs were more effective for predicting fire response

trajectories in the vegetation community dominated by non-resprouting, seroti-

nous shrubs (mallee-heath) than that dominated by resprouting, serotinous

trees (mallee). The underlying vegetation assembly model and PFTs used appear

suitable for broader application in fire-prone communities with competitive

dominance by non-resprouting, serotinous shrubs, but less so in communities

dominated by other PFTs.

Introduction

Fire is a perturbation that shapes vegetation patterns

and plant community composition in seasonally dry

landscapes worldwide (Bond & van Wilgen 1996; Bond

et al. 2005; Verdu & Pausas 2007). Fires consume bio-

mass and promote plants with functional traits that

enable survival, recruitment and/or reproduction during

and shortly after fire.

Plant functional types (PFTs) are groupings of plant taxa

that share particular functional traits. Whilst the traits used

in such classifications necessarily vary depending on the

purpose of the study and the mechanisms through which

responses occur (Noble & Gitay 1996), the PFT approach

Journal of Vegetation ScienceDoi: 10.1111/j.1654-1103.2012.01434.x© 2012 International Association for Vegetation Science 1

has been widely used in interpreting plant community

changes in response to a variety of environmental pertur-

bations, including fire (Noble & Slatyer 1980; McIntyre

et al. 1995; Pausas et al. 2004; Keith et al. 2007). There is

growing evidence from empirical studies that PFT

approaches can effectively predict and generalize vegeta-

tion changes under specific fire regimes (Pausas et al.

2004; Keith et al. 2007), and hence support decisions on

the timing and type of management interventions for bio-

diversity conservation.

The application of a deductive PFT approach to under-

standing community changes after disturbance is under-

pinned by two key components: (1) the identification of

an appropriate underlying vegetation assembly model that

supports interpretation of vegetation changes; and (2) the

identification of traits relevant to the processes of vegeta-

tion change in the adopted vegetation assembly model

(Keith et al. 2007; Keith 2012).

The ‘initial floristic composition’ vegetation assembly

model (Egler 1954) proposes that changes in plant commu-

nities after fire are driven by two main processes. First, all

plant species present during the succession re-establish

shortly after fire, and dispersal of new species into the area

and their establishment during the inter-fire period is negli-

gible. Second, changes in community composition with

time since fire reflect the differential growth rates, competi-

tiveness and longevities of component species (Collins et al.

1995; Capitanio & Carcaillet 2008). Support for this model

of vegetation assembly in fire-prone communities has

accrued through numerous studies showing peak richness

of above-ground plants immediately after fire, combined

with no evidence over the inter-fire period for the whole-

sale community change indicative of concordance with

alternative vegetation assembly models (Russell & Parsons

1978; Grace & Keeley 2006; Gosper et al. 2012).

How species respond to time since fire in terms of the

processes that underpin the initial floristic composition

model, such as growth rate, competiveness and longevity,

will be determined by their functional traits. Using a

deductive approach (Pausas & Lavorel 2003; Keith 2012),

the critical functional traits affecting post-fire community

assembly and recovery under the initial floristic composi-

tion model include: the degree of individual persistence

(resprouting or non-resprouting); propagule persistence

and storage type (persistent in the canopy – serotiny, per-

sistent in the soil or transient seed bank); plant height

(reflecting a competitive hierarchy during the inter-fire

period based on vertical stature); and individual longevity

(Pausas & Lavorel 2003; Keith et al. 2007; Table 1). The

PFT approach predicts that species with particular combi-

nations of the above traits will respond in a similar and pre-

dictable way to time since fire or a particular fire regime.

So, for example, the richness and cover of long-lived, res-

prouting, serotinous, upper stratum species might be

expected to remain stable with time since fire, compared to

that of long-lived, non-resprouting, persistent soil seed

bank, lower stratum species, which would be predicted to

decline with time since fire.

To better establish the predictive capacity and generality

of the PFT approach underpinned by the initial floristic

composition model, we evaluated it in two contrasting veg-

etation types, mallee and mallee-heath, in the globally sig-

nificant biodiversity hotspot of south-western Australia.

Mallee andmallee-heath are prominent and diverse vegeta-

tion types in this mediterranean climate region, that occur

in amosaic across topographically subdued, fire-prone land-

scapes (Beard 1990). The two communities have competi-

tive dominance by different PFTs, with mallee-heath being

dominated by non-resprouting, serotinous shrubs and mal-

lee by resprouting, serotinous trees (Gosper et al. 2010; Par-

sons & Gosper 2011). Previous detailed analyses have

demonstrated the effectiveness of the PFT approach in fire-

prone shrublands dominated by non-resprouting, seroti-

nous shrubs (Keith et al. 2007), hence its relative success in

predicting the vegetation dynamics of fire-prone communi-

ties dominated by other PFTs is of particular interest.

Additionally, we tested whether a PFT approach suffi-

ciently captures time since fire responses in our study com-

munities, or whether floristic compositional data provide

additional insights relevant to management. Use of PFTs

allows for prediction and generalization (Keith et al.

2007), but few studies have compared analyses of func-

tional vs species composition with respect to their value for

interpreting vegetation change at the local level.

Table 1. Key traits (numbered, grey shading) that combine to form plant

functional types (PFTs) (RStree etc.) in fire-prone mediterranean climate

vegetation. For each combination of the traits of longevity, individual per-

sistence and seed bank type there are three potential placements in the

competitive hierarchy based on (4. Plant Height): upper (tree), mid-stratum

(shrub) and ground-layer (low).

1. Longevity 2. Individual

Persistence

3. Seed Bank Persistence and Type

Persistent

Canopy

Persistent

Soil

Transient

Soil

Long (>6 yr) Resprouter RStree [ RNtreeA ]

RSshrub [ RNshrubA ]

– [ RNlowA ]

Non-resprouter NStree NNtree –

NSshrub NNshrub –

– NNlow –

Short (� 6 yr) Resprouter –[ Ephem

B]

Non-resprouter –

– = no representatives known in our communities. See Table 2 for

expanded PFT codes. For the purposes of our analyses, these PFTs were

merged across persistent and transient seed bank types (A) or seed bank

type, individual persistence and competitive hierarchy (B) (see Methods).

Journal of Vegetation Science2 Doi: 10.1111/j.1654-1103.2012.01434.x© 2012 International Association for Vegetation Science

Plant functional type changes with time since fire C.R. Gosper et al.

Methods

The study was conducted in the south-eastern wheat belt

inWestern Australia. All nature reserves and parcels of un-

allocated crown land were considered for sampling in the

50 km 9 70 km area bounded by Newdegate (33°04′ S,119°04′ E), Lake King, Cocanarup and Pingrup. The region

has a dry mediterranean climate, with average annual

rainfall at Lake Grace (the nearest long-term weather sta-

tion) of 354 mm, mainly falling in winter. Mean monthly

daily temperature maxima range from 15.4 °C to 31.4 °C,and mean monthly minima from 5.6 °C to 15.1 °C(Bureau of Meteorology 2008). The region supports a

mosaic of mallee, mallee-heath and woodland, with vege-

tation type determined by climate and especially edaphic

factors (Beard 1990), and influenced by historic distur-

bance patterns.

The mallee-heath community is characterized by a

diverse shrub layer dominated by serotinous, non-resprouters

(often Proteaceae), with scattered emergent mallees, most

frequently Tallerack (Eucalyptus pleurocarpa) (Gosper et al.

2010). Thematuremallee community is characterized by a

close-spaced canopy of mallees (most frequently E. scypho-

calyx, E. phaenophylla and E. flocktoniae), over a sparse layer

of mostly resprouting shrubs (especially Melaleuca spp.)

and sedges (Parsons & Gosper 2011). Mallees are long-

lived Eucalyptus spp. characterized by numerous aerial

stems, a narrow canopy zone, and a large lignotuber from

which plants resprout after disturbances (Noble 2001).

Experimental design

Five replicates (except where indicated) were located in

each of nine mallee-heath and eight mallee vegetation age

treatments: 2 yr since the last fire (four samples, mallee-

heath only), 3–4 yr, 6 yr (six samples in mallee-heath), 18

–20 yr, 25 yr, 30 yr, 35 yr, 45 yr and ‘long unburned’

(eight samples inmallee-heath). Long unburned should be

interpreted as the site not having experienced fire since at

least 1956, which we have allocated an age of 55 yr post-

fire for analyses (although actual age is likely to be sub-

stantially higher in many cases; see Gosper et al. 2012 for

further details). Information on aspects of fire regime other

than age (such as intensity, previous fire intervals) was not

available. For simplicity in presentation and reflecting gaps

in the span of vegetation ages sampled, we aggregated ages

into ‘young’ (<10 yr post-fire), ‘mature’ (18–35 yr) and

‘old’ (>40 yr).

Our ‘space-for-time’ approach assumed that floristic

composition at each of the different sites is comparable (or

at least that differences between them are randomly dis-

tributed across fire ages; Hurlbert 1994; Oksanen 2001)

and that fire event effects (Bond & van Wilgen 1996) do

not confound time since fire effects. We took a number of

steps to minimize uncertainty in attributing differences to

time since fire. In particular, replicates were spread across

the available range of individual fire events and across the

study area where possible, and where multiple samples

were placed within an individual fire scar, samples were

spaced at least 150-m apart (described further in Gosper

et al. 2012).

Sampling

Plots of 10 m 9 10 m were placed at a random point 20–

150 m into the vegetation from an access track. In spring

2007, we recorded all vascular plant species present and

determined abundance using a line intercept technique by

systematically placing a 12.5-mm diameter pole vertically

at 50 points spread across the plot in a grid. Abundance for

any species was the proportion of points at which any of its

leaves, stems or inflorescences intercepted the pole. This

technique provided an objective measure of abundance

reflecting, but not equivalent to, projected cover, and

encompasses growth of individual plant canopies as well as

changes in plant density. This measure is hereafter referred

to as ‘cover’. Species that were present but not recorded at

point intercepts were allocated a nominal proportional

abundance of 1%.

Plant functional type classification

We classified species on the basis of traits relevant to vege-

tation assembly through the initial floristic composition

model. These were: (i) the capacity to resprout from fire-

resistant organs (e.g. lignotubers, rhizomes, etc.); (ii) the

location and persistence of the seed bank (i.e. persistent

canopy, persistent soil, transient soil); (iii) competitive stra-

tum (upper, mid and ground), largely reflecting plant

height; and (iv) longevity of standing plants (i.e. species

divided into those that grow, reproduce and senesce pri-

marily in the immediate post-fire period (� 6 yr post-fire)

and those that do not (Table 1).

Not all of the resultant 36 possible PFT combinations

were represented in the sampled flora (Table 1). Further,

following Keith et al. (2007), we combined some allied

PFTs to increase sample sizes and thus the capacity to

detect changes, leaving 11 PFTs used in analyses. For resp-

routers, we combined species with transient and persistent

soil-stored seed banks into one PFT per competitive stra-

tum. Due to trade-offs with the capacity for persistence,

recruitment in resprouters is often low (Bond & Midgley

2001), rendering the seed bank a secondary means of per-

sistence in many cases. Representatives of short-lived spe-

cies were combined into a single PFT (post-fire ephemeral

herbs, graminoids and shrubs), occurring across the lower


C.R. Gosper et al. Plant functional type changes with time since fire

two competitive strata, as these species are largely func-

tionally equivalent in avoiding competition with other

PFTs through rapid growth and reproduction post-fire,

then retreating below-ground (in seeds or dormant tubers

or rhizomes) through the bulk of the inter-fire period (see

Keith et al. 2007).

For all species recorded in plots, we used field observa-

tions and published sources to allocate them to PFTs

(Appendix S1). Of the 305 taxa recorded in mallee-heath,

16.4% could not be allocated to a PFT, while in mallee this

was 22.2%of the 243 recorded taxa. Theseunallocated taxa

weremostly in low abundance, as taxa of an unknown PFT

contributed only 5.8%of all cover acrossmallee-heath sites

and 11.4% formallee sites. For each plot, richness and total

cumulative cover of eachPFTwere calculated.

Predicted changes in cover and richness of PFTs with

time since fire

-For each of the four critical traits affecting post-fire com-

munity assembly under the initial floristic composition

model (Table 1), we predict that the 11 subsequent PFTs

would respond as listed in Table 2, based on the following

population and community processes. Due to their limited

longevity (1. Longevity; Table 1) and fire-stimulated germi-

nation, post-fire ephemerals are likely to decline rapidly in

richness and cover with time since fire (Bond & van Wil-

gen 1996).

The capacity to resprout (2. Individual Persistence) gener-

ally confers resistance to change in both richness and cover

with time since fire, as individual plants are often both

highly persistent (Bond & Midgley 2001) and recover bio-

mass rapidly after fire compared to non-resprouters (Keith

& Bradstock 1994; Pausas 1999; Lamont et al. 2011). As

recruitment among resprouters is often low (Bond &Midg-

ley 2001), we suggest that the form of the seed bank

(3. Seed Bank Persistence and Type) will have only minor

effects on changes with time since fire in resprouting PFTs.

Resprouters can decline depending on their position in the

competitive hierarchy (4. Plant Height) through the inter-

fire period, with a simple competition model in which

competitive interactions are determined by the relative

height of individuals, having proven useful in explaining

vegetation changes under specific fire regimes (Keith &

Bradstock 1994; Keith et al. 2007).

Hence, we predict that resprouting PFTs in the domi-

nant upper stratum will have stable richness and stable or

increasing cover with time since fire. In the lowest stratum

most subject to competitive effects in older vegetation, we

predict that resprouting PFTs will either remain stable in

richness due to high levels of persistence of individuals, or

decline in richness (at the plot scale) and decline in cover if

less tolerant of competition. In the middle stratum, at

intermediate levels of competition, changes with time

since fire are predicted to lie between these extremes.

Both serotiny and persistent soil-stored seed banks (3.

Seed Bank Persistence and Type) ensure persistence of propa-

gules through inter-fire intervals, ready to take advantage

of improved conditions for recruitment in the post-fire

environment. Each approach to ensuring seed persistence

has different implications for the longevity of adult plants,

however. For serotinous species, seeds stored on dead

plants or shed after inter-fire plant deaths are typically lost.

This, combined with the importance of maximizing seed

bank size at the time of fire in environments where matu-

ration is slow, inter-fire recruitment is limited and post-fire

Table 2. Plant functional types and their predicted response to increasing periods since fire. %, percentage of all taxa per habitat allocated to each PFT.

Plant Functional Type (code) Mallee Mallee-Heath Predicted Response

% % Richness Cover

Resprouting Serotinous Trees (RStree) 7.0 2.3 Stable Stable or increase

Non-Resprouting Serotinous Trees (NStree) 2.5 3.9 Stable Increase

Resprouting Non-Serotinous Trees (RNtree) 0 1.3 Stable Stable or increase

Non-Resprouting Non-Serotinous Trees & Climbers (NNtree) 2.1 3.6 Stable or decrease Intermediate peak

Resprouting Serotinous Shrubs (RSshrub) 3.3 6.9 Stable Stable

Non-Resprouting Serotinous Shrubs (NSshrub) 10.7 10.2 Stable Increase

Resprouting Non-Serotinous Shrubs (RNshrub) 9.1 8.9 Stable or decrease Stable or decrease

Non-Resprouting Non-Serotinous Shrubs (NNshrub) 21.0 17.4 Decrease Intermediate peak

Resprouting Non-Serotinous Dwarf Shrubs,

Herbs & Graminoids (RNlow)

11.5 15.7 Stable or decrease Decrease

Non-Resprouting Non-Serotinous Dwarf Shrubs,

Herbs & Graminoids (NNlow)

5.3 6.9 Decrease Decrease

Post-Fire Ephemeral Herbs, Graminoids & Shrubs (Ephem) 5.3 6.6 Post-fire only Decrease

Unknown (NA) 22.2 16.4 – –

Total Taxa (n) 243 305



conditions are conducive to seed survival and recruitment

(Lamont et al. 1991, 2007), indicates that high longevity

(relative to typical inter-fire intervals) in individuals is

important for population persistence in non-resprouting

(2. Individual Persistence) serotinous PFTs. Hence, we predict

non-resprouting serotinous PFTs across all competitive

strata (4. Plant Height) will have stable richness with time

since fire, and increasing cover as individuals develop from

small seedlings to large adult plants (Pausas 1999).

As non-resprouting (2. Individual Persistence), persistent

soil-stored seed (3. Seed Bank Persistence and Type) PFTs do

not need to be present as standing adult plants at the time

of fire for population persistence (Weston 1985), high indi-

vidual longevity is less important. Low and mid strata

(4. Plant Height), non-resprouting, non-serotinous PFTs

could avoid competition with dominant PFTs by early

reproduction and accumulating a substantial seed bank

prior to periods of peak competition (Keith & Bradstock

1994; Keith et al. 2007). Individuals could then succumb

to competition and have reduced growth and/or increased

mortality with little implication for population persistence,

provided a fire occurred before the loss of viability of the

soil-stored seed bank. Indeed, a negative relationship

between growth rate and longevity, and a positive rela-

tionship between age of reproductivematurity and longev-

ity, has been established in other systems (Loehle 1988).

If these relationships hold, we predict declining above-

ground richness and declining or an intermediate peak in

cover in subdominant non-resprouting, non-serotinous

PFTs. The dominant canopy non-resprouting, non-

serotinous PFT may, however, show lesser effects of

competition, with stable to decreasing richness and an

intermediate peak in cover predicted.

Statistical analyses

PRIMER analysis software (version 6.1.11, PRIMER-E,

Plymouth, UK) was used for ordination analyses of flo-

ristic and PFT composition. As regional differences in the

flora associated with high rates of species turnover and

endemism in south-western Australia (Cowling et al.

1994) were likely to obscure any time since fire effects

on community composition based on analyses of raw

floristic data, plant species recorded from only a single

part of the study area were omitted from species-level

analyses (not PFT-level analyses). The study region was

broadly divided in two, approximately north–south, by

Lake Magenta and associated salt lakes. Only species

that occurred on both sides of this band of unsuitable

habitat were included in analyses. Although there were

some cases of individual vegetation ages in each habitat

distributed only to one side of the Lake Magenta system,

there was a reasonably even distribution of sites when

ages were aggregated into ‘old’, ‘mature’ and ‘young’

age classes (see Fig. S1 of Gosper et al. 2012). This

reduced total taxa per habitat from 305 to 168 in

mallee-heath, and 243 to 116 in mallee.

Cover data was square-root transformed for both species

and PFT analyses. This transformation gives a stronger

weighting to larger and/or more abundant species or PFTs

than presence/absence, but reduces the influence of larger

and/or abundant species/PFTs compared to when untrans-

formed. We used non-metric, multi-dimensional scaling,

with the Bray-Curtis dissimilarity metric, and PERMANO-

VA and PERMDISP to test for differences in statistical loca-

tion and dispersion, respectively, among vegetation ages.

Significant differences in dispersion between vegetation

ages would indicate an important change in the nature of

the vegetation in its own right, along with possible con-

straints in the interpretation of significant PERMANOVA

analyses. The SIMPER algorithm was used to determine

which species contributed most to similarity within and

dissimilarity between fire ages.

Analysis of variance (ANOVA), using Statistica (Ver-

sion 7.1, Statsoft, Tulsa, OK, US), was used to test for

differences in richness and cover of PFTs and total vege-

tation cover due to vegetation age (young, mature and

old) in each vegetation community. To homogenize

variances, square-root (x + 1) transformation was

applied to mallee total cover, richness of NNtree (see

Table 2 for PFT codes) in mallee-heath and cover of

RNlow in mallee; and natural log (x + 1) transformation

to richness of NA, RNlow and Ephem in mallee-heath,

and cover of NStree, NNtree and RNshrub in mallee and

NStree, RNtree, NNlow and Ephem in mallee-heath.

Due to the absence of RNtree from mallee, and Ephem

from the old age class in mallee, ANOVA was not used

in these cases.

Results

Species composition

Time since the last fire exerted a detectable effect on spe-

cies cover in both habitats. In mallee-heath, sites less than

10 yr post-fire were orientated in one direction on the

ordination, mature (18–35 yr) in another, with old

(>40 yr) somewhat intermediate (Fig. 1a). There were

differences in composition but not dispersion (Table 3)

between time since fire groups, with pair-wise compari-

sons indicating that all mallee-heath age groups were dis-

tinct. Mean between vegetation age group dissimilarity

was greatest between young and mature, but old was on

average more similar to mature than to young (Appendix

S2). Total cover inmallee-heathwas least in young vegeta-

tion, but reached a plateau across both mature and old

vegetation (Table 4).



For analyses of mallee-heath cover, non-resprouting,

serotinous shrubs were the greatest contributors to simi-

larity within vegetation age groups and to dissimilarity

between vegetation age groups (Appendix S2). As pre-

dicted, representatives of this PFT had much lower cover

in young vegetation, higher cover in old and mature veg-

etation, but variable patterns of increase or decrease

between mature and old, probably depending on individ-

ual species’ longevity. Among the species contributing

highly to similarity/dissimilarity within/between vegetation

32

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EUCFLOFLO

EUCPHAPHAEUCSCY

GAHANC

LEPBRU

LEPPUBMELHAM

SPYCOR

3D Stress: 0.18

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BANERYERY

BANRUFCHE

BEAMICMIC

BEASCH

CRYLEU

EUCPLE

HAKCYGCYGHAKPANCRA

MELVILNEUALO

3D Stress: 0.16

Age post-fire 1 = < 10 yrs 2 = 18-35 yrs 3 = > 40 yrs

3

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NSshrub RNshrub

NNshrubRNlow

3D Stress: 0.15

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NStree

NNtree

RSshrub

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RNlow

NNlow

Ephem

3D Stress: 0.14

Age post-fire 1 = < 10 yrs 2 = 18-35 yrs 3 = > 40 yrs

Floristic cover(a)

(b) Cover of PFTs

Fig. 1. Non-metric multi-dimensional scaling ordination of sites in each habitat (left, mallee-heath; right, mallee) by (a) floristic cover and (b) cover of plant

functional types (PFT), with age class indicated by numbers. MDS on square-root transformed data, 100 runs, random start configurations and three-

dimensional final solutions, with bubble size showing the third dimension. Vectors are (a) those for the top three species contributing to similarity within

and dissimilarity between times since fire (Appendix S2, with species names and PFTs; also see main text) and (b) PFTs (see Table 2) with a Pearson

correlation coefficient >0.5.

Table 3. PERMANOVA results for the effect of time since fire on the species and plant functional type (PFT) cover of mallee-heath and mallee vegetation,

and PERMDISP results for differences in dispersion. All mallee-heath df 2, 45; all mallee df 2, 37. Young (Y), < 10 yr; Mature (M), 18–35 yr; Old (O), > 40 yr

post-fire. Pair-wise comparisons show t-value.

PERMANOVA Pair-wise Comparisons Dispersion (Mean ± SE) PERMDISP Pair-Wise Comparisons

Pseudo-F Y vs M Y vs O M vs O Y M O F Y vs M Y vs O M vs O

Species level

Mallee-heath 4.30*** 2.45*** 1.96*** 1.60** 35.8 ± 0.8 33.7 ± 0.9 32.8 ± 0.9 2.59 – – –

Mallee 1.92*** 1.48** 1.29* 1.35* 46.7 ± 2.0 42.9 ± 0.8 43.8 ± 1.6 2.01 – – –

PFT Level

Mallee-heath 8.34*** 3.59*** 2.83*** 1.64** 14.3 ± 0.5 11.7 ± 0.7 10.6 ± 0.6 8.94** 4.10* 2.98* 1.08

Mallee 2.21** 1.60* 1.71** 1.10 17.3 ± 1.2 14.6 ± 0.7 18.2 ± 1.3 4.23* 2.11 0.51 2.76**

***P � 0.001; **P � 0.01; *P < 0.05.



age groups, vectors of some were orientated in ordination

along the division between vegetation ages, whilst others

were associated with particular locations across the study

area. Of those orientated with the division between vege-

tation age groups, the direction of vectors matched PFT

predictions, with resprouters (Melaleuca villosisepala, MEL-

VIL on Fig. 1a) associated with young vegetation and

non-resprouting, serotinous trees (Hakea pandanicarpa,

HAKPANCRA) and shrubs (Banksia erythrocephala, BA-

NERYERY; H. cygna, HAKCYGCYG) with mature vegeta-

tion. Resprouters, including trees, serotinous shrubs and

non-serotinous graminoids, contributed the highest

within-group similarity in young vegetation cover

(Appendix S2). The strongest contributors to similarity in

mature and old vegetation were all non-resprouting,

serotinous trees or shrubs. Between-group dissimilarity

reflected this change from resprouter to non-resprouter

dominance with increasing time since fire.

For mallee vegetation, species cover differed with time

since fire, but dispersion did not (Table 3). Pair-wise com-

parisons indicated that all mallee age groups were distinct

(Table 3), but this was not clearly apparent on the ordina-

tion (Fig. 1a). Further, there was much lower similarity

within ages overall than in mallee-heath (Appendix S2).

Between vegetation age group dissimilarity was greatest

between young and both older groups. The strongest con-

tributors to within/between vegetation age group similar-

ity/dissimilarity were all resprouters and among the

species with greatest covers overall (Appendix S2). Cover

of these resprouters was high at all times since fire,

although contrary to predictions, there was no evidence

for differences in response to time since fire between vege-

tation ages in resprouters occurring in different vegetation

strata. Total cover remained similar across all vegetation

age categories (Table 4).

PFT composition

Time since fire exerted a significant influence on the

PFT composition of both vegetation communities, in

similar ways to analyses based on the composition of

plant species. In mallee-heath, young sites had greater

Table 4. Richness and cover of each plant functional type (PFT) between vegetation age classes in mallee and mallee-heath. See Table 2 for PFT codes.

Mean ± SE for each age class per habitat shown, with F-values and significance levels (****P < 0.0001; **P < 0.01; *P < 0.05) from ANOVA. Different

superscripts indicate significant differences between age classes according to post-hoc Newman-Keuls tests. Young, < 10 yr post-fire; mature, 18–35 yr;

old, > 40 yr. Grey shading indicates differences (or lack thereof) inconsistent with predictions.

PFT Mallee-heath PFT richness Mallee PFT richness Prediction

Young Mature Old F2,45 Young Mature Old F2,37

RStree 1.67 ± 0.2 1.25 ± 0.2 1.69 ± 0.2 1.62 3.40 ± 0.3b 4.65 ± 0.3a 3.80 ± 0.4ab 4.51* Stable

NStree 2.40 ± 0.3 2.85 ± 0.2 2.85 ± 0.3 0.84 0.50 ± 0.3 0.45 ± 0.2 0.90 ± 0.4 0.87 Stable

RNtree 0.47 ± 0.2 0.65 ± 0.2 0.38 ± 0.1 0.80 0 0 0 – Stable

NNtree 2.47 ± 0.5 2.80 ± 0.3 1.69 ± 0.2 2.41 0.60 ± 0.2 0.85 ± 0.2 0.40 ± 0.2 1.36 Stable or decrease

RSshrub 8.67 ± 0.6 7.80 ± 0.5 8.92 ± 0.5 1.21 1.70 ± 0.2 1.70 ± 0.2 2.10 ± 0.4 0.88 Stable

NSshrub 10.8 ± 0.6b 12.3 ± 0.3a 11.6 ± 0.4ab 3.46* 3.10 ± 0.6 3.15 ± 0.6 2.90 ± 0.9 0.03 Stable

RNshrub 7.27 ± 0.7a 5.90 ± 0.4ab 5.31 ± 0.4b 3.70* 4.50 ± 0.3 4.45 ± 0.4 4.40 ± 0.3 0.01 Stable or decrease

NNshrub 9.53 ± 0.6a 9.85 ± 0.3a 6.92 ± 0.6b 10.4*** 8.50 ± 1.2 6.70 ± 0.5 6.50 ± 0.9 1.70 Decrease

RNlow 18.7 ± 1.2a 15.4 ± 0.5a 17.0 ± 0.9a 3.23* 6.90 ± 0.7 7.20 ± 0.7 6.20 ± 0.7 0.43 Stable or decrease

NNlow 4.47 ± 0.5a 2.60 ± 0.3b 2.92 ± 0.6b 6.13** 2.20 ± 0.6 1.95 ± 0.3 1.20 ± 0.3 1.62 Decrease

Ephem 3.07 ± 0.7a 0.35 ± 0.1b 0.62 ± 0.2b 16.7*** 2.60 ± 0.7 0.05 ± 0.1 0 – Post-fire only

NA 9.87 ± 0.9a 6.45 ± 0.3b 5.54 ± 0.6b 11.1*** 7.90 ± 1.0 6.90 ± 0.5 5.30 ± 1.0 2.56 –

Mallee-heath PFT % cover Mallee PFT % cover

RStree 8.80 ± 1.9 9.45 ± 2.0 7.65 ± 1.8 0.22 30.8 ± 3.7a 42.5 ± 2.7b 48.4 ± 3.1b 6.12** Stable or increase

NStree 3.33 ± 0.7a 20.2 ± 2.9b 15.1 ± 3.9b 24.5*** 0.80 ± 0.6 0.90 ± 0.4 5.40 ± 3.4 2.44 Increase

RNtree 0.47 ± 0.2 2.15 ± 0.8 0.54 ± 0.2 2.24 0 0 0 – Stable or increase

NNtree 4.07 ± 1.1a 10.2 ± 1.6b 4.77 ± 1.2a 7.07** 1.80 ± 1.2 1.55 ± 0.4 2.30 ± 1.5 0.10 Intermediate peak

RSshrub 27.4 ± 3.5 22.2 ± 2.1 32.4 ± 4.3 2.67 13.0 ± 4.3 12.0 ± 2.1 16.9 ± 5.0 0.55 Stable

NSshrub 33.9 ± 6.6c 79.3 ± 4.8b 100.2 ± 9.8a 22.6*** 7.40 ± 1.7 6.75 ± 1.6 7.20 ± 2.8 0.03 Increase

RNshrub 14.4 ± 2.0 11.1 ± 1.3 9.77 ± 1.7 1.95 12.5 ± 2.4 9.75 ± 1.0 17.5 ± 4.2 2.55 Stable or decrease

NNshrub 16.7 ± 1.8b 25.5 ± 1.9a 18.4 ± 2.9b 5.13** 23.5 ± 6.6 21.9 ± 2.9 25.6 ± 5.2 0.18 Intermediate peak

RNlow 41.6 ± 5.2 35.9 ± 2.9 31.1 ± 3.7 1.61 22.7 ± 7.3 20.1 ± 2.3 20.0 ± 4.4 0.01 Decrease

NNlow 6.40 ± 1.0a 2.90 ± 0.3b 3.00 ± 0.5b 8.40*** 6.50 ± 2.2 9.45 ± 2.8 5.60 ± 2.3 0.57 Decrease

Ephem 6.73 ± 2.1a 0.35 ± 0.1b 0.62 ± 0.2b 16.9*** 3.90 ± 1.4 0.05 ± 0.1 0 – Decrease

NA 15.3 ± 2.1 10.5 ± 1.2 12.8 ± 2.4 1.92 19.0 ± 2.5 15.1 ± 2.0 18.1 ± 4.8 0.56 –

Total cover 179 ± 7.6b 230 ± 5.7a 236 ± 9.4a 17.3*** 142 ± 10.8 140 ± 5.9 167 ± 16.5 1.80



variability in PFT composition than the other vegetation

ages (Table 3). Young sites also clearly differed in posi-

tion in ordination space, as did old and mature vegeta-

tion, but to a lesser extent (Table 3; Fig. 1b). All time

since fire groups differed in PFT composition in pair-

wise comparisons (Table 3).

Vectors showing the orientation of PFTs largely sup-

ported predictions (Table 2). Post-fire ephemerals, and

non-resprouting and resprouting, non-serotinous dwarf

shrubs, herbs and graminoids were associated with sites

<10 yr post-fire (Fig. 1b). Non-resprouting, serotinous

trees and shrubs had greater cover in old andmature vege-

tation. Resprouting, serotinous shrubs and non-resprout-

ing, non-serotinous trees appeared less responsive to time

since fire, with the orientation of these vectors largely per-

pendicular with the main division between young and not

young vegetation in ordination. Mean similarity within

vegetation ages peaked in mature vegetation (Appendix

S2), with the greatest within vegetation age similarity con-

tributed by resprouting, non-serotinous dwarf shrubs,

herbs and graminoids for all vegetation ages; resprouting,

serotinous shrubs in young vegetation; and non-resprout-

ing, serotinous shrubs in mature and old vegetation. Dis-

similarity between vegetation ages was greater between

young and mature and young and old, than between old

andmature (Appendix S2).

Mallee PFT composition also differed with time since

fire. Times since fire differed in dispersion, although pair-

wise comparisons were inconsistent (Table 3). Given these

differences in dispersion, there is uncertainty as towhether

the significant PERMANOVA result indicates differences in

PFT composition between time since fire groups (Table 3).

The pair-wise comparisons indicate that differences in PFT

composition do exist, as differences largely contradict those

in dispersion, with young vegetation being different from

old and mature, but old and mature vegetation being simi-

lar (Table 3).

As there was poor distinction of vegetation age groups

in ordination (Fig. 1b), interpretation of PFT vector orien-

tation is of little value. Within vegetation age group simi-

larity peaked in mature over young and old vegetation

(Appendix S2). Dissimilarity in PFT composition of vegeta-

tion groups largely increased with the age difference

between them.

Richness and cover of many PFTs varied according to

vegetation age, mostly in the directions predicted

(Table 4). Of the 11 PFTs in mallee-heath, ten responded

as predicted to time since fire for richness and ten in cover.

One exceptionwas richness in non-resprouting, serotinous

shrubs, which was not stable. The other was cover in res-

prouting, non-serotinous dwarf shrubs, herbs and grami-

noids, which had a non-significant decline in cover with

age when a decline was predicted.

In mallee, PFT response to time since fire matched pre-

dictions less well, with seven of the ten PFTs represented

responding as predicted in richness but only four of ten for

cover. The only PFT other than post-fire ephemerals show-

ing a response in richness or cover to time since fire inmal-

lee was resprouting, serotinous trees, which unexpectedly

were most rich in mature vegetation (although pair-wise

comparisons were inconsistent; Table 4) and, as predicted,

increased in cover from young tomature and old.

Of those species with an unknown response to fire, no

differences with time since fire were recorded in mallee,

but more species occurred in young than in mature or old

mallee-heath, but without time since fire differences in

their overall cover (Table 4).

Discussion

Changes in cover and richness of PFTs with time since the

last fire were broadly predicted. This supports the utility of

PFTs as a framework for predicting and interpreting vege-

tation change (e.g. McIntyre et al. 1995; Keith et al.

2007). The use of PFTs based on the initial floristic compo-

sition model of vegetation assembly could thereby contrib-

ute to improved ecological fire management through

predictions of vegetation change under specific fire

regimes supporting decisions on the timing and type of

management interventions.

More of the predicted trajectories of change in PFTs

were upheld inmallee-heath than in mallee. This suggests,

first, that careful consideration should be given to the eco-

logical processes underlying vegetation dynamics when

applying PFTs to predict community responses to fire, and

to other disturbances. The functional classification and

vegetation assembly model applied here may thus be less

useful in predicting changes with time since fire in vegeta-

tion communities not competitively dominated by non-

resprouting, serotinous PFTs. Second, supporting an

increasing body of evidence derived from analogous com-

munities, both in south-western Australia (Yates & Ladd

2010; Gosper et al. 2012) and elsewhere (Horton & Krae-

bel 1955; Bond 1980; Trabaud & Lepart 1980; Keith et al.

2007), our findings indicate substantial changes in compo-

sition with increasing time since fire in fire-prone shrub-

lands, such as mallee-heath.

The most plausible explanation for the differences in

response to time since fire between communities relate

to the relative dominance (at least in cover) of different

PFTs at different times since fire. The dominant, long-

lived resprouters in mallee resist change, suggesting that

communities dominated by resprouters can be inherently

robust to large variation in times since fire (Keeley

1986). In contrast, the dominant, non-resprouting, seroti-

nous PFTs in mallee-heath contributed proportionally



much less to cover in the years immediately after fire, as

they recruited from seed and often lagged in growth

behind resprouters, but became increasingly dominant

over time.

Old vegetation was more distinct from mature vegeta-

tion in mallee-heath than in mallee, with declines in cover

and/or richness of non-resprouting, non-serotinous trees

and shrubs, and resprouting, non-serotinous shrubs,

between mature and old mallee-heath. As most represen-

tatives in these PFTs have persistent soil-stored seed banks,

these changes reflect a transition from above-ground

plants to existing in the soil seed bank. This transition

could suggest vulnerability of mallee-heath to long inter-

vals between fires, depending on seed bank longevity,

which is poorly known for local species. Although seed

banks of some species are very long-lived (Weston 1985),

substantial variation in seed bank longevity between co-

occurring species (Auld 1995; Auld et al. 2000) compli-

cates generalization.

There was a tendency for the richness of post-fire

ephemerals and resprouting, non-serotinous dwarf

shrubs, herbs and graminoids to increase in old mallee-

heath (differences were not significant and inconsistent

in pair-wise comparisons, respectively), relative to

mature vegetation, which was unpredicted. This suggests

that these PFTs have some capacity for expansion in the

absence of a fire-cued establishment event to capitalize

on newly available resources following the senescence

(Bond 1980; Gosper et al. 2012) of some components of

the vegetation. The most plausible mechanisms are

through gradual loss of seed dormancy (Orscheg & En-

right 2011) and recruitment in gaps for post-fire ephe-

merals, and vegetative growth and lateral spread to

avoid competition among resprouting, non-serotinous

dwarf shrubs, herbs and graminoids (Keith et al. 2007).

The capacity for inter-fire recruitment or lateral expan-

sion is often overlooked in studies aimed at establishing

appropriate fire return intervals for vegetation communi-

ties; however it can be significant in some circumstances

(Ooi et al. 2006).

Contrary to predictions, there were no declines in

richness and/or cover in mallee of non-resprouting,

non-serotinous shrubs, or non-resprouting and resprout-

ing, non-serotinous dwarf shrubs, herbs and graminoids.

Why this occurred is unclear, but these results indicate

either that these PFTs tolerate competition, or are capa-

ble of recruitment in the absence of fire (Ooi et al.

2006). There may be less competition for resources

(light, moisture) in mallee than in mallee-heath, as the

overall cover of vegetation is lower (Table 4), especially

in lower strata in mature and old vegetation (Parsons &

Gosper 2011). An additional or alternative possibility is

that hydraulic redistribution of groundwater by the

dominant mallees (Brooksbank et al. 2011) may facili-

tate, rather than reduce, understorey diversity by pro-

viding additional soil moisture during dry periods. There

are no plausible ecological explanations for the unex-

pectedly lower richness of resprouting, serotinous trees

(all mallee Eucalyptus) in young mallee. However, rich-

ness may have been underestimated in young mallee

due to the difficulty in identifying Eucalyptus in the

absence of reproductive material.

Statistical and technical problems may have contrib-

uted to unexpected responses, and if these could be

overcome it may enhance the utility of the PFT

approach. Some PFTs (e.g. resprouting, non-serotinous

trees) had few representatives and contributed little

cover; hence there was limited statistical power to detect

changes between times since fire. More information on

the fire responses of species that could not be assigned

to a PFT may have improved this situation. Grouping of

aligned PFTs, while increasing sample sizes and thus

potential statistical power, may have had the contrary

effect if trait differences unexpectedly contributed to

divergent responses. Variability in functional responses

to fire (Vivian et al. 2010) and misclassifications of spe-

cies based upon this could also have contributed to

unexpected responses. Finally, the age of ‘old’ vegeta-

tion was more than 40 yr (although some sites may

have been substantially older than this; Gosper et al.

2012). As some mallee Eucalyptus are known to live for

centuries (Wellington & Noble 1985), it is possible that

predicted changes in species and PFT composition may

only become apparent over longer time scales than

those sampled. Improved estimation of the age of long-

unburned vegetation (e.g. Clarke et al. 2010) may

improve predictive ability.

The floristic composition and PFT approaches produced

very similar results and implications for management.

While having the outcomes replicated at different levels of

aggregation adds to the robustness of the conclusions, it

also suggests that using a single approach would be more

efficient and not substantially less informative. Each

approach may offer advantages depending on context.

Using PFTs has the advantage of time since fire changes

being less subject to effects driven by regional differences

in the flora, which in our case required the omission of

some species from the analyses. Further, the similarity of

the PFT model and in the responses of PFTs to time since

fire to that found by Keith et al. (2007) demonstrates a

generality in responses in fire-prone shrublands domi-

nated by non-resprouting, serotinous shrubs, indicating

that the model may be applicable to such communities

more broadly. Alternatively, there may be cases where a

lack of trait data for defining PFTs may place greater limits

on analyses than using floristic composition.



Acknowledgements

This study was jointly funded by the Department of Envi-

ronment and Conservation’s (DEC) Saving Our Species

Initiative and CSIRO Ecosystem Sciences (CES). The spa-

tial distribution of sampling was based in part on remote

sensing data derived from the research of Dr Li Shu, in

digital image processing and remote sensing at Fire Man-

agement Services, Regional Services Division, DEC. We

thank Anne Rick for assistance with floristic surveys, and

Georg Wiehl (CES), Blair Parsons (CES), Tanya Llorens

(DEC) and Hafeel Kalideen (DEC) for field and technical

assistance. Janet Franklin and David Keith provided valu-

able comments on themanuscript.

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Supporting Information

Additional supporting information may be found in the

online version of this article:

Appendix S1. All taxa recorded, with their plant

functional type classification.

Appendix S2. Species and plant functional types

contributing most to similarities within and differences

between times since fire age classes in mallee-heath and

mallee.

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