Ecological patterns and biological invasions: Using regional species inventories in

macroecology

Marc W. Cadotte1*, Brad R. Murray2 and Jon Lovett-Doust3

1: Complex Systems Group, Ecology and Evolutionary Biology, 569 Dabney Hall,

University of Tennessee, Knoxville, TN, 37996-0001, USA.

2: Institute for Water and Environmental Resource Management, University of

Technology Sydney, Gore Hill, NSW 2065, Australia.

3: Department of Biological Sciences, University of Windsor, Windsor, ON, N9B 3P4,

Canada.

* Author for correspondence, [email protected]

Abstract

Macroecology depends heavily on a comparative methodology in order to identify

large-scale patterns and to test alternative hypotheses that might generate such

patterns. With the advent and accessibility of large electronic databases of species and

their life-history and ecological attributes, ecologists have begun seeking generalities,

and examining large-scale ecological hypotheses involving core themes of range,

abundance and diversity. For example, combinations of ecological, life history and

phylogenetic data have been analysed using large species sets to test hypotheses in

invasion biology. Analysis of regional species inventories can contribute cogently to our

understanding of invasions. Here we examine several ways in which database analysis

is effective. We review 19 studies of comparative invasions biology, each using > 100

species of plants in their analyses, and show that invader success is linked to seven

correlates: short life cycle, abiotic (mostly wind) dispersal, large native range size, non-

random taxonomic patterns (emphasizing certain families or orders), presence of clonal

organs, occupying disturbed habitats, and earlier time of introduction). These

phylogenetically influenced, comparative analyses using regional species inventories

are only just beginning and have much potential.

KEYWORDS: Database analyses, comparative studies, macroecology, plant invasions,

phylogenetic analyses, ecological generalities, predictability

 3
“Given that we must renounce our quest for case-by-case predictability, what should we

do? There is pattern in the data we possess on invasions. The next efforts in the study

of invasions should be self-consciously statistical... But for this we will need the raw

data on-line in computer databases accessible to all researchers.” Gilpin (1990 )

Introduction

A. S. Watt’s conclusion in his presidential address to the British Ecological

Society —“Pattern and Process in the Plant Community“ (1947)— continues to resonate

today, in the emergent field of invasions biology and in the larger emergence of

macroecology. Watt first showed how a plant community may be described from these

two points of view —for diagnosis and classification, and as a working mechanism. For

Watt, “the ultimate parts of the community are the individual plants, but a description of

it in terms of the characters of these units and their spatial relations to each other is

impractible at the individual level. It is, however, feasible in terms of aggregates of

individuals and of species which form different kinds of patches“. Later, another B.E.S.

president, John Harper (1982) suggested that it was only natural that early stages in the

emergence and growth of any science should consist of the description and ordering of

the material for study. The next stage is to search for correlations and possible causes

underpinning what has been described.

Harper (1982) showed how historically the field of plant ecology had been

dominated by two major themes: 1) description of vegetation (i.e., assemblages of

species that are treated as objects for classification or ordination; with an array of
 4
environmental features regarded, variously, either as drivers or simple correlates); and

2) autecological single-species descriptions (e.g., Biological Flora of the British Isles, in

Journal of Ecology, or the Biology of Canadian Weeds series in Canadian Journal of

Plant Sciences), intended to produce detailed treatments of individual species ---their

nomenclature, form, distribution, population ecology, response to environmental factors,

etc. Again the emphasis was on the species. As Harper (1982) put it: “These two broad

categories of description give ecologists the equivalent of the telephone directory and

’Yellow Pages’, the one describing who we can find where, and the other describing

who (plant or community) does what (and again where they can be found).“

Both kinds of investigation allow predictions to be made —about where we will

find particular vegetation or species, and about what sorts of behaviour, or morphology,

or life history we may expect to find among the species in a particular region. Harper

(1982) warned that, whereas almost inevitably ecological studies use the species as the

basis for description, it remains far from clear “that the conservative and stable

characters and the breeding isolation that may be used to define such taxa are

appropriate to define ecological units ---knowing as we do the wide range of ecologically

different behaviours that are included within single species“. Harper suggested that

questions be formulated in general as: What are the limitations in form and behaviour of

organisms that account for present distributions and behaviours? Today this still seems

an apt and useful guide.

If as Gilpin (1990) suggests (in the opening quotation), ecologists are unable to

predict individual ecological outcomes, then the focus must turn to uncovering broader
 5
patterns of ecological generality. Since the advent of the science of ecology (e.g.,

Clements 1905, Warming 1909) there have been only a limited number of ecological

generalities uncovered. Conspicuous examples include the competitive exclusion

principle (e.g., Grinnell 1925; Gause 1934; Hardin 1960), the relative abundance

distributions of rare and common species (Preston 1948); species-area and isolation

relationships (MacArthur and Wilson 1963); and see too Bell (2001) and Hubbell (2001)

on patterns in neutral macroecology. The paucity of ecological generalities appears to

be inversely related to the general extent of knowledge of any particular phenomenon,

and the complex multiplicative and often non-linear interactive nature of many

ecological processes. The difficulty is due, in part, to the absence of any standardized

method for critically testing and evaluating generalities. For example, does highlighting

and noting a limited number of examples of a phenomenon provide enough support for

a theoretical generality? Similarly, does determination of particular variants or

exceptions necessarily disprove an ecological generality? And of course, even when we

think we have identified a strong ecological generality, is causation necessarily implied?

At present, the major mechanism used to test ecological generalities is critical

review, and meta-analysis of previous published results. While these provide valuable

insight into ecological problems, they are subject to two potential problems: bias in the

selection of the study organism and difficulties due to censored data. The first, bias in

study species (see Bonnet et al. 2002), confounds many meta-analyses in ecology. For

example, has the ecology of sedges been studied to the same extent as that of lycopod

mosses or oak trees, or indeed of salamanders, etc.? Several reasons exist for this,
 6
including availability of funding for research on certain (charismatic, relevant,

serendipitous) organisms over others, emotional attachment, ease of study, etc.; see

Bonnet et al. 2002). Organisms and whole systems may be selected precisley because

they may lend themselves to clearly supporting or equally disproving a hypothesis; or

because they may easily lend themselves to being manipulated to answering the

important questions, or because of an a priori belief that these organisms or systems

exhibit a certain pattern. An example of this kind of bias is seen among studies

examining the effects of forest fragmentation, where there has been a significant bias

toward using birds as the organisms of study (Simberloff 2000). While fragmentation

studies have lead to some interesting and important results (e.g., Bierregaard et al.

1992; Trzcinski et al. 1999), reviews and meta-analyses examining effects of forest

fragmentation will be biased towards this unique and exceptionally mobile group of taxa.

The second potential bias, data censoring, is perhaps more insidious and difficult

to control. Primary papers —which form the basis for reviews and meta-analyses— are

the source of censored data. Journals (like all publications) need to sell themselves and

hold a market position (involving scientific standards, reputation, first-publication record

for important results, theories, etc.) and are probably biased towards the publishing of

substantial results over ‘non-results’ (Murtaugh 2002). Thus papers reporting, say, an

absence of any apparent differences between rare and common species, will probably

tend not to occupy the pages of top-ranked ecological journals.

If organisms may be selected in a biased manner, and reports preferentially

publish positive results, do reviews and meta-analyses have a role in the elucidation of
 7
ecological generalities? Of course they do, but as an intermediary step toward

developing it. We see reviews and meta-analyses as combining disparate sources of

information and results into workable hypotheses, which can then be more thoroughly

tested. An overall framework for testing ecological generalities is suggested in Fig. 1.

We do not argue that the vast array of ecological complexities can necessarily be

attributed to a small number of ecological generalities; rather, we see a structure by

which ecologists can test the relative importance of proposed generalities, and also the

importance of variation or departure from a potential generality. Recently Daehler

(2001) rightly noted that a number of floras needed to be analysed in order to

adequately test hypotheses of plant naturalization.

With the ability to maintain electronically and transmit large datasets, scientists

can analyse complete species datasets (usually maintained by government agencies

and conservation organizations (e.g., USGS invasive species, FishBase, IUCN

endangered species, etc.)). Difficulties include differing levels or extents of species

inventories, ranging from a few pre-selected species to complete species lists of a

region. We see four, broadly different levels of species compilation, available for

comparative analyses (Table 1). First is a complete (or near complete) inventory where

there has been a concerted effort to record all extant taxa within a large (socio-politically

delineated) region or jurisdiction (see also below, ‘Inventory’). Second is a large species

database, which does not attempt to enumerate all the species within an entire region

but is, rather, a complete list of species that occur in a large, representative habitat or

ecosystem. Third is a species list, which is a limited subset of species, selected on

 8
some a priori criteria (e.g., sampled from a particular habitat type or biome, or from an

individual taxonomic family). The final type is made up of pre-selected small groups of

species, such as a few pairs of closely-related species. These have received much of

the focus thus far, given their utility in controlling for phylogenetic effects, and the sense

of comparative value. In what follows, we emphasize analysis using the first two, less

studied levels: species inventories and large species databases.

Given that variables relevant to a hypothesis are available for analysis (e.g.,

abundance, longevity, body size, fecundity, growth rate, etc.), then a hypothesis (

suitablyrefined by theory and meta-analysis) can be tested, minimizing taxon-specific

bias. Such analyses of a complete inventory or large species database may then be

combined in order to determine the extent of dependence the results may have on

geography and taxon. The use of large species inventories and databases to test

hypotheses explaining large-scale patterns in ecology and as evolutionary biology has

led recently to important and diverse conclusions (macroecological studies: Kelly 1996;

Kelly and Woodward 1996; Westoby et al. 1996; Pyšek 1998; Hegde and Ellstrand

1999; Lovett-Doust and Kuntz 2001; Cadotte and Lovett-Doust 2001, 2002; Khedr et al.

2002; Murray et al. 2002; Lovett-Doust et al. 2003; evolutionary studies: Eriksson and

Bremer 1992; Ricklefs and Renner 1994; Dodd et al. 1999; Gaston and Blackburn 2000;

among inventories: Leishman et al. 1995). Most of these examples probably coincide

with the recent completion and availability of electronic inventories. Here we wish simply

to highlight the importance and value of using large databases for the discovery and
 9
validation of ecological generalities. We highlight certain limitations, which bear

recognition by ecologists.

Generalities in invasions biology: much potential for database analyses

By the middle of the past century, invasions by non-native species had begun to

be recognized as a significant biological phenomenon and an issue of general concern

(Elton 1958; and see reviews of early invasions in Mack 2000; Cadotte in press). Since

then, numerous examples of species thriving outside their native range have been

described, with authors inferring ‘rules’ or generalities of invasiveness (e.g., Baker 1974;

Rejmanek 1996). Although these synthetic attempts drew on results or examples from

only a limited number of studies and taxa, and were subject to the potential errors

described above, they did shape and direct our understanding of biological invasions.

Moreover there is no clear consensus concerning generalities and the capacity of

biologists to predict future invasions. Although recent attempts range from relative

agnosticism to sanguine confidence (examples from this spectrum, presented in a

general range from confident to pessimistic, include: Pheloung et al. 1999; Daehler and

Carino 2000; Reichard and Hamilton 1997; Kolar and Lodge 2001; Rejmanek and

Richardson 1996; Pimm 1989; Goodwin et al. 1999; Mack et al. 2000; Scott and

Panetta 1993; Williamson 1999), there remain relatively few analyses of large

databases probing characteristics of invasive species (but see Table 2). In a review of

16 invasion studies, each containing at least 20 species (though definitely not complete

species inventories), Kolar and Lodge (2001) showed that generalizations appeared to
 10
be emerging, at least among plants and birds. Kolar and Lodge also showed that this

type of investigation (using larger datasets) is only just commencing, with thirteen of the

studies occurring in the past ten years.

The inventory

For purposes of examining issues in invasions biology, an inventory of species

may include all of the species, both native and introduced, or just the exotics, depending

upon the questions being asked. A bare-bones species inventory is just that ---a list of

all (or nearly all) species in a given region. How big should the region be? This depends

upon the scale of the question, and could limit inferential power. To test potential

ecological generalities, the species list should be at a sufficient scale as to transcend

the influences of local environments, and individual species’ dispersal capabilities (to

remove effects that transient species may have on local assemblages), and the

vagaries of idiosyncratic and random (including historical) influences. Thus for example,

the species inventory for the island Krakatau may be interesting in its own right, but may

not be an effective example for studying invasions of intact communities. At the same

time the list cannot be so large as to transcend the regional and geographical forces

affecting species distributions and abundances. Global databases (e.g., IUCN red lists)

are very important in their own right, and extremely useful for conservation and

evolutionary analyses (e.g., Purvis et al. 2000), but may not answer ecological

questions without further regional information. For example, constructing a global list of

invaders would be difficult, not only because of definitional problems (e.g., Colautti and
 11
MacIsaac 2004), but also for historical and ecological ones. Not all species have equal

opportunity to expand to potentially new habitats, nor are invaders going to be

successful in every habitat they come across.

We prefer a large (>100,000 km2) political jurisdiction, for several reasons. First,

due to the structure of government agencies and their funding lines, political regions will

be a logical level of focus for the initial construction of species inventories, especially

considering governmental efforts to control invasive species; secondly, large political

regions are largely biogeographically arbitrary (excluding small islands). Such

arbitrariness means that there is no a priori reason for habitat-level structuring to be a

factor in the species inventory. Large political regions (e.g., some European and Central

American countries, and Canadian, Australian and large US provinces/states) will often

contain a number of different geological and biogeographical realms. Third, political

regions are repeatable. That is we can locate arbitrary regions of similar size, climate,

species number, etc, for comparative analyses. Finally, data collection at this level will

likely contain important ‘on the ground’ information, such as relative abundances and

area of occupation. Analyses of this type of database will compare species which that

are likely to interact with one another.

A species list on its own is not so very interesting. However once ancillary,

correlative evolutionary or ecological information is added, parsing the inventory can

become particularly fruitful. Any number of biologically meaningful variables may be

added, including but not limited to phylogenetic information, life history attributes,

habitat associations, relative abundances, invasion status, and so on. Unfortunately,

 12
there is no central repository for databases of this sort, where researchers add variables

or fill missing information (but see Peterson and Vieglais 2001), but with the blossoming

of bioinformatics as a relatively well-funded area of science, database development and

maintenance should continue to grow rapidly. We note the presence of several excellent

repositories of large databases ---USGS invasive species, FishBase, IUCN endangered

species. For example Ruesink (2003) used FishBase (www.fishbase.org) to show that

the majority of non-indigenous fish had ecological effects on the invaded communities,

and that these impacts were more severe in communities with high endemism and

longer amount of time since invasion in the community.

Database analyses

Large databases are amenable to many commonly used statistical analyses (e.g.

Sokal and Rohlf 1994, Zar 1998), especially correlation, regression, ANOVA (including

MANOVA and ANCOVA) and their nonparametric equivalents. However, it is crucial that

the data points be seen for what they are. One may probe the dataset using individual

species as independent datapoints, or by using a phylogenetically-informed analysis.

These two approaches can reveal quite different results (Harvey et al. 1995; Westoby et

al. 1995) and the researcher should be aware of what their results mean and the

potential limitations, or crucial assumptions attached to their analysis.

Incorporating phylogenetic information in database analysis

 13
Ecological generalities in invasions biology are often sought on the premise that

invasive species possess a distinctive suite of biological traits (e.g., Kolar and Lodge

2001). To test this premise, traits of invasive species have been compared with those of

native species in invaded habitats (Cadotte and Lovett-Doust 2001), and with those of

non-invasive species from the species pool in the source region (Prinzing et al. 2002),

as well as among non-native species of differing levels of ecological success (Cadotte

et al. 2004, in review). Such comparisons are possible where trait values for each

species contribute single units of information in statistical analyses (Fig. 2). These

cross-species comparisons are of consequence because they describe present-day

relationships among species (Grafen 1989; Price 1997; Ackerly and Donoghue 1998;

Westoby et al. 1998; Harvey and Rambaut 2000). Moreover value is given to the

present-day functionality of the traits of each species. Cross-species analyses suggest

that certain traits, such as clonal growth and hermaphroditism for instance, are

functionally important in promoting invasiveness among plants (Thompson et al. 1995;

Cadotte and Lovett-Doust 2001).

Cross-species comparisons have traditionally been conducted without explicit

consideration of phylogenetic relatedness among species. Coupling information on the

traits of extant species with phylogenetic information can detail the extent to which

change in one trait (e.g., invasiveness) may be correlated with change in another (e.g.,

seed mass) throughout a particular phylogeny (or correlated-divergence analysis sensu

Westoby et al. 1998).

 14
The importance of phylogenetically-structured comparisons is that each radiation

or node in a phylogeny, rather than each species, contributes a single piece of

information in the analysis (Ridley 1983; Felsenstein 1985; Grafen 1989; Harvey and

Pagel 1991; see Fig. 2). The database of trait values for extant species can be used to

calculate values at each node in a phylogeny (a procedure that has been referred to as

‘hanging a variable on a tree’, Grafen 1989). In the simplest case, values at nodes are

effectively calculated as differences in trait values between two species diverging from

the same node (Felsenstein 1985; see Martins and Hansen 1996 for discussion of more

complex situations). The important feature here is that values determined for nodes are

independent when they describe separate radiations in a tree. Hence values at nodes

can be used in place of trait values of species to perform correlated-divergence analysis

in a variety of statistical techniques, including correlation, regression, multiple

regression, principal components analysis, and analyses of variance and covariance

(Felsenstein 1985; Garland et al. 1999).

The utility of a complementary approach in database analysis, where both cross-

species and correlated-divergence analyses are performed, is that biologically

informative patterns may emerge in the comparison of the different analytical methods

(Ricklefs and Starck 1996; Price 1997; Brown 1999). For instance, there might be a

positive relationship between seed mass and invasiveness within genera or families.

Yet this pattern could be obscured in cross-species analysis because of large

differences in basic natural history between families or other major clades. The results

of cross-species analysis might reveal no relationship or even a negative relationship

 15
between invasiveness and seed mass (for a good example of this see Nee et al. 1991

for the relationship between body size and abundance in birds).

A number of techniques are available for formally incorporating phylogenetic

information into comparative analyses (Harvey and Pagel 1991; Martins and Hansen

1996). For two programs that can be used to perform correlated-divergence analysis,

packaged, powerful software is available as free downloads from the Internet. CAIC

(Comparative Analysis by Independent Contrasts) (Purvis and Rambaut 1995) is

available (at www. bio.ic.ac.uk/evolve/software/caic/), as is the phylogenetic regression

(Grafen 1989) (see users.ox.ac.uk/~grafen/phylo/), as at July 2004. For application of

CAIC see Prinzing et al. (2002), and for application of the phylogenetic regression see

Murray et al. (2002). Both these programs require a working phylogeny of the study

species (see Grafen 1989). A good starting point for construction of a working

phylogeny for plants is that of the Angiosperm Phylogeny Group (2003). The APG

(2003) provides a classification for orders and families of flowering plants. For further

phylogenetic resolution, consultation with systematists or additional literature searches

are required.

Database studies of plant invasions

There has been a recent spate of published articles using database analyses to

examine patterns and test hypotheses about plant invasions. Table 2 summarizes 19

such studies. Several patterns are evident. First, there is considerable variation in

database size and in the objects of comparison, with comparisons of natives vs. exotics
 16
(e.g., Crawley et al. 1996), both within particular groups of invaders (Daehler 1998), and

between invaders and non-invaders (Goodwin et al. 1999).

Relatively few studies have employed phylogenetic information in their analyses,

with most using species as independent data points (See Table 2). Many of these

studies test very different hypotheses (usually because the databases contain different

information). Though the database studies outlined in Table 2 are not readily amenable

to examination of how widespread certain generalities are (due to differing sizes and

types of information), they do show the great potential for this type of study. From Table

2 it is apparent that there may be widespread patterns emerging, though an adequate

test would need more standardized analyses (full species lists and incorporating

phylogenetic analyses). Seven possible generalities (patterns occurring in multiple

studies) are evident, relating the success of plant invaders to: short life cycle; abiotic

(mostly wind) dispersal syndrome; large native range size; non-random taxonomic

patterns (emphasizing certain families or orders); presence of clonal organs; occupation

of disturbed habitats; and early time-since-introduction.

Issues of concern

As long as researchers continue to collect life history and population-level data

(about mating systems, genetic variability, competitive interactions, etc.), the information

contained in inventories will continue to grow. We believe that ecologists and agencies

should strive to develop biotic inventories, which future researchers may build on (see

Grime et al. 1988). As long as these datasets remain ‘public’, that is they are accessible
 17
to other researchers to use or to add information to, these rich datasets would go a long

way toward answering questions about how general and pervasive certain patterns are.

Several cautions need particular mention.

Data collection

Individuals with differing degrees of training do database construction and

augmentation. Workshops and meetings devoted to methods of database construction

and analyses (e.g., a special session at the invasive species conference in Beijing, June

2004) attest to the reality that researchers are thinking about ways to make large

species databases more accurate. However, the analyst must be cognizant that these

databases are not peer-reviewed and that the overall accuracy of the data must be

assumed. Some of the aforementioned organized databases (e.g., FishBase) do have

rigorous methods of data inclusion. However the methods of construction and inclusion

for other databases are less clear. The Ontario plant database used by Cadotte and

others (Cadotte and Lovett-Doust 2001, 2002; Cadotte et al. in review) was compiled

from local botanical lists compiled by regional municipalities and the provincial

government, with more limited indication of overall rigour. Additional data (life-history

attributes, habitat associations, etc.) were compiled by the authors from literature

searches. How valid is the assumption that the data are accurate? This can only be

answered by systematic review of databases.

Further, these databases are delineated by political boundaries, and so likely do

not match geographical or biogeographical regions. Again, from the Ontario plant
 18
database, many rare Ontario species are relatively common in the USA. Abundance

scores recorded in this database are affected by this range truncation. How much this

effected the results of Cadotte and Lovett-Doust (2002) are not immediately obvious,

though an interesting approach to it would be to compare the edge-of-range rare

species with those having their range centers in Ontario.

Generality does not equal predictability

There have been calls for ecologists to do more to help policy-makers arrive at

informed environmental decisions (Shrader-Frechette 2001; Ehrlich 2002). This concern

is a prudent response to prevent the invasions of exotic species. In an article highly

critical of invasions studies, Shrader-Frechette (2001) argues that the major problem

associated with predicting invasive species “is primarily one of scientific method”, and

involves the caveat that “politics and economics play a role” (p. 507). She goes on to

say that this scientific inadequacy is three-pronged. First, there is no firm definition of

exotic species; second, current theory has brought no predictive power; and finally,

there are no empirical generalizations. Criticism such as this (but see also Ehrlich 2002)

contends that ecologists have not met the needs of the conservation community mainly

because ecologists have not been able to predict the outcome of certain events,

perhaps in a manner parallel to that of classical physics, or haven’t yet found the

mechanistic understanding that, say, geneticists have, with DNA.

True universal predictability in ecology may never be fully attainable, though

generalizations may be. Ecology is Complex. Many phenomena are time and place
 19
dependent and are likely to have very complex causes, influences and patterns

(McMahon and Cadotte 2002). Similarly, the ability to predict precisely which species

will have what kind of impact may not be possible (Williamson 1999), due to the

complex nature of species interactions with other species and their environments. If we

were to select a single strong predictor of invasive plant success, it would probably be

showy flowers, since most plant introductions are from horticulture (Mack and Lonsdale

2001; Lovett-Doust 2003). Of course this would have little biological meaning as to how

well a species performs in a new community, competes with other species, or utilizes

resources.

Ecology’s goal as an applied science need not be to predict the next purple

loosestrife or zebra mussel. Rather, we view ecology’s applied role as one of examining

patterns, discerning generalities and assessing potential consequences. Do we expect

climatologists to predict when and where the next hurricane will hit land? No, the

meteorological events giving rise to hurricanes are far too complex and stochastic to be

precisely predicted. But what these experts can do is identify high-risk locations and

determine the phenomena that may form a hurricane. Much the same should obtain in

invasions biology. The fact that ballast water of trans-oceanic ships has been the target

of legislation shows that scientists are identifying high risk activities.

Database analyses, by their statistical nature, will never be able to produce a

case-by-case predictive application. Rather, the best it can do is highlight trends and

generalities and produce statistical predictions, such as with discriminant analyses (e.g.,

Reichard and Hamilton 1997). In understanding statistical patterns in large data sets,
 20
ecologists gain a better understanding of how species interact with other species and

the abiotic environment.

Acknowledgements

We thank Jim Carlton, Petr Pyšek, Mark Westoby, Ian Wright and an anonymous

reviewer for helpful comments, Sean McMahon for useful discussions and the hurricane

analogy, and Jim Drake for suggesting we write this paper.

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 31
ble 1: Types of comparative analysis and their potential uses for analyzing species inventories.

Type of comparison Number of species¶ Potential utility Example

entory Complete species >80% of species in region. Determined in part by Cadotte and Lovett-
inventory of a large region. history, ecology, and Doust 2002
evolution.
tabase Large representative group 20-80% of species in a region, Likely to be Murray et al. 2002
of species. or 100% from sub-regions. phylogenetically random.
t Species list. <20% or 50-200 species. Not phylogenetically Eriksson and Bremer
random, selected with prior 1992
rationale (e.g., community
type)
oups Selected species. Few species, e.g., Not random; limited Rabinowitz and Rapp
comparisons among 20 inferential space. 1981
species in single genus.

umber is arbitrary.
 32
ble 2: Plant invasion studies using >100 species. ‘Data points’ refers to whether studies used species as independen

a points or followed some form of phylogenetic analyses/controls.

thors Dataset Type of Data points General findings

comparison

(from Table 1)

derson 1) 40 native species of disturbed List Species Dispersal strategies seem importa

95 habitats; 2) 61 aliens of disturbed (e.g., wind for invaders of natural

habitat; and 3) 52 aliens in natural habitats).

habitats in Denmark.
 33
adotte and 1814 species found in southwestern Inventory Species Taxonomically non-random and

vett-Doust Ontario, 484 of which are exotics. exotics were short-lived with longe

01 flowering periods, and small, wind

dispersed fruits.

adotte et al. 1153 non-native taxa found in Ontario. Inventory Phylogenetic Successful exotics were over-

04, in Comparisons involved common and represented by: arriving earlier, fr

view rare species. Europe and Eurasia, having an

organ of clonal growth, longer

flowering periods, and ability to

occupy variable soil moisture

regimes and disturbed habitats.

 34
awley et al. 1515 natives and 1169 aliens in the Inventory Phylogenetic Aliens were over-represented by

96 British Isles. heavier seeds, tall height and

protracted seed dormancy.

aehler 1998 1348 ‘serious’ agricultural weeds, 1041 Inventory Species Agricultural weeds were over-

widespread weeds, and 381 natural represented by rapidly reproducin

area invaders. abiotically dispersed herbs, while

natural area invaders were

represented by larger families.

aehler and 54 invasive and 57 non-invasive non- List Species Using Pheloung et al methodology

arino indigenous species in Hawaii they accurately identified >90% of

the invasive species.

 35
oodwin et 165 pairs of species native to Europe. List Species Most important trait appears to be

1999 First in pair is invasive to New geographic range size in Europe.

Brunswick, Canada and second is not.

nsdale 463 intentionally introduced species in Database Species <1% were found to be ‘useful’

94 northern Australia. without becoming problem specie

Also, grasses were more likely tha

legumes to be weedy.

eloung et 370 non-native species in Australia Database Species They created a list of factors

1999 and New Zealand. (biogeography, ecology,

reproduction, dispersal, etc.) whic

predicted >85% ‘weeds’ as being

concern.
 36
nzing et al. 183 invaders in Buenos Aires province Database Species and Four generalities contributing to

02 and 74 in Mendoza province, phylogenetic invader success: frequency in nat

Argentina. range; abiotic niche; native range

which covers multiple floristic

regions; and ruderal habit.

šek 1997 1) 457 aliens in Central Europe; 2) Multiple Phylogenetic Analysis of the influence of clonal

World’s most aggressive weeds (n = clonals appear more successful in

207); 3) Aliens of Auckland region (n = wetter, colder climates. Clonals ha

615); and 4) 64 natural area invaders dispersal disadvantage but an

in South Africa. advantage once established.

 37
šek 1998 26 regions around the world, with 57- Multiple Species* Most effective invaders belong in:

975 alien species per region. Papaveraceae, Chenopodiaceae,

Amaranthaceae, Cruciferae, and

Polygonaceae.

šek et al. 132 permanently established invasives List Species Aliens vs natives: Taxonomic, life

95 in Czech Republic compared to native form, life strategies, dispersal, and

flora. abiotic requirement differences.

šek et al. 668 species, non-indigenous to the Inventory Species Species adapted to disturbed

03 Czech Republic and introduced after habitats (e.g., early flowering time

1500. annual life cycle.) were more likely

have arrived earlier in history than

other species.
 38
eichard and 235 woody plants in North America. Database Species Used discriminant analysis to

amilton confirm weedy status. Traits such

97 species being related to an invade

and rapid vegetative reproduction

appear important.

ott and 242 established South African plants in List Species These ‘weeds’ seems to have wid

anetta 1993 Australia. native range size, are weeds

elsewhere, have been resident in

Australia for long time, and are

related to congeneric weeds.

 39
ébaud and 651 species in two datasets: 1) Database Species Species not taller in introduced

mberloff European species in California and ranges.

01 vice versa; and 2) European species in

the Carolinas and vice versa.

ompson et 211 species that have expanded List Species Invasives are more likely to be clo

1995 ranges in England, Scotland, the and polycarpic perennials.

Republic of Ireland and the

Netherlands vs natives.

lliamson Established aliens vs natives of Britain Inventory Species Factors important for invasives:

d Fitter (n = 1777). Native range size, size of plant, a

96 propagule pressure.

his study used species to compare taxonomic patterns of invasions

40
 41

LEGEND TO FIGURES

Fig. 1: Framework for using primary literature, meta-analyses, and complete species

inventories for testing hypotheses of general ecological patterns. Converging arrows

represent the consolidation of independent analyses; diverging arrows represent the

testing (via multiple independent tests) of the hypotheses, strengthened at the meta-

analytic level.

Fig. 2: A hypothetical phylogeny linking eight species, numbered 1 to 8. In cross-

species analysis, the trait values of each of the eight species are treated as

independent data points (n = 8). In a fully resolved phylogeny, there will be n – 1 nodes

or radiations; in this example there are 7 nodes, labeled A to G. Phylogenetically-

structured comparisons use values calculated at nodes as the independent data points,

as these values constitute phylogenetically independent contrasts.

 42

Primary studies

Primary reviews and meta-analyses

Test specific working hypotheses using

species invetories or databases

Secondary review or meta-analysis

Ecological generality: supported,

unsupported, and conditions

Fig. 1
 43
1
A
2
E
3
B
4
G

5
C
6
F
7
D
8

Fig. 2