Classificação de Comunidades de Formigas

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A Classification of Australian Ant Communities, Based on Functional Groups Which Parallel
Plant Life-Forms in Relation to Stress and Disturbance
Author(s): Alan N. Andersen
Source: Journal of Biogeography, Vol. 22, No. 1 (Jan., 1995), pp. 15-29
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Joumal of BiogeographV (1995) 22, 15-29 
A classification of Australian ant communities, based 
on functional groups which parallel plant life-forms in 
relation to stress and disturbance 
ALAN N. ANDERSEN Division of Wildlife and Ecology, CSIRO Tropical Ecosystems Research Centre, PMB 44 
Winnellie, NT 0821, Australia 
Abstract. A system is proposed whereby ant functional are structurally analogous to grassy forests. The distribution of 
groups are used as structural attributes to classify ninety-four ant functional groups is considered in relation to stress and 
Australian ant communities in a manner analogous to the disturbance by adopting Grime's (1979) triangular ordination 
classification of vegetation according to predominant life- concepts and nomenclature, with ant community structural 
forms. In terms of their responses to stress and disturbance, types being analysed in terms of the relative importance of 
Dominant Dolichoderinae (DD) are considered analogous to competition, stress and disturbance as factors regulating com- 
trees, functionally subdominant Generalized Myrmicinae munity structure. DDO and DD1 structural types are stress-tol- 
(GM) to shrubs and ruderal Opportunists (OPP) to grasses. erant, or ruderal, communities; DD2 and DD3 types are 
Community types DDO (twenty-two sites), DDl (twenty-two competitive communities when Generalized Myrmicinae are 
sites), DD2 (eight sites), DD3 (thirty-nine sites) and DD4 abundant, and competitive ruderal or competitive stress-toler- 
(three sites), respectively, are defined as having the relative ant ruderal when Opportunists are predominant among non- 
abundance of Dominant Dolichoderinae 90%. They are structurally analogous ruderal. In temperate regions, seasonal changes in ant com- 
to treeless plant communities, open woodlands, woodlands, munity structure parallel those occurring along biogeographi- 
forests and plantations, respectively. DDO communities are cal gradients spanning comparable temperature regimes. A 
classified as DDOGM (analogous to shrublands) when Gener- positive relationship was found between the abundance of 
alized Myrmicinae predominate, DDOOPP (analogous to functionally dominant ants (DD + GM) and species richness. 
grasslands) when Opportunists predominate and DDOCS Plant and ant communities often differ from each other in their 
(analogous to cold-adapted heathlands) when neither func- responses to the same stress or disturbance, such that there is 
tional group is abundant. Similarly, the relative abundances of often a poor correspondence between ant and plant community 
Generalized Myrmicinae and Opportunists are used to classify structural type at any particular site. 
DD 1-3 communities in a manner analogous to the 
classification of woodlands and open forests according to 
understorey type. DD30PP communities, for example, where Key words. Ant communities, community classification, 
the relative abundance of Dominant Dolichoderinae is 30- community structure, competition, disturbance, functional 
70% and Opportunists are predominant among remaining ants, groups, stress. 
INTRODUCTION 
Plant ecologists have long sought to produce community 
classification schemes based on the identification of func- 
tional groups whose relative abundances vary predictably 
in response to external limiting factors. Examples include 
the life-form system of Raunkier (1934) and the 'vital 
attributes' model of Noble & Slatyer (1980). These 
schemes allow for the identification of global patterns of 
community structure (Box, 1981; Woodward, 1987) and 
provide a framework for analysing the responses of plant 
communities to environmental stress and disturbance 
(Grime, 1979; Tilman, 1982, 1988; Werger et al., 1988). 
Such global models of community structure and dynamics 
are notably lacking for animals. This paper aims to provide 
such a model for Australian ant communities. 
One explanation for the absence of global classification 
schemes for animal communities is that they are not poss- 
ible, due to fundamental differences between plants and 
? 1995 Blackwell Science Ltd. 15 
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16 Alan N. Andersen 
animals in community structure. For example, climate de- 
termines vegetation structure to an extent that might not 
occur in animal communities, and interspecific competition 
is a dominant force among plants (Grime, 1979; Tilman, 
1982, 1988) but not necessarily animals (Strong et al., 
1984). Moreover, vegetation classification is based on 
clearly defined structural attributes (plant life-forms) that 
may have no equivalent in animal communities. 
In many important respects ant colonies behave more 
like plants than animals, and this has important implica- 
tions for community structure (Andersen, 1991a). Like 
plants, ants are modular organisms, consisting of an inde- 
terminate number of repeated units of multicellular struc- 
ture (modules; Harper, 1977, 1981). Both ants and plants 
'nest' in a fixed position, usually in the ground, and re- 
source capture is achieved through the ramification of 
foraging modules (Harper, 1985). Most plants have broadly 
similar ecological requirements (Grime & Hodgson, 1987; 
Latham, 1992), using roots and stolons to forage in the soil 
for water and nutrients, and shoots and leaves to forage 
above-ground for sunlight and carbon dioxide. Therefore, 
there is considerable ecological overlap between plant taxa 
varying enormously in morphology, such as canopy trees 
on one hand and herbs of the forest floor on the other hand. 
A similar situation also occurs in ant communities. Most 
ants have similar ecological requirements (H1ldobler & 
Wilson, 1990), foraging on the ground and vegetation as 
generalized scavengers, predators and collectors of plant 
exudates. Moreover, the prevalence of interference compe- 
tition means that ecological interaction often also occurs 
between ant species utilizing different resources. 
These attributes mean that competition for space and 
resources is more pronounced in plants (Grime, 1979; 
Tilman, 1982, 1988) and ants (Holldobler & Wilson, 1990) 
than in most other biological communities (Strong et al., 
1984). Moreover, the ecological overlapping of component 
species means that the conventional partitioning of animal 
communities into guilds based on resource utilization (Ter- 
borgh & Robinson, 1986) is of limited use for plants and 
ants. In these two taxa, functionalRoger, Hypoponera Santschi, Monomo- 
rium talpa Emery, Oligomyrmex Mayr, Plagiolepis Mayr, 
Ponera Latreille, Quadristruma Brown, Solenopsis 
(Diplorhoptrum Mayr), Sphinctomyrmex Mayr, Stru- 
migenys F. Smith, Trachymesopus Emery. 
Opportunists 
Aphaenogaster Mayr, Cardiocondyla Emery, Doleromyr- 
ma Forel, Ochetellus glaber (Mayr) group, Odontomachus 
Latreille, Paratrechina Motschoulsky, Rhytidoponera 
Mayr, Tapinoma Forster, Technomyrmex Mayr, Tetramori- 
um Mayr. 
Generalized Myrmicinae 
Crematogaster Lund, Monomorium Mayr (part), Pheidole 
Westwood. 
Specialist predators 
Anochetus Mayr, Bothroponera Mayr, Cerapachys F. 
Smith, Colobostruma Wheeler, Epopostruma Forel, Lep- 
togenys Roger, Mesostruma Brown, Myrmecia Fabricius, 
Platythyrea Roger. 
? 1995 Blackwell Science Ltd, Journal of Biogeography, 22, 15-29. 
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	Article Contents
	p. 15
	p. 16
	p. 17
	p. 18
	p. 19
	p. 20
	p. 21
	p. 22
	p. 23
	p. 24
	p. 25
	p. 26
	p. 27
	p. 28
	p. 29
	Issue Table of Contents
	Journal of Biogeography, Vol. 22, No. 1 (Jan., 1995), pp. 1-160
	Front Matter [pp. ]
	Special Paper: Is Greenland a Zoogeographical Unit of Its Own? [pp. 1-6]
	Special Paper: What Determines the Probability of Discovering a Species?: A Study of South American Oscine Passerine Birds [pp. 7-14]
	Southern Hemisphere Zoogeography
	A Classification of Australian Ant Communities, Based on Functional Groups Which Parallel Plant Life-Forms in Relation to Stress and Disturbance [pp. 15-29]
	Biogeography of the Australian Dynastinae, Rutelinae, Scarabaeinae, Melolonthini, Scitalini and Geotrupidae (Coleoptera: Scarabaeoidea) [pp. 31-48]
	A Mediterranean Origin for the Veldrif (South Africa) Artemia Leach Population [pp. 49-59]
	Insular Biogeography of Birds on Mountain-Tops in North Western Argentina [pp. 61-70]
	Northern Hemisphere Zoogeography
	Co-Occurrence of Cetaceans and Seabirds in the Northeast Atlantic [pp. 71-88]
	Biotic Affinities in a Transitional Zone Between the Atlantic and the Mediterranean: A Biogeographical Approach Based on Sponges [pp. 89-110]
	Geographical Ecology of Mongolian Desert Rodent Communities [pp. 111-128]
	The Effects of Forest Fragmentation on Butterfly Communities in Central Spain [pp. 129-140]
	Habitat Distribution of Terrestrial Coleoptera in Iceland as Indicated by Numerical Analysis [pp. 141-148]
	Correspondence
	On the Holocene Palaeoenvironmental Record from Lake Tyrrell, Northwestern Victoria, Australia-a Reply [pp. 149-152]
	Holocene Palaeoenvironments at Lake Tyrrell-Response to Sluiter and Parsons [pp. 152-156]
	Book Reviews
	Animal Acts in the Heath [pp. 157-158]
	Tales from the River Bank: A Valuable Contribution [pp. 158-159]
	Perhaps not Worth the Climb [pp. 159]
	What's in a Name? [pp. 159-160]
	Back Matter [pp. ]groups are based on a 
range of morphological and behavioural attributes that have 
striking parallels with each other (Andersen, 1991a). 
The parallels between ant and plant community structure 
suggest that ant communities might usefully be classified 
according to structural attributes that parallel those adopted 
in vegetation science. The objective of this paper is to 
provide such a classification for Australian ant communi- 
ties, and to use it as a basis for analysing the responses of 
ant community structure to stress and disturbance. It must 
be emphasized that this paper deals specifically with pat- 
terns of community structure occurring on a biogeographi- 
cal scale (as does vegetation classification), and does not 
purport to be a comprehensive treatment of community 
dynamics at individual sites. 
ANT FUNCTIONAL GROUPS 
Greenslade (1978) has proposed a functional group 
classification of Australian ants based on their postulated 
competitive interactions, habitat requirements and evol- 
utionary history. This scheme has subsequently been 
modified to place a greater emphasis on community dy- 
namics than on evolutionary history (reviewed by An- 
dersen, 1990, 1992), and provides the structural attributes 
upon which the community classification proposed here is 
based. The functional groups have been discussed exten- 
sively elsewhere (see references below), and are only 
briefly outlined here. There is often a strong relationship 
between the systematics and ecological behaviour of ani- 
mal species (Brooks & McLennan, 1991; Spence & An- 
dersen, 1994), and this is reflected in a taxonomic basis for 
some of the groups. Such a taxonomic basis is also recog- 
nized for functional groups of other insect taxa, such as 
butterflies (Hodgson, 1993). The taxa (in most cases gen- 
era) assigned to each group, along with their nomenclatural 
authorities, are listed in the Appendix. 
Dominant Dolichoderinae The dolichoderine genus 
Iridomyrmex is virtually ubiquitous in the Australian en- 
vironment and consists of abundant, highly active and 
aggressive species that exert a major competitive influence 
on other ants (Greenslade, 1976, 1979; Andersen, 1992; 
Andersen & Patel, 1994). They are particularly abundant 
and diverse in hot and open habitats, which allow for high 
rates of foraging activity, and are often absent from heavily 
shaded sites. Iridomyrmex is replaced by Anonychomyrma 
as the dominant dolichoderine genus in cooler and wetter 
regions of southern and eastern Australia. Oecophylla is 
also a competitively dominant ant (Holldobler & Wilson, 
1990) but, because of its restricted distribution (tropical 
forests) and arboreal habit, I place it in another functional 
group (Tropical climate specialists; see below). I consider 
it as a taxon that can achieve dominance only in the 
absence of Iridomyrmex. 
Subordinate Camponotini Camponotine formicines, 
especially species of Camponotus, are also virtually ubiqui- 
tous in the Australian environment, with up to twenty or 
more species occurring at a single site (Andersen & Yen, 
1985). They are behaviourally submissive to Iridomyrmex, 
and therefore competitively subordinate in their presence, 
but can be competitively dominant in their absence (An- 
dersen & Patel, 1994). Despite their ubiquity and richness, 
their relative abundance in any community is generally 
low. 
Climate specialists These taxa have distributions 
heavily centred on one of three distinct climatic zones: the 
arid zone (Hot climate specialists), the humid tropics 
(Tropical climate specialists) and cool-temperate regions 
(Cold climate specialists). Both Cold and Tropical climate 
specialists are characteristic of habitats where the 
abundance of Dominant dolichoderines is low and, apart 
from their habitat tolerances, are often unspecialized ants 
in terms of foraging ecology. Hot climate specialists, on 
the other hand, are characteristic of sites where Domi- 
nant dolichoderines are most abundant, and possess a range 
of physiological, morphological and behavioural 
? 1995 Blackwell Science Ltd, Journal of Biogeography, 22, 15-29. 
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Classification of Australian ant communities 17 
specializations relating to their foraging ecology, which 
reduce their interaction with other ants. The most striking 
example is that of Melophorus species, which are excep- 
tionally thermophilic (Christian & Morton, 1992), foraging 
when few or no other ants are active. 
Cryptic species These are small to minute species, 
predominantly of the subfamilies Myrmicinae and Poneri- 
nae, which nest and forage almost exclusively within soil 
and litter, and therefore probably have little interaction with 
other ants. 
Opportunists These are ruderal species (Grime, 
1979), characteristic of sites where stress (e.g. waterlogged 
soils, low food availability, cold climate) or disturbance 
severely limit ant productivity and diversity. They are 
unspecialized, poorly competitive species whose distribu- 
tions appear to be strongly influenced by competition from 
other ants. 
Generalized Myrmicinae The three cosmopolitan 
myrmicine genera Pheidole, Monomorium and Cremato- 
gaster are ubiquitous and often highly abundant in Aus- 
tralian ant communities. They have generalized habits in 
relation to nesting and dietary requirements, but are ex- 
tremely competitive at rich food sources, recruiting rapidly 
to them, and being able to defend them even from Domi- 
nant dolichoderines (Andersen, Blum & Jones, 1991). Gen- 
eralized myrmicines depend on rapid recruitment and mass 
mobilization for their success-unlike Dominant doli- 
choderines, individuals are not highly active and aggress- 
ive, and they tend to have small foraging ranges. 
Specialist predators This group is used here for the 
first time, replacing (but including) Greenslade's (1978) 
'Large, Solitary Foragers'. It comprises medium- to large- 
sized species which are specialist predators of other 
arthropods. Group raiders such as Leptogenys and Cera- 
pachys are included, along with solitary foragers such as 
Myrmecia (which are considered 'specialist' predators in 
terms of size, rather than type, of prey taken), Pachycondy- 
la and non-cryptic dacetines. They tend to have little 
competitive interaction with other ants due to their special- 
ized diets and typically low population densities. 
CLASSIFYING AUSTRALIAN ANT COMMUNITIES 
Attributes used for classification 
The scheme proposed here uses three of the above func- 
tional groups as structural attributes. These are: Dominant 
Dolichoderinae, Generalized Myrmicinae and Oppor- 
tunists. Dominant Dolichoderinae provide a fundamental 
structural framework in ant communities; under favourable 
conditions (discussed later) they contribute substantially to 
total ant biomass, and exert a pervasive competitive 
influence over other taxa. Generalized Myrmicinae can also 
be functionally dominant, but their competitive influence 
tends to be expressed more locally, such as by the monop- 
olization of a rich food source close to where they nest. 
They never attain the biomass that can be achieved by 
Dominant dolichoderines, but are abundant in a broader 
range of habitats. Opportunists are poorly competitive in 
relation to most other ants, and predominate only under 
conditions of disturbance or stress. They tend to be the 
major functional group in ruderal habitats. For the remain- 
der of this paper Dominant Dolichoderinae, Generalized 
Myrmicinae and Opportunists will be abbreviated to DD, 
GM and OPP, respectively. 
The classification scheme directly uses only three of the 
seven functional groups. Subordinate Camponotini and 
Specialist foragers are present at most sites, but rarely if 
ever numerically dominant. They are therefore of limited 
use in discriminating structuraltypes. Climate specialists 
and Cryptic species have far more restricted distributions 
but, as outlined later, these are correlated with the three 
functional groups used directly. 
The data set 
For this analysis all published studies that quantify ant 
species composition and relative abundance at an Aus- 
tralian site have been compiled. There are ninety-four such 
sites and each has been assigned a code (Table 1) that will 
be used throughout this paper. The sites are concentrated in 
three biogeographic regions (Fig. 1): the semi-arid and 
mesic southeast (thirty-one sites), the Mediterranean south 
west (twenty-eight sites) and the north western seasonal 
tropics (twenty-five sites). The central arid zone, Queens- 
land and New South Wales are all poorly represented, and 
no sites are available from Tasmania. Despite this patchy 
geographical coverage all major vegetation types are repre- 
sented, including various grasslands, heathlands, shrub- 
lands, woodlands, savannas, open forests and rain forests 
(Table 1). However, it should be noted that humid tropical 
rain forests are not represented at all, nor is any alpine 
vegetation. The data set also includes numerous disturbed 
sites affected by mining, fire, grazing, plantation forestry 
and urbanization. 
In most cases, a site is a plot of 1 ha or less. The 
exceptions are ENEA 1-3, each representing data from 
several small plots within a few kilometres of each other, 
and KUN, MORG and CARID 1-3, each of which repre- 
sents regional compilations from areas of the order of 100 
km2. Data on species occurrences were obtained by a 
variety of methods, but in all but two cases the quantitative 
data on species relative abundances are results from pitfall 
traps. The two exceptions are EVALE 1 and 2, where 
relative abundances are based on colony densities. There is 
obviously considerable variation in sampling intensity 
which affects the reliability of the data, especially in rela- 
tion to species richness. For each site, sampling intensity 
has been rated according to a five-point scale (Table 1), 
where '1' indicates that a site was sampled at low to 
moderate intensity on a single occasion, and '5' indicates 
that a site was intensively studied over at least a 12-month 
period. For analyses of species richness, only sites with 
C 1995 Blackwell Science Ltd, Journal of Biogeography, 22, 15-29. 
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18 Alan N. Andersen 
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Classification of Australian ant communities 19 
CO 
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0 
lCC 
.= oo CC) ONONONONONONONONONONONON CONONONONONO' C' - oo oo oo oo oo oo oo oo oo oo oo oo Ch cr e 000000 c c c e 
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? 1995 Blackwell Science Ltd, Journal of Biogeography, 22,~C 15-29 
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20 Alan N. Andersen 
HOLN 
yKIM RANG 
BROC LHILL 
X ENEA 
SW 
DWEL R EVALE 
ARID * RG 
LOFY EL G 
WPROM 
FIG. 1. Location of study sites used for ant community classification 
(see Table 1). 
sampling intensity rated three or greater (n = 44) are con- 
sidered. 
Community classifications 
The first distinction made is between communities with and 
without abundant DD. The relative abundance of DD 
ranges from 0 to 100% (Fig. 2), and 10% is used as an 
arbitrary threshold for defining whether or not DD is 
structurally dominant. 
There are twenty-two (23%) sites with %DD 30 
and %OPP 50; and (3) four sites with %GM 15 and > %OPP (DD1GM, nine sites); 
%OPP > 20 and > %GM (DD1OPP, ten sites); and 
%GM %OPP (non-DD) 
DD3GM 
%OPP (non-DD) > 20 and S %GM (non-DD) 
DD30PP 
%GM (non-DD)2 
- -60 SWAN I1 
60 RANGE GLEN 2 DD2,3GM O- 6 RANG 4 () SSWAN 12 RAN4 
t;; WPROM 10 / tj; ENEA7 
DWELS3 L 17 ,WPROMB 6 
WPROM8 
a40 SWAN SWAN9 R / | C40 -- M . D 
? 1;2WPROM3,DWEL2 CA,03 , 0 WGROM I / GM o 
iL 1 'm eLHLL 6 LHILLS 0 
L 
MUNM. WPNOM7 
20: , LHILL2 RANG8 EVALE1 20 POM * EVALE2 
WROSLHILLS GSLEN I SWELl *ML M W 
N 
S 
MUNM 
3 
'SWAN 7 MUNM4 
MORG CAIDD2 LH LL;5 KAP MELT ENEAS 6 4WPROM 12 M 
0WPROMB 
9 r I 
0YP2.WPRM14 LOFTY 2 
o 20 40 60 80 100 0 20 40 60 80 
Generalized Myrmicinae (%) Generalized Myrmicinae (%) 
100 
DDOOPP 
(c) 
80 - KIMB 8 
SWANS 3 
ROT 3 
SWAN 3 
BROO 
RANG 3 
60 - SWAN 6 
- 000DDOGM 
C 1 WPROMS 2 |} 40 - , -ROM 2 I 
HOLM 
DDOCS KIMB 3S 
ROTS 2 MB 2 
20; XSPROMS 
RGFIOT 
1 KIMB7 M 4 
WPROM 15,16 KIB IMBi1 
/ WPROM15 , 
KIMB 5 
KIMBM 
SWAN 5 
0 20 40 60 80 100 
Generalized Myrmicinae (%) 
FIG. 3 (a) Classification of DDO ant communities (Dominant Dolichoderinaedominant 
ants (dominant Dolichoderinae + generalized Myrmicinae) and total ant 
species richness. Only sites where data were collected from single plots 
and sampling intensity is rated at least three (Appendix; n = 44) are 
included. The regression equation is y = 0.34x + 14.28 (r2 = 0.178, 
P 0.05 
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24 Alan N. Andersen 
etation processes based on the recognition of three primary 
life-history strategies in plants in relation to stress and 
disturbance. According to Grime, competitors (C) predomi- 
nate at sites experiencing low stress and low disturbance, 
stress-tolerators (S) occur under conditions of high stress 
and low disturbance, and ruderals (R) are characteristic of 
sites experiencing low stress and high disturbance (environ- 
ments experiencing both high stress and high disturbance 
are not viable for vascular plants, and so there is no fourth 
strategy). The three different environments are subject to 
K-, A- and r-selection respectively (MacArthur & Wilson, 
1967; Greenslade, 1983), and can be represented on a 
triangle with axes corresponding to each of the three major 
selective pressures (competition, stress and disturbance; see 
also Southwood, 1977). The merits of CSR theory in 
relation to life history attributes have been hotly debated 
(Loehle, 1988; Southwood, 1988; Midgley, 1993; Oksanen, 
1993), and this debate is not entered into here. Rather, 
Grime's nomenclature is used as a framework for repre- 
senting ant community structure in relation to stress and 
disturbance; instead of using the triangular ordination in 
relation to history strategies, it is used to map and predict 
community-level structural attributes. 
Three primary types of communities can be identified 
(Fig. 7a): competitive (C),where both stress and disturb- 
ance are low, and competition is the primary factor regulat- 
ing community structure; stress-tolerant (S; stress high, 
disturbance low), where stress is the primary factor regulat- 
ing community structure; and ruderal (R; stress low, dis- 
turbance high), where disturbance (or disturbance-induced 
stress) is the primary factor regulating community structure 
(Andersen, 1991a). Secondary communities can also be 
recognized (Fig. 7a) where stress and/or disturbance are 
moderate: competitive ruderal (C-R), with low stress and 
moderate disturbance; stress-tolerant ruderal (S-R), with 
stress and disturbance both moderate, and stress-tolerant 
competitive (C-S), with moderate stress and low disturb- 
ance. 
Table 4 classifies the different structural types of ant 
communities in relation to stress and disturbance, and these 
are plotted on the CSR habitat template in Fig. 7b. DDO 
and DD1 structural types, where Dominant Dolichoderinae 
are poorly represented, are stress-tolerant or ruderal com- 
munities. They are stress-tolerant (S) when GM predomi- 
nate, ruderal (R or SR) when OPP predominate and, with 
one exception, 'super' stress-tolerant (SS; occupying cool 
and shaded sites) when neither GM or OPP are abundant 
(Cold climate specialists predominate). 
DD2 and DD3 structural types are competitive (C) when 
GM are abundant, stress-tolerant competitive ruderal (CSR) 
or competitive ruderal (CR) when OPP are predominant 
among non-DD, and competitive stress-tolerant (CS) when 
neither GM nor OPP are abundant. DD2,3GM (C) com- 
munities are restricted to hot and open habitats, and have 
highest diversity (Fig. 4). They all include at least several 
species of Melophorus. DD2,30PP structural types occur 
under a wide range of conditions (Table 4). At hot and 
open sites with stony soils, Generalized myrmicines tend to 
be replaced by large species of Rhytidoponera, producing 
CSR communities. Large species of Rhytidoponera are also 
(a) 
C 
competition disturbance 
~C-S \C-R/ 
/ X t-S-F4\ / \ 
/ S ".,S-R/ R 
stress 
(b) 
0D2,3GM 
I/ DDZ3OPP: \,DD2,3CS/ / 
bXD2,30PR / 
DD0,1GM\k^01yD0lPP' DD0,10PP 
O DOiCS\ 8 
FIG. 7. Classification of communities in relation to stress and 
disturbance following the nomenclature of Grime (1979). A generalized 
habitat template is shown in (a). Three primary community types are 
recognized: competitive (C), stress-tolerant (S) and ruderal (R) which, 
respectively, occur under conditions of low stress and disturbance, high 
stress and low disturbance and low stress and high disturbance. Various 
secondary community types are also recognized, occurring when stress 
and/or disturbance are moderate (see text for details). In (b), the major 
structural types of ant communities identified in the present study are 
plotted on the CSR triangular ordination. 
abundant at highly disturbed sites (CR communities), but 
species richness is low and Melophorus is poorly repre- 
sented. In open habitats of temperate regions (CSR com- 
munities), species of Anonychomyrma are often the most 
abundant Dominant dolichoderine, Melophorus is poorly 
represented and small species of Rhytidoponera (metallica 
and allies) are abundant. DD4 structural types are all 
low-diversity, CR communities occurring at highly dis- 
turbed sites. 
DISCUSSION 
The classification of Australian ant communities proposed 
here has several potential uses. First, it provides simple 
pigeon-holes to facilitate communication between re- 
searchers. Secondly, it forms a framework for biogeograph- 
ical comparisons of ant communities from a diverse range 
of environments, even when species pools have little or no 
overlap. This allows for the identification of continent-wide 
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Classification of Australian ant communities 25 
patterns of ant community structure. Thirdly, it can help 
identify the major factors (relating to stress, disturbance 
and competition) structuring ant communities at a biogeo- 
graphical scale. As discussed below, low-temperature 
stress, determined through a combination of climate and 
vegetation structure, is highlighted here as a key factor 
determining ant community structure, particularly through 
its effect on dominant species. Finally, the classification 
allows for the prediction of ant community structure at sites 
where empirical data are unavailable. For example, ant 
community structure at the two most important Australian 
biomes not represented in the current data set, humid 
tropical rain forest and alpine vegetation, can be predicted 
from the analysis presented here. Humid tropical rain 
forests are likely to support structurally similar communi- 
ties to those occurring in monsoonal rain forests (i.e. 
primarily DDOGM communities), but presumably with a 
greater representation of Tropical climate specialists. 
Alpine vegetation is likely to support DDO, 1OPP or 
DDO1CS communities, as occur in lowland, cool-temper- 
ate Australia. 
Stress and disturbance 
Low-temperature stress is highlighted as a key factor regu- 
lating ant community structure; in particular, it controls the 
abundance of Dominant Dolichoderinae, and therefore 
largely determines competitive dynamics within the com- 
munity. Vegetation structure plays a key role in regulating 
low-temperature stress through its effects on microclimate. 
For example, at Wilson's Promontory in cool-temperate 
southern Victoria, ant community structure ranges from 
DDOCS and DDICS (SS) at the most heavily shaded sites 
(WPROM 12-17), to DDOOPP and DDlOPP (SR; 
WPROM 1,2,7,8,11) and DD2CS (CS; WPROM 5,9) and 
DD30PP (CSR; WPROM 3,6,10) as the level of insulation 
on the ground increases. Similarly, in monsoonal Australia, 
savanna vegetation tends to support DD3GM(C) ant com- 
munities containing numerous Hot climate specialists 
(KAP, MUNM 1-3), whereas rain forest characteristically 
supports DDOGM(S) communities containing few, if any, 
Hot climate specialists, but numerous Tropical climate 
specialists (HOLM, KIMB 1-7). 
The seasonal changes in ant community structure (based 
on foraging activity) that occur in temperate regions (Table 
2) mirror changes associated with biogeographical gradi- 
ents spanning comparable temperature regimes. For exam- 
ple, the structural type in a temperate woodland (WPROM 
1) during winter (DDOCS) is the same as that overall in 
more heavily shaded habitats of the region (WPROM 13, 
TABLE 4. Classification of structural types of Australian ant communities in relation to stress and disturbance, based on the nomenclature 
of Grime (1979). Competitive (C) communities occur where both stress and disturbance are low, stress-tolerant (S) communities occur where 
stress is high and disturbance is low, and ruderal (R) communities occur where stress is low and disturbance high. See text for details. 
Structural 
type Sites C-S-R Dominant habitat features 
A. DDO AND DD1 TYPES 
DDO, 1GM ROT 1,2; SWAN 4,5,7; S Cool/mild and moderately shady 
LOFTY 2; WPROM 4 
HOLM, KIMB 1-7; MUNM 4-6 S Hot and shady 
EVALE 2, ENEA 5 S ? 
DDO, 1OPP ROT 3; BROO; RANG 2-4 R Highly disturbed, and cool or shady 
SWAN 1,3,6,11; WPROM 1,2, SR Cool/mild and shady 
7,8,11 
KIMB 8, GLEN 3 SR Warm/hot and very shady 
ENEA 7 SR ? 
DDO, 1CS WPROM 12-17 SS Cool and very shady 
ENEA 3 R Highly disturbed 
B. DD2 AND DD3 TYPES 
DD2, 3GM MUNM 1-3; LHILL 2,3,5,8; C Warm/hot and open 
ENEA 4,6; LOFTY 1; KUN; 
MORG; CARID 1,2; MELT; 
GLEN 1; DWEL 1; RANG 8; 
EVALE 1; KAP; WYP 1,2 
DD2, 3OPP ENEA 1,2; RANG 5,6; DWEL CR Highly disturbed, with both shady and open patches 
2- 4 
LHILL 1,4,6,7,9; RANG 7 CSR 
Hot and open, with either stony soils or heavy litter 
CARID 3; GLEN 2; SWAN CSR Cool/mild and open 
2, 8,9,10,12; WPROM 
3,6,10DD2, 3CS WPROM 5,9 CS Cool and shady 
C. DD4 TYPE 
RANG 1; DWEL 5,6 CR Highly disturbed 
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26 Alan N. Andersen 
15-17), whereas in summer (DD20PP) it is the same as 
that overall in similar vegetation types experiencing 
warmer climates (e.g. GLEN 2, SWAN 12). 
A positive relationship was found between the abun- 
dance of functionally dominant ants and species richness. 
This directly contradicts Wilson's (1990) 'dominance-im- 
poverishment rule', which states that an inverse relation- 
ship exists between species richness and the degree of 
competitive dominance in ant communities. The positive 
relationship shown here suggests that both competitive 
dominance and species richness respond similarly to en- 
vironmental stress and that, on a continental scale, the 
positive effects of decreasing stress on richness over-ride 
the negative effects of increasing competitive pressure. 
This continental pattern should not be confused with local 
patterns of species richness, where competition from domi- 
nant species has been shown to reduce diversity (Andersen, 
1992; Andersen & Patel, 1994). 
The only time that high competitive dominance was 
associated with species-poor communities was at heavily 
disturbed sites (DD4 communities). It is noteworthy that 
disturbance is also implicated in the association of high 
competitive dominance with low diversity elsewhere in the 
world (Wilson, 1990), following invasion by highly com- 
petitive exotic species such as Solenopsis invicta, Linep- 
ithema humile and Pheidole megacephala (Haskins & 
Haskins, 1988; Porter & Savignano, 1990). Colonization by 
P. megacephala following disturbance has also been shown 
to cause a marked reduction in ant diversity in Queensland 
(Majer, 1985b). 
It was previously mentioned that the major effects of site 
disturbance on ant communities is often secondary and 
stress-related, particularly when disturbance is not severe. 
This is illustrated by the effects of fire which, by reducing 
vegetation cover, reduces low-temperature stress. In the 
monsoonal tropics, for example, unburnt open forests 
(MUNM 5,6) support stress-tolerant communities, which 
change to competitive communities under a regime of 
annual burning (MUNM 1,2; Andersen, 1991b). Ant com- 
munity succession following severe disturbance also shows 
clear patterns in relation to stress, as shown by ant com- 
munity development following revegetation of waste rock 
at Ranger uranium mine in the seasonal tropics (Andersen, 
1993a). A DD4 community (CR; RANG 1) is the first to 
develop, which is replaced by DDOOPP communities (R; 
RANG 2-4) once the site becomes dominated by acacias. 
These become DD30PP communities (CR; RANG 5,6) 
when patches of bare ground occur, following a trajectory 
towards a DD3GM community (C; RANG 8), which is 
characteristic of undisturbed sites in the region. It should be 
noted that, in both the above cases, the structural responses 
of ant communities to disturbance differ from those of 
plants. In primary succession, for example, plant communi- 
ties begin as ruderal and pass directly to competitive (Ma- 
jer, 1989). 
Responses of ant and plant communities to 
the same stress 
As has just been mentioned, ant and plant communities can 
respond to disturbance in different ways. The same applies 
to stress. In terms of their analogous structural attributes 
there is often a poor correspondence between ant and plant 
community structural types at any particular site. That is, 
for example, DD3 ant communities are analogous to forests 
in terms of their structural attributes, but they are not at all 
characteristic of forest habitats. Such a lack of correspon- 
dence occurs because the same stress is not equally import- 
ant to both plants and ants. For example, whereas low 
temperature is arguably the most critical stress for ant 
communities, low moisture availability is arguably the 
most critical stress determining plant community structure. 
Thus, the arid zone typically supports competitive ant 
communities (DD3GM, analogous to rain forests, as out- 
lined below) because of low low-temperature stress, but 
supports stress-tolerant plant communities (open shrub- 
lands, woodlands and hummock grasslands) because of 
high water stress (competition can be important for desert 
plants (Tilman, 1988), but the structure of desert vegetation 
is fundamentally determined by low moisture availability). 
There are cases, however, when ant and plant communi- 
ties appear to be structured by the same stress. This occurs 
at very cold sites, where low temperatures severely limit 
both ant and plant productivity. Such sites support 
DDO,1CS ant communities (i.e. Cold climate specialists are 
predominant Fig. 3a,b), which both occur in, and are 
analogous to, heathlands of cold climates (high latitudes 
and altitudes), where low temperatures prevent structural 
dominance by tall woody plants, and cold-adapted taxa 
predominate. 
A direct correspondence between ant and plant structural 
attributes at the same site can also occur when sunlight is 
a critical limiting factor, such as on the floor of tropical rain 
forests. Although tropical rainforests are competitive plant 
communities overall, the forest floor is a stress-tolerant 
environment for plants as it is for ants. Interestingly, in 
tropical rain forests Dominant dolichoderines are largely 
confined to the canopy, which is a competitive environ- 
ment. This applies to Philidris and Dolichoderus in Aus- 
tralia and SE Asia, and to Azteca and Dolichoderus in the 
New World (Majer, 1993). In tropical rain forests of the 
Old World, competitively dominant roles are played by 
species of Oecophylla and Crematogaster (Greenslade, 
1971; Room, 1971; Majer, 1976, 1990), which are again 
arboreal. 
The analogy between DD relative abundance and canopy 
cover breaks down at very high ( > 70%) values. A closed 
forest plant community can be structurally complex (such 
as in the humid tropics), because a closed canopy does not 
necessarily preclude the occurrence of other plant life- 
forms. However, by definition a 'closed forest' of DD 
implies that few other ant species occur, and that the 
community is structurally simple. As stated previously, 
DD4 communities appear to be restricted to highly dis- 
turbed habitats, and are analogous to plantations rather than 
to closed forests. The richest (often> 100 spp/ha), most 
productive and most structurally complex ant communities 
are DD3GM types), and these are the ant analogues of 
tropical rain forests. Just as tropical rain forests include a 
rich array of life-forms (e.g. liannes, epiphytes, parasites) 
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Classification of Australian ant communities 27 
adapted to co-exist with canopy trees, DD3GM ant com- 
munities typically include numerous species of Hot climate 
specialists whose morphological, physiological and be- 
havioural specializations are suggestive of a long history of 
interaction with competitively dominant ants. 
The future 
In conclusion, the classification proposed here identifies 
structural types of ant communities which vary predictably 
in response to stress and disturbance. This provides a 
general framework for the analysis of ant community struc- 
ture, integrating the roles of both biotic and abiotic factors 
(Dunson & Travis, 1991). It also has an important appli- 
cation in the use of ants as bio-indicators of environmental 
change (Andersen, 1990), which requires a predictive 
understanding of the response of ant communitystructure 
to environmental variables. 
Several important questions emerge from this study as 
themes for future research. First, how successfully does the 
analysis presented here describe ant community dynamics 
elsewhere? That is, how useful is the system as a basis for 
a global classification of ant communities? This will be 
addressed in a subsequent paper. Secondly, to what extent 
is there a relationship between ant community structure and 
the various roles of component species in relation to 
ecosystem function? Such a relationship between com- 
munity structure and ecosystem function is evident in 
plants: for example, rain forest trees characteristically pro- 
duce large, often fleshy fruits that support a diverse array of 
frugivorous birds and mammals (Willson, Irvine & Walsh, 
1990; French, O'Dowd & Lill, 1992; Osunkoya, 1994), 
which are absent from sclerophyll habitats. How does the 
distribution of the various functional roles of ants (e.g. 
granivory, seed-dispersal and other mutualisms) vary with 
ant community type? 
Finally, to what extent are continental or global 
classifications possible for communities of other animal 
groups? Considerable progress has been made in identify- 
ing environmental patterns of diversity in other faunal 
groups, including birds (Gentilli, 1992) and mammals 
(Smith, May & Harvey, 1994), but not of community 
structure. Vegetation classification is possible because a 
small number of external factors (relating to climate and 
soils) can be readily identified as fundamental determinants 
of plant community structure. The same is true for ants, 
where climate and vegetation structure are of prime import- 
ance. The success or otherwise of classification systems for 
other animal groups will probably depend on the 
identification of similarly small sets of factors that are of 
universal importance in community structure. 
ACKNOWLEDGMENTS 
This paper is a tribute to the pioneering work on ant 
functional groups by P .J. M. Greenslade. I thank S. P. 
Cover, D. Haig, J. D. Majer, G. Orians and B. H. Walker 
for their comments on the manuscript. This is TERC library 
contribution no. 857. 
REFERENCES 
Andersen, A.N. (1983a) A brief survey of ants in Glenaladale 
National Park, with particular reference to seed-harvesting. Vict. 
Nat. 100, 233-237. 
Andersen, A.N. (1983b) Species diversity and temporal distri- 
bution of ants in the semi-arid mallee region of northwestern 
Victoria. Aust. J. Ecol. 8, 127-137. 
Andersen, A.N. (1986a) Diversity, seasonality and community 
organization of ants at adjacent heath and woodland sites in 
southeastern Australia. Aust. J. Zool. 34, 53-64. 
Andersen, A.N. (1986b) Patterns of ant community organization in 
mesic southeastern Australia. Aust. J. Ecol. 11, 87-99. 
Andersen, A.N. (1988) Immediate and longer-term effects of fire 
on seed predation by ants in sclerophyllous vegetation of south- 
eastern Australia. Aust. J. Ecol. 13, 285-293. 
Andersen, A.N. (1990) The use of ant communities to evaluate 
change in Australian terrestrial ecosystems: a review and a 
recipe. Proc. Ecol. Soc. Aust. 16, 347-357. 
Andersen, A.N. (1991a) Parallels between ants and plants: impli- 
cations for community ecology. Ant-Plant interactions (ed. by 
C.R. Huxley and D.F. Cutler), pp. 539-558. Oxford University 
Press, Oxford. 
Andersen, A.N. (1991b) Sampling communities of ground-forag- 
ing ants: pitfall catches compared with quadrat counts in an 
Australian tropical savanna. Aust. J. Ecol. 16, 273-279. 
Andersen, A.N. (1991c) Responses of ground-foraging ant com- 
munities to three experimental fire regimes in a savanna forest 
of tropical Australia. Biotropica, 23, 575-585. 
Andersen, A.N. (1992) Regulation of 'momentary' diversity by 
dominant species in exceptionally rich ant communities of the 
Australian seasonal tropics. Am. Nat. 140, 401-420. 
Andersen, A.N. (1993a) Ants as indicators of restoration success 
at a uranium mine in tropical Australia. Restoration Ecol., 1, 
156-167. 
Andersen, A.N. (1993b) Ant communities of the Gulf region of 
Australia's semi-arid tropics: species composition, patterns of 
organization, and biogeography. Aust. J. Zool. 41, 399-414. 
Andersen, A.N., Blum, M.S. & Jones, T.M. (1991) Venom alka- 
loids in Monomorium 'rothsteini' Forel repel other ants: is this 
the secret to success by Monomorium in Australian ant com- 
munities? Oecologia, 88, 157-160. 
Andersen, A.N. & Burbidge, A.H. (1991) The ants of a vine 
thicket near Broome: a comparison with the northwest Kimber- 
ley. J. Roy. Soc. W.A. 73, 79-82. 
Andersen, A.N. & Burbidge, A.H. (1992) An overview of the ant 
fauna of Cape Arid National Park, W.A. J. Roy. Soc. W.A. 75, 
41-46. 
Andersen, A.N. & McKaige, M.E. (1987) Ant communities at 
Rotamah Island, Victoria, with particular reference to disturb- 
ance and Rhytidoponera tasmaniensis. Proc. R. Soc. Vict. 99, 
141-146. 
Andersen, A.N. & Majer, J.D. (1991) The structure and biogeogra- 
phy of rainforest ant communities in the Kimberley region of 
northwestern Australia. Kimberley rainforests of Australia (ed 
by N.L. McKenzie, R.B. Johnston and P.J. Kendrick.), pp. 
333-346. Surrey Beattie & Sons, Sydney. 
Andersen, A.N., Myers, B.A. & Buckingham, K.M. (1991) The 
ant fauna of a mallee outlier near Melton, Victoria. Proc. Roy. 
Soc. Vic. 103, 1-6. 
Andersen, A.N. & Patel, A.D. (1994) Meat ants as dominant 
members of Australian ant communities: an experimental test of 
their influence on foraging success and forager abundance of 
other species. Oecologia, 98, 15-24. 
Andersen, A.N. & Reichel, H. (1994) The ant (Hymenoptera: 
Formicidae) fauna of Holmes Jungle, a rainforest patch in the 
? 1995 Blackwell Science Ltd, Journal of Biogeography, 22, 15-29. 
This content downloaded from 134.225.1.226 on Sat, 3 Jan 2015 12:20:01 PM
All use subject to JSTOR Terms and Conditions
http://www.jstor.org/page/info/about/policies/terms.jsp
28 Alan N. Andersen 
seasonal tropics of Australia's Northern Territory. J. Aust. Ent. 
Soc. 3, 153-158. 
Andersen, A.N. & Yen, A.L. (1985) Immediate effects of fire on 
ants in the semi-arid mallee region of northwestern Victoria. 
Aust. J. Ecol. 10, 25-30. 
Box, E.O. (1981) Macroclimate and Plant Forms; An Introduction 
to Predictive Modelling in Phytogeography. Junk Press, The 
Hague. 
Briese, D.T. & Macauley, B.J. (1977) Physical structure of an ant 
community in semi-arid Australia. Aust. J. Ecol. 2, 107-120. 
Brooks, D.R. & McLennan, D.A. (1991) Phylogeny, ecology and 
behaviour: a research program in comparative biology. Univer- 
sity of Chicago Press, Chicago. 
Christian, K.D. & Morton, S.R. (1992) Extreme thermophilia in a 
Central Australian ant, Melophorus bagoti. Physiol. Zool. 65, 
885-905. 
Dunson, W.A. & Travis, J. (1991) The role of abiotic factors in 
community organization. Am. Nat. 138, 1067-1091. 
French, K., O'Dowd, D.J. & Lill, A. (1992) Fruit removal of 
Coprosma qudrifida (Rubiaceae) by birds in south-eastern Aus- 
tralia. Aust. J. Ecol. 17, 35-42. 
Gentilli, J. (1992) Numerical clines and escarpments in the geo- 
graphical occurrence of avian species; and a search for relevant 
environmental factors. Emu, 92, 129-140. 
Greenslade, P.J.M. (1971) Interspecific competition and frequency 
changes among ants in Solomon Islands coconut plantations. J. 
appl. Ecol. 8, 323-352. 
Greenslade, P.J.M. (1976) The meat ant Iridomyrmex purpureus 
(Hymenoptera: Formicidae) as a dominant member of ant com- 
munities. J. Aust. Ent. Soc. 15, 237-240. 
Greenslade, P.J.M. (1978) Ants. The physical and biological 
features of Kunoth Paddock in Central Australia (ed. by W.A. 
Low.), pp. 109-113. CSIRO Division of Land Resources Tech- 
nical Paper No. 4, Canberra. 
Greenslade, P.J.M. (1979) A Guide to ants of South Australia, p. 
44. South Australian Museum, Adelaide. 
Greenslade, P.J.M. (1983) Adversity selection and the habitat 
templet. Am. Nat. 122, 352-365. 
Greenslade, P.J.M. (1985) Some effects of season and geographi-cal aspect on ants (Hymenoptera: Formicidae) in the Mt Lofty 
Ranges, South Australia. Trans. Roy. Soc. S.A. 109, 17-23. 
Greenslade, P.J.M. & Halliday, R.B. (1983) Colony dispersion and 
relationships of meat ants Iridomyrmex purpureus and allies in 
an arid locality in South Australia. Ins. Soc. 30, 82-99. 
Grime, J.P. (1977) Evidence for the existence of three primary 
strategies in plants and its relevance to ecological and evol- 
utionary theory. Am. Nat. 111, 1169-1194. 
Grime, J.P. (1979) Plant strategies and vegetation processes. John 
Wiley, Chichester. 
Grime, J.P. & Hodgson, J.G. (1987) Botanical contributions to 
contemporary ecological theory. New Phytologist, 88 (Sup- 
plement), 283-295. 
Harper, J.L. (1977) Population biology of plants. Academic Press, 
New York. 
Harper, J.L. (1981) The concept of population in modular organ- 
isms. Theoretical ecology, 2nd edn (ed. by R.H. May), pp. 
53-77. Blackwell Scientific Publications, Oxford. 
Harper, J.L. (1985) Modules, branches, and the capture of re- 
sources. Population biology and the evolution of clonal organ- 
isms (ed. by J.B.C. Jackson, L.W. Buss and R.E. Cook), pp. 
1-33. Yale University Press, New Haven. 
Haskins, C.P. & Haskins, E.F. (1988) Final observations on 
Pheidole megacephala and Iridomyrmex humilis in Bermuda. 
Psyche, 95, 177-184. 
Hodgson, J.G. (1993) Commonness and rarity in British but- 
terflies. J. appl. Ecol. 30, 407-427. 
Holldobler, B. & Wilson, E.O. (1990) The ants. Belknap Press, 
Cambridge, Mass. 
Latham, R.E. (1992) Co-occurring tree species change rank in 
seedling performance with resources varied experimentally. E- 
cology, 73, 2129-2144. 
Loehle, C. (1988) Problems with the triangular model for repre- 
senting plant strategies. Ecology, 69, 284-286. 
MacArthur, R.H. & Wilson, E.O. (1967) The theory of island 
biogeography. Princeton University Press, Princeton, New 
Jersey. 
Majer, J.D. (1976) The maintenance of the ant mosaic in Ghana 
cocoa farms. J. appl. Ecol. 13, 123-144. 
Majer, J.D. (1977) Preliminary survey of the epigaeic invertebrate 
fauna with particular reference to ants, in areas of different land 
use at Dwellingup, Western Australia. For. Ecol. Mngmnt, 1, 
327-334. 
Majer, J.D. (1985a) Seasonality of ants (Formicidae) in southwest- 
ern Australia. Soil and litter invertebrates of Australian Med- 
iterranean-type ecosystems (ed. by P. Greenslade and J.D. 
Majer), pp. 23-30. WAIT School of Biology Bulletin Number 
12, Perth, W.A. 
Majer, J.D. (1985b) Recolonization by ants of rehabilitated min- 
eral sand mines on North Stradbroke Island, Queensland, with 
particular reference to seed removal. Aust. J. Ecol. 10, 31-48. 
Majer, J.D. (1989) Long-term colonization of fauna in reclaimed 
land. Animals in primary succession-the role of fauna in 
reclaimed lands. (ed. by J.D. Majer), pp. 143-178. Cambridge 
University Press, Cambridge. 
Majer, J.D. (1990) The abundance and diversity of arboreal ants in 
northern Australia. Biotropica, 22, 191-199. 
Majer, J.D. (1993) Comparisons of the arboreal ant mosaic in 
Ghana, Brazil, Papua New Guinea and Australia-its structure 
and influence on arthropod diversity. Hymenoptera and biodi- 
versity (ed. by J. LaSalle and I.D. Gauld), pp 115-141. CAB 
International, Wallingford, UK. 
Majer, J.D., Sartori, M., Stone, R. & Perriman, W.S. (1982) 
Recolonisation by ants and other invertebrates in rehabilitated 
mineral sand mines near Eneabba, Western Australia. Reclama- 
tion Revegetation Res. 1, 63-81. 
Midgley, G. (1993) Grime's triangle: synthesis, senseless or sign- 
post? Bull. S.Afr. Inst. Ecol. 12, 15-18. 
Noble, I.R. & Slatyer, R.O. (1980) The use of vital attributes to 
predict successional changes in plant communities subject to 
recurrent disturbances. Vegetatio, 43, 5-21. 
O'Dowd, D.J. (1985) Seasonal patterns in the activity of ants in 
Belair Recreation Park, South Australia. Soil and litter inverte- 
brates of Australian Mediterranean-type ecosystems (ed. by P. 
Greenslade and J.D. Majer), pp 70-72. WAIT School of Bi- 
ology Bulletin Number 12, Perth, W.A. 
Oksanen, L. (1993) Plant strategies and environmental stress: a 
dialectic approach. Plant adaptation to environmental stress 
(ed. by L. Fowden, T.A. Mansfield and J.L. Stoddart), pp 
313-333. Chapman & Hall, London. 
Osunkoya, O.O. (1994) Postdispersal survivorship of north 
Queensland rainforest seeds and fruits: effects of forest, habitat 
and species. Aust. J. Ecol. 19, 52-64. 
Porter, S.D. & Savignano, D.A. (1990) Invasion of polygyne fire 
ants decimates native ants and disrupts arthropod community. 
Ecology, 71, 2095-2106. 
Raunkier, C. (1934) The life-forms of plants and statistical plant 
geography. Clarendon Press, Oxford. 
Room, P.M. (1971) The relative distribution of ant species on 
Ghana's cocoa farms. J. anim. Ecol. 40, 735-751. 
Rossbach, M.H. & Majer, J.D. (1983) A preliminary survey of the 
ant fauna of the Darling Plateau and Swan Coastal Plain near 
Perth, Western Australia. J. Roy. Soc. W.A. 66, 85-90. 
? 1995 Blackwell Science Ltd, Journal of Biogeography, 22, 15-29. 
This content downloaded from 134.225.1.226 on Sat, 3 Jan 2015 12:20:01 PM
All use subject to JSTOR Terms and Conditions
http://www.jstor.org/page/info/about/policies/terms.jsp
Classification of Australian ant communities 29 
Smith, F.D.M., May, R.M. & Harvey, P.H. (1994) Geographical 
ranges of Australian mammals. J. anim. Ecol. 63, 441-450. 
Southwood, T.R.E. (1977) Habitat, the templet for ecological 
studies? J. anim. Ecol. 46, 337-365. 
Southwood, T.R.E. (1988) Tactics, strategies and templets. Oikos, 
52, 3-18. 
Spence, J.R. & Andersen, N.M. (1994) Biology of water striders: 
interactions between systematics and ecology. Ann. Rev. Ento- 
mol. 39, 101-128. 
Strong, D.R., Simberloff, D., Abele, L.G. & Thistle, A.B. (eds) 
(1984) Ecological communities: conceptual issues and the evi- 
dence. Princeton University Press, New Jersey. 
Terborgh, J. & Robinson, S. (1986) Guilds and their utility in 
ecology. Community ecology: patterns and processes (ed. by J. 
Kikkawa and D.J. Anderson), pp. 65-90. Blackwell Scientific 
Publications, Melbourne. 
Tilman, D. (1982) Resource competition and community structure. 
Princeton University Press, New Jersey. 
Tilman, D. (1988) Plant strategies and the dynamics and structure 
of plant communities. Princeton University Press, New Jersey. 
Werger, M.J.A., van der Aart, P.J.M., During, H.J. & Verhoeven, 
J.T.A. (eds) (1988) Plant form and vegetation structure: adap- 
tation, plasticity and relation to herbivory. SPB Academic 
publishing bv, The Hague. 
Willson, M.V., lrvine, A.K. & Walsh, N.G. (1990) Vertebrate 
dispersal syndromes in some Australian and New Zealand plant 
communities, with geographic comparisons. Biotropica, 21, 
133-147. 
Wilson, E.O. (1990) Success and dominance in ecosystems: the 
case of the social insects. Ecology Institute, Oldendorf/Luhe. 
Woodward, F.I. (1987) Climate and plant distribution. Cambridge 
University Press, Cambridge. 
APPENDIX 
Classification of taxa into functional groups. 
Dominant Dolichoderinae 
Anonychomyrma Donisthorpe, Iridomyrmex Mayr, Frog- 
gattella Forel, Papyrius Shattuck. 
Subordinate Camponotini 
Calomyrmex Emery, Camponotus Mayr, Opisthopsis 
Emery, Polyrhachis F. Smith. 
Hot climate specialists 
Adlerzia Forel, Anisopheidole Forel, Melophorus Lubbock, 
Meranoplus F. Smith, Monomorium Mayr (part), Ochetel- 
lus flavipes (Kirby). 
Cold climate specialists 
Dolichoderus Lund, Heteroponera Mayr, Monomorium 
Mayr (part), Myrmechorhynchus E. Andre, Notoncus 
Emery, Podomyrma F. Smith (part), Prolasius F. Smith, 
Stigmacros Forel. 
Tropical climate specialists 
Mayriella Forel, Monomorium Mayr (part), Oecophylla F. 
Smith, Pheidologeton Mayr, Podomyrma F. Smith (part), 
Solenopsis geminata (Fabricius), Tetraponera F. Smith. 
Cryptic species 
Acropyga Roger, Aenictus Shuckard, Amblyopone Erich- 
son, Bothriomyrmex Emery, Brachyponera lutea (Mayr) 
group, Discothyrea