The Diversity Succession of on INCO · 2005. 2. 9. · INCO Ltd. of Copper Cliff, Ontario has generated over 450,000 tonnes of taiiings material contained in an area of roughiy 2,225

The Diversity and Succession of Wandering Spider Communities on INCO Ltd.

Reclaimed Tailings Habitats

David Peter Shorthouse

THESIS SUBMITTED M PARTIAL RnFILLMENT OF THE REQUTREMENTS

FOR THE DEGREE OF MASTER OF SCIENCE IN BIOLOGY

SCHOOL OF GRADUATE STUDIES

LAURENTIAN UNIVERSITY

SUDBURY, ONTARIO

1998

O David Peter Shorthouse, 1998

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Canada

Master of Science

S io logy

1998

TITLE:

NAME:

SUPERVISOR:

NCTMBER OF PAGES:

Laurentian University

Department of Biology

Sudbury, Ontario

The Diversity and Succession of

Wandering Spider Cornmunit ies on INCO

Ltd. Rechimed Tailings Habitats

David Peter Shorthouse

Y. Alarie, Ph D.

146

ABSTRACT:

INCO Ltd. of Copper Cliff, Ontario has generated over 450,000 tonnes of taiiings

material contained in an area of roughiy 2,225 ha. An additionai 270,000 tonnes of

deposition is anticipated within 30 yean. INCO Ltd. taiiings are fhely ground, waste

bedrock generated fiom the milling process that contains little to no nutrients but elevated

levels of sulfide-complexed heavy metals (Cu, Ni, Fe). The naturally occurring oxidation

of these metal sulfdes in the presence of rainwater leads to the acidification of ground

and subsurface waters, a process known as "ac id-rnine drainage". Mining companies are

legall y bund to reclaim disturbed land to the best approximation of what was present

before their activities begaa iNCO Ltd. is in the process of ameliorating their disturbed

property while attempting to eliminate funue legacies of acid-generat hg land. The

technique employed coasists of spreading massive amounts of lime, fertilizer and straw

directly on top of the tailings subsaate. M e r the tailings pH is deemed adequate for plant

growth, various species of grasses and herbs are seeded. FoUowing a growth period of

approxhately 5 yeas, tree species are planted. It is hoped that the eventual formation of

soi1 and a thick humus mat, will intercept much of the rainwater and reduce oxygen

diffusion, thus reducing the level of sulfuric acid formation and heavy metal runoff. It is

also hoped that prospective habitats will support a biotic community in concert with their

relatively undisturbed surroundings. The INCO Ltd. tailings sites fonn of an

anthro pogenic gradient fiom bue, unrec laimed tailings ( year O), through grassland

habitats (5 years poa-reciamation), to a rnixed-wood habitat with a rich understory of

grasses and herbs (1 5 years post-reclarnation), ending in a monoculture of conifers with

little understory (30 years pst-reclarnation). It is assumed that animal life also follows a

similar successional pattern.

In order to assess the biological success and sustainability of the reclamation

process, a guild of ground-dweliing spiders (Araneae) was inventoried. This %andering

spider" guild as it is called (Uetz 1 979), is composed of the abundant cursorial families,

Lycosidae, Gnaphosidae, Clubionidae, Philodromidae, Thomisidae, and some members

of the families Hahniidae and Agelenidae. These spiders share a similar habitus because

of their body shape and size and mode of prey capture. They are ideal ecological

indicat ors because of their a bundance (do wing for diversit y assessments), because the y

are short-lived (thus adjusting more rapidly to changes in the environment), and because

their taxonomy is well understood (assuring correct identification) (Clausen 1 986, Allred

1975). One disadvantage of the taxonomy of this group however. is the difficulty in

identifying immature specirnens because species are identified based on the examination

of omate sexual organs present ody in the adult lifestage.

Wandering spiders were pitfàll trapped on each of the four tailings habitat types

throughout the surnmer of 1 996. The diversity of these spider communities was compared

to the diversity of communities on control sites located within the Sudbury region,

assumed to represent healthy, or equüibrium States. Three of the four control sites were

chosen to reflect the h c e area, physiognomy, and age of three reclaimed tailings

habitats. This was accomplished by selecting sites that had natually recovered fiom

forest fire disturbance. A fourth control site was chosen as a representative of Sudbury's

widespread and unique Birch Transiîion ecotone (Amiro and Courtin 198 1 ).

Based on determinations of general habitat preferences of ten of the moa

abundant species, a qualitative hunistic assessrnent alluded to potential physical and/or

biologicd constraints associated with MC0 Ltd. tailings sites. These potential constraints

may help to explain varying abundances of species such as Gnuphosa p m f a Banks,

Zelotesfia~ris Chamberlin, and Purdosa moesta Banks between tailings and control sites.

The a (alpha) diversity, of wandering spider cornmunities was compared via

abundance-based species richness estimations and Shannon- Wiener indices. The results

fkom these analyses revealed that there were more species on the control sites than on the

tailings sites and that some tailings sites were not as diverse as their control site

counterparts.

The P (beta) diversity, or species turn-over, of wandering spiders on tailings and

control sites was examined using the Morisita-Hom and Bray-Curt is comrnunity

similarity coe flic ients. These revealed that the wandering spider cornmunit ies on the two

oldea, hypothesized to be the healthiest, reclaimed tailings sites were moa similar to

control sites. These meanires also demonstrated that early successio na1 reclaimed tailings

sites were somewhat more similar to older, later successional reclaimed tailings sites.

This suggested an avenue of wandering spider arriva1 and establishment. In addition, a

dendogram constnicted with clusters of Bray-Curtis coefficients illustrated that barren,

unreclaimed tailings supported a community of spiders dissimilar to aii other

communities examined.

Al1 of the wandering spider communities on control sites were deemed mature

and varied naturai communities, significantly fating the tnuicated log n o 4 model. The

bare tailings and 30-year-old tailings cornmunit ies on the other hand, significantly fit the

geometric model suggeaing that these sites represent either stressed ecosystems or

ecosystems in the early stages of succession.

The results obtained in this study add to the knowledge of wandering spider

diversity in the Sudbury region as well as provide insight into the health of INCO Ltd.

reclaimed tailings habitats. These results should be heeded should INCO Ltd. wish to

undertake responsible mine c losure.

iii

This thesis is dedicated to my wife, Sara Shorthouse (née Marcantognini)? for her

patience and encouragement.

ACKNO WLEDGMENTS

I am grateful to my supervisor, Dr. Yves Alarie, for providing his tirne and

energy. 1 am also gratefùl to rny comminee members. Dr. Gerard Courtin and Dr.

Giuseppe Bagatto, and to my extemal examiner, Dr. Henri Goulet, for their exacting

evaluations, Dr. Goulet is a research scient ist with the Eastern Cereal and Gilseed

Research Centre (ECORC), a Research Branch of Agriculture and Agri-Food Canada in

Ottawa, Ontario. I am especially indebted to Dr. Bagatto for his field assistance and his

extensive stat ist ical h o wledge.

1 am gratefbl of CO-workers who have assisted me with field-work. These are, in

no particular order, Sara S horthouse, Daniel Paquette, C d e y Tissington, Monica Turk,

and Liane Dumas. Without their help, this thesis would not have k e n possible.

1 am also inde bted to my parents for their support. 1 have been forninate to have a

father as a faculty member of Laurentian University's Biology Department. He has

provided support at both ernotional and professional levels; somethuig unique in any

institution.

1 am especially thankful of Sara who has endured my late nights with

understanding. She has been a fantastic field assistant and continues to be my best fiend.

Financial support for this project was provided by grants &om INCO Ltd. Support

for my education included a Laurentian University bursary, an Ornario Graduate

Scholarship, and two Graduate Teaching Assistantships.

TABLE OF CONTENTS

PAGE

Abstract

Dedication

Acknowledgments

Table of Contents

List of Figures

List of Tables

1. General Introduction

II. An Introduction to MC0 Ltd. Tailings

III. An Introduction to Spider Biology

A. The Classification of Spiders

B. Spider Anatomy

C. Anatomy as it Relates to Classification

D. Wandering Spider Ecology

IV. Site Descriptions

A. Study Locale

B. Study Sites

i) INCO Ltd. Tailings Sites

iii Control Sites

V. General Materials and Methods

M. Faunisic Assessrnent of Wandering Spiders on MC0

Ltd. Tdings and Control Sites

A. Introduction

B. Methods

C. Redts and Discussion

1

iv

v

vi

viii

X

W. Wandering Spider a and P Diversity and Community

Modeling on MC0 Ltd. Tailings and Control Sites

A. Introduction

B. Materials and Methods

C. Results

1 . Species Richness

a. Control Sites

b. Tailings Sites

2. Abundance-based Coverage Estimations (ACE)

a. Control Sites

b. Tailings Sites

3. Shannon- Wiener Diversity (H') and Evenness(E)

a. Control Sites

b. Tailings Sites

4. Morisita-Horn (CmH) and Bray-Curtis (BC)

Communit y Similarity

5. Abundance Modeis

a, Control Sites

b. Tailings Sites

D. Discussion

VIII . General Discussion

IX. Appendices

X. Literature Cited

vii

LIST OF FIGURES

FIGURE PAGE

Clarabeiie Mill, INCO Ltd.

Polyethylene tailings pipes.

Dry ing and oxidizing tailing S.

C ladogram of the arachnid orders.

Idkaorder Araneomorphae cladogram

Anatomy of a generalized spider.

Spider eye arrangement.

Simple male pedipalp.

Complex male pedipalp.

Female reproductive system

Localities of the study sites.

Localities of INCO Ltd. tailings study sites.

Tailings Site A.

Tailings Site B.

Tailings Site C .

Tailings Site D.

Control Site 1 .

ControI Site 2.

Control Site 3.

Control Site 4.

Collection locallies of Gnuphosu parvula Banks.

. . . of Zelotesfratris Chamberlin.

. . . of Hognu fiondicola (Emerton).

. . . of Pardosu distincta (B lackwall).

. . . of Pardosa modica (Blackwall).

. . . of Pardosa moesta Banks.

FIGURE PAGE

. . . of Pirata minutus Emerton.

. . . of Schizocosa sa1ta~i.x (Hentz) .

. . . of Trochosa terricola Thorell.

H ypo thetical raddabundance p Io t . ACE and ICE representation.

Species richness on controls and tailings.

ACE for spiders on controls and tailings.

Jack-knifed Shannon-Wiener (H') diversity.

Jack-knifed Shannon- Wiener Evemess @).

Bray-Curtis dendogram.

Species abundance distributions on controls.

S pecies abundance distribut ions on tailing S.

LIST OF TABLES

TABLE

Latitude and longitude of study sites.

Total wandering spider catches on aii sites.

Biogeographical support for species presence.

Numbers of species found in published reports.

Additional species not collected.

Morisita-Hom (Cd) community similarities.

Chi-square tests for control abundance models.

Chi-square tests for taïlings abundance rnodels.

PAGE

1. GENERAL INTRODUCTION

As Gunn (1 995) aptly points out, Sudbury is well known as a poiluted region. It is

one of the largest point sources of sulfur dioxide emissions, contains approxirnately

1 7,000 ha of industrial barrens, over 7,000 acid-damaged lakes, and many other aartliog

examples of industrial devastation. The goal is to mine and refine copper and nickel

( including other metals of lesser concentration) and the environmental repercussions have

long been overlooked. The symbol of devastation has k e n the 381 -rn tail "Superstack".

However, for Little more than thirty years, Sudburians have been quashing Sudbury' s

negative image by expending enormous effort and resources directed at "re-greening"

their backyards. Sudbury's image has taken an about- face. Nevertheless, concealed

within the mining giant's grounds is a poignant sight that could threaten rising spirits.

Prior to the 19503, the townsfolk of Copper Cliff, the center of the INCO Ltd.

refinery, complained of air-borne dust. This fine particulate material was wepi up with

the prevailing winds, wafted over steep embankments, and filled the air of nearby streets.

The dust's source could be traced to massive containment areas for MC0 Ltd. tailings:

the ground, waste-rock fi om their mining process. The sought-after copper and nickel are

in concentrations of 2 6 % of deeply buried bedrock. The initial extracthg process

involves milling the rock into liquid slwries whereby the met& are decanted off the top

and the wastes are pumped to drained lakebeds or natural depressions in the ground.

Upon drying, this h e l y grouml waste was fiee to be picked up by winds.

Shortiy d e r 1957, the early pioneering efforts of agriculhirist Tom Peters led to

the creation of a vegetative cap on the tailings surface, thus damping the potential for air-

borne dust (Peters 1995). It was later discovered that water percolating through bare

tailings contained heavy metals and an acidic pH, providing M e r incentive to 'te-

vegetate." It was hypothesized that a self-sustajning ecosystem on top of their wastes

would not only reduce the amount of blowing tailings dust but would also intercept

rainwater. It was also hoped that the legacy could be bwied by a living system in concert

with its Unmediate surroundings. Since tailings continue to be produced through ongoing

extractions, the agricultural efforts resuh in a human-made, or anthro pogenic, succession

of life. This thesis is one of few studies occupied with INCO Ltd. tailings ecosystems and

great attention will be devoted to their description.

Any introductory ecology textbook will detine succession and will include several

illustrative examples. In contemporary usage, the term succession refers to a sequence of

changes in the species cornposit ion of a cornmdty, which is supposed to be associated

with a sequence of changes in its structural and functional properties. The fhdamental

property assigned to succession is that the changes are progressive or directionai, and it is

possible to predict which species will replace others in the course of a succession. The

succession of life on INCO Ltd. tailings is certainly directional because an outcome for

its development has been determined a priori. Under natural conditions however, an

endpoint is often too dmcult to hypothesize, and some would argue that endpointsper se

are only found in the imaginations of ecologists. Nevertheless, the succession of Me on

reckimed tailings sites is Wrely comparable to that following a natural disturbance such

as flooding or forest f ~ e . There are certainly important functional dflerences between the

two types of events, but the structural components bear remarkable similarity. If one asks,

"Just how similar are the successive communities of life on the tailings to life following a

forest fire event," an introductory eco logy textboo k will not provide any guidance.

In acniality, there is an abundance of methods to assist in analyzing succession.

These measures are associated with three main subdivisions of diversity: alpha (a), beta

(P) and gamma (y) (Whittaker 1 972). Alpha diversity and gamma diversity pertain to the

number of taxa at local and regional spatial scales, respectively, and their units are

number of species or other suitable measure of diversity. Beta diversity measures the

turnover of species between local areas and is usuaily treated as a dimensionless constant.

Within these subdivisions are both qualitative and quantitative meamres. if the biology of

a test group is well known, successional communities may be chanicterized based on

species composition alone. When two or more sites are compared however, mathematics

becomes an essential tool. The next moa taxing question Zone is charged with makuig a

cornparison between tailings communities and naturally successional communities is,

"What group of organisms would be a good qualitative and quantitative hdicator?"

Certain species signal changes in biotic or abiotic conditions. It is important to

inventory these sensitive species because they can becorne the important moniton.

Indicator species reflect the quality of environmental conditions and aspects of

community composition. Useful attributes of indicator species that are applicable to a

wide range of organisms in a variety of ecosystems include (modified fiom Pearson 1995

and Brown 1991 ):

high taxonornic and ecological diversity;

close association with, and identification of the conditions and responses o c

other species;

relat ively high abundance and damped fluctuations (Le. they are always

present and easy to locate in the field);

m o w endemism or, if widespread, well differentiated (either locally or

regionally) ;

we il known taxonomy and easy identification;

good background information;

predictable, rapid, sensitive, analyzable and linear response to disturbance.

1 rnight also add that, fiom a management and conservation point of view, an

indicator species shoukl also be one diat decision-makers and the public can appreciate

and recognize.

Spiders make an ideal indicator group. Numerous workers have shown that

different environments have specific spider faunas. and in gradient analyses, the species

are not evenly or randomiy distributed. The general impression is that spider faunas give

a pattern similar to that of vascular plants (Curtis 1978, Allred 1 975). This is not the same

as saying the number of spider species fluctuates with the number of plant species. In

fact, the two variables do not correlate well but depend largely on the spatial structure

and microclimate of t he enviro nment (Greenstone 1984). Like plants, difkent spider

species have difEerent requirements. Spider fàunas on trees have been studied in relation

to Sa pollution by Gilbert (1971); they have k e n analyzed in rehtion to heavy met&

(Rabitsch 1995, Strojan 1978); and they have k e n intensively studied in relation to

succession (Crawford et al. 1995, Bultman and Uetz 1982, Bultman 1980, DufTey 1978,

Peck and Whitcomb 1 978, Huhta 197 1, Lowrie 1 948 and others reviewed by Uetz 1 99 1 ).

They have also k e n studied in industrial landscapes (Luczak 1987, 1984) and dong

pollution gradients (Koponen and Niemela 1995, 1993), and even on reclaimed strip

mines (Hawkins and Cross 1982). Spiders are also prevalent in our liteninue, folklore,

and applied sciences, making them apparent to the public.

A group of spiders, called the wandering spiders, are adapted to or specialized for

ninning and are particularly suited to the examination of INCO Ltd. successional habitats.

The individuals of this group share similar body shapes and feeding strategies, informally

defining their association as an ecological guild (Adams 1985). Although spiders are a

very visible and permanent part of any ecosystem, strangely, they are innequent ly studied

by eco log ist s assembling invertebrate inventories. Perhaps because of this, the y are rarely

included in undergraduate course work. A review of spider taxonorny and biology will be

included in this study to direct people desiring spcific information.

Wandering spiders wiU be used in this nudy to explore the health of MC0 Ltd.

reclaimed taiiings habitats. Ecosystem health is gauged by equilibrium resilience

(Perrings 1995). Resilience in the traditional sense is a measure of resistance to

disturbance and speed of r e m to an equilibrium state. Community and systems

ecologists contend that there exists a well-defmed relationship between fùnctional

diversity and ecosystem resilience (Perrings 1995). Furthemore, the diversity and

complexity of ecological systems cm be traced to a m l 1 set of biotic processes (Holling

1992). Control sites of similar age and physiognomy, also forming a successional

gradient, were chosen as standards or as an approximation to the equilibrium state. The

srnall set of biotic processes on N C O Ltd. tailings habitats, such as the succession of

wandering spider species, will be qualitatively and quantitatively examined. Qualitative

rneasures will include faunistic considerations and the quantitative meastues will include

species richness estimations and biodiversity indices, both at the level of a diversity. and

comrnunity similarity indices, at the level of B diversity. Beyond these diversity

classifications, wandering spider species abundance distributions will be examined via

community modeling . Assemblages of wandering spiders will be exarnined in their own right using

species richness estimations and biodiversity indices whereas the tailings ecosysterns in

general will be exarnined from an abstract and theoretical perspective using similanty

indices and species abundance distribution models.

This thesis was supported by MC0 Ltd. contracts awarded to Dr. Giuseppe

Bagatto and Dr. Joseph S h o ~ h o w of the Department of Bio logy, Laurentian University.

These researchers were charged with an initial examination of the health and

sustainability of MC0 Ltd. tailings habitats via a census of carabid ground beetle

(Coleoptera: Carabidae) biodiversity. Later contracts were awarded to undertake

vegetation surveys, heavy metal uptake studies, and a census of soi1 mite biodivenity.

This thesis marks the first census of wandering spiders on INCO Ltd. tailings. The

ult imate purpose of this study is to gauge the health and sustainability of these

rudimentary ecosysterns, to help guide futw land management decisions.

II. AN INTRODUCTION TO iNCO LTD. TAILINGS

The INCO Ltd. tailings storage area is the largest repository of tailings in Canada.

Approxirnately 2,225 ha of land retains approximately 450,000 tonnes of taihgs (Puro et

al. 1995). Tailings have been deposited at this site since the 1930's and deposition will

likely continue for another 30 years, resulting in an ultimate deposition of more than

720,000 tonnes (Puro et al. 1995). Storing huge amounts of tailïngs without darnage to

the local environment is a difficult task.

Copper and nickel ores in the Sudbury basin average only a few percent of base

metals by weight and must be miIled using a process called beneficiation This procedure,

carried out in the Clarabelle Mill (Figure l) , prepares and concentrates the ore by

removing wastes (gangue). The concentrates of the desired metals are sent to the smelter

for refining and the unwanted gangue is sent to the tailings impoundment areas up to 6

km away.

Tailings are pumped as aqueous slurries under high pressure in huge rubber-lined,

high-density polyethylene pipes (Figure 2). At enclosed disposal sites, the pipes are

spigotted to release the taiiings and water mixture. Excess water flows over the d a c e

and is returned to the rnilVsmeher area for purification while the fine tailings material

senles (Figure 3). In tirne, the surface becomes suficient ly dry and strong to support

agricu hural equipment for reclamat ion work.

The enclosed disposal sites consist of a series of basins that follow bedrock ridges

whereby naturally o c c ~ g rock outcrops are used as bunresses for the tailings.

Additional dams are c o n m a e d with local till material and are successively raised as the

basins fil1 wit h tailings. The dams maintain structural stability by allowing seepage.

INCO Ltd. tailings consist of two primary areas: one area was fiiled between 1936

and 1988 and the second area has been receiving l?esh tailings since 1986. A variety of

grasses. legumes and trees have been seeded or planted on the first area. The second area

is devoid of vegetation and is almost completely exposed with the exception of a thin

layer of chernicals for dust controL

Figure 1 . The INCO Ltd. Carabelle Mill located on the northeast side of the property. The

input material is mined ore and the output material is base metal concentrate (transported

to the smelter) and tailings.

Figure 2. Rubber-lined polyethylene pipes transponing tailings under pressure from the

Clarabelle Mill to the tailings containment area, approxirnately 6 km away.

Figure 3. Tailings disposal site beguining to dry and oxidize. Excess water was retumed

to the mil1 and smelter for purification and reuse.

The objectives for reclaiming tailings are to reduce wind and water erosion,

reduce acid and heavy metal leaching, and to develop ecosystems in iwmony with their

surroundings. However, tailings are difficult materials on which to establish vegetation.

A combination of physicai, chernical, and biological constraints act in concert to fb tmte

plant growth. The primary physical constraint is soil compaction resulting fitom the small

and d o m particle size of tailings materials. Secondary sources of repression include

stratification during deposition and unfàvourable porosity, aeration, water filtration and

percolation properties. The suite of problems is exacerbated by the presence of iron

pyrites (FeS2) not removed during ore beneficiation, which leads to acid-generating

waste. Pyrite- bearing wastes c m oxidize within months to produce extreme acidit y. This

degradation is assisted by ferrous ion oxidizing bacte& ïWobucillus ferrooxidans,

which thnve at pH 1.5 to 3.0. Biological constraints to plant growth on tailings include

low concentrations of essential nutrients, nitrogen and phosphorus, b i t e d water holding

capacity, no soil structure, a low pH and no organic matter (Crowder et al. 1982). This is

accentuated by the absence of an organic fiaction within the surface horizon. The sterility

of tailings has to be rectified prior to adequate plant growth.

Agricuhural limestone is added to raise the pH of the surface to approximately 5.0

as well as large arnounts of fertilizers (550 kgha of 6-24-24 NPK) to increase

rnacronutrient levels (Peters 1995). Since the microclimate of large expanses of tailings

rnay be extreme enough to prevent the establishment of vegetation (Crowder et al. 1 982),

straw is spread to help stabilk the surface, increase cohesion and aggregation, provide

nirface roughness, improve water fihration and retention, and to dampen temperature

extremes. Straw also acts as a seed trap for local indigenou species and, as it

decomposes, it serves as a source of organic matter. Seeding occurs in late nimmer and

early fdl and may be repeated for a couple of yean afterwards. The gras seed mirmire is

composed of Canada bluegrass (Poo pratensis L.), t h t h y (Phleum pratense L.), tall

fescue (Festuca arundinucea Schreb.) aod creeping red fescue (Festuca rubru L.).

Legurnes suc h as birdsfoo t trefoil (Lotus cornicuIahis L.) are seeded the folio wing

season.

The development of tailings ecosystems is slow even with these reclarnation

procedures. Ho wever? natural recover y would be even slo wer because some ear ly pioneer

species, such as lichens and mosses, are highly sensitive to acidity and metal

contamination Even afler a grass cover has been established on tailings, recurring

problems of re-acidification can threaten plant growth (Peters 1984). Like that of other

disturbances, the general effect of depositing tailings in an area is to push back

succession to an early stage.

Alt hou& the pro blems of establishing sustainable vegetation on tailings may

seem next to impossible, in most areas, a mat of vegetation bas developed on the surface.

Approximately 10 years from the firn seeding of grasses, a 2-3 cm organic horizon

begins to f o m Despite problematic drainage, and seepage waters fiom the tailings, it

appears that establishing a cover of vegetation reduces the amount of infiltrathg water by

intercept hg prec ipitat ion, reduc ing evapo transpirat ion and by forming an oxygen-

consuming barrier (Peters 1995). Thus, it seerns that the severity of acid-mine drainage is

diminished.

Little is known about the ecology of MC0 Ltd. tailings ecosystems. Nevertheless,

it is suspected that maturing ecosystems with diverse flora and huna will be the most

sustainable. About 70 species of vo lunteer plants and several species of birds and

mamrnals have become established since the work of Peters; however. few detailed

studies on the invertebrate huna have been undertaken,

III. AN INTRODUCTION TO SPIDER BIOLOGY

The spider fauna in any terrestrial region of the world may be as suitable for the

characterization of biotopes as are vascular plants (Clausen 1986). Furthemore, spiders

are excellent indicaton of atmospheric S 0 2 (Gilbert 1971) and heavy metal burden

(Clausen 1984, Price et al. 1974). Because al1 spiders are predaceous, there is a potential

for biological concentration of toxic matter. "Spiders ( a h ) constitute one of the best

indexes for the investigation of community structure, stratification, and succession"

(Bames 1953). There are three suppoaing reasons for this sweeping generalization. First,

spiders are generally abundam in terrestrial cornrnunities ensuring large enough samples

for numerical analyses. Second, their high degree of adaptive radiation also provides a

variety of forms to fùlfill a variety of ecological niches and third, the order is small

enough so that a working knowledge of the taxonomy is not beyond the capabilities of a

single worker.

Any diversity study, regardless of the number of contnbutors, requires that a

thorough knowledge of the study organism be integrated with meanirements of

abundance, richness, or some combination of the two. This knowledge takes the form of

taxonomie discernent, ecolog ical awareness, and the perception of reg ional and local

distribution patterns. All levels of diversity are nested within these three resources. A

regional faunistic assessrnent of wandering spiders wiil be taken into account in a later

section; this section explores their taxonomy and ecology. Wiîh a weil-rooted

background, the wandering spider guild may then be examined for its appropriateness as

an index of the sustainability of INCO Ltd. reckimed tailings ecosystems.

m. A. The Classification of Spiders

Spiders, ticks, horseshoe-crabs, harvestmen, scorpions, pycnogonids and other

like invertebrates are grouped in the arthropod subphylum Chelicerata The class

Arachnida includes the eleven orders Palpigradi (rnicrowhipscorpions), Araneae

(spiders), Amblyp yg i (tailless whipsco pions), Thel yphonida ( whipscorpions or

vhegaroons), Schizomida (no common name), Ricinulei (no common narne), Acari

(mites and ticks). Opiliones (harvestmen), Scorpiones (scorpions), Pseudosforpiones

( pseudoscorpions) and Soli fbgae (sunspider, windscorpion, or so lpug id). In generai,

members of the class Arachnida are terrestrial arthropods that have the body divided in

two main regions and have a pair of chelicera, which end in pincers or fangs.

The traditional view has placed the order Araneae as sister group of the order

Amblypygi. M e r much cladisitic analysis, spiders are now postulated to be a sister order

to the Pedipalpi ( Amblypyg i, Schizo mida, and Thelyphonida) (Coddington and Levi

1991, Shultz 1990) (Figure 4). Before 1880, spider classification was based on broad

categories of lifest y les result ing in a paraphy let ic arrangement. Today, the

monophylogeny of Araneae is supported by several complex and unique

synapomorphies. The most important of these are abdominal appendages modified as

spinnerets, silk glands and associated spigots, che liceral veno m glands. male pedipalpal

tarsi modified as sperm transfer organs, and loss of abdominal segmentation (Coddington

and Levi 1991).

Within the Araneae, three major groups are generally recognized: Mesothelae,

Mygalomorphae, and Araneomorphae. The suborder Mesothelae contains the single

family Liphistiidae (2 genera, 40 species) limited to China, Japan, Southeast Asia, and

Sumatra (Platnick and Sedgwick 1 984). The idkaorder Mygalomorp hae ( 1 5 families, 260

genera, 2200 species) include the Theraphosidae (baboon spiders or tarantulas),

Ctenizidae, Actinopodidae, and Mig idae (trapdoor spiders), Atypidae (purse- we b

spiders), Hexathelidae (fuiuiel web spiders), and several groups with no common name.

The infraorder Araneomorphae (90 families, 2700 general, 32,000 species), somet imes

referred to as "true" spiders, includes all remaining spider taxa.

Wandering spiders form a guild, ail of whose members are within the infiaorder

Araneomorphae. The guild as first described by Breymeyer (1966) included the families

Clubionidae (sac spiders), Gnaphosidae (ground spiders), Lycosidae (wolf spiders),

Pisauridae (fishing spiders), Thomisidae (crab spiders), and some representatives of the

Agelenidae (grass spiders) and Hahniidae (no commo n name). Salt icidae (jumping

spiders) and Philodrornidae (crab spiders) have since been added to this list (Uetz 1975).

These families are similar in size, general body form. and mode of prey capture. The

Thomisidae is an exception with regard to body foxm (legs 1 and II are laterograde raîher

Figure 4. Cladogram of the arachnid orden (after Shultz 1990).

than prograde) but have been observed to forage in the same manner as other members of

the guild (ninning down or pouncing on prey) (Uetz 1 975). The wandering spider guild

roughly f t s h o the "RTA Clade", or the "retrolateral tibia1 apophysis clade"

(Coddington and Levi 199 1) (Figure 5). The RTA clade lies at the same taxonornic level

as the Orbiculariae. the orb weavers.

The two most diverse families within the wandering spider guild are the

Lycosidae and the Gnaphosidae. Globally, the Lycosidae includes nearly 2000 species.

Of the 300 species in North America, 107 species are found in Canada and Alaska The

Gnaphosidae are not as taxonomically well known; nevertheless, they comprise an

estimated 1 500 species. Of the 330 species of Gnaphosidae in North America, 100

species are represented in Canada and Alaska.

In. B. Spider Anatomy

The spider body consists of two main parts, an anterior portion, the prosoma (or

cephalothorax) and a posterior part, the opisthosoma (or abdomen). A nanow s d k , the

pedicel, connects these parts (Figure 6). With respect to functions. the prosorna serves

mainly for locomotion, for food uptake, and for nervous integration (as the site of the

central nervous system). In contrast, the opisthosoma fulfills tasks associated with

digestion, circulation, respiration, excretion, reproduction, and silk production.

The prosoma is covered by a dorsal and a ventral plate: the carapace and the

sternum, respectively. It serves as the place of attachment for six pairs of appendages:

one pair of bit ing chelicerae and one pair of leg-like pedipalps are situated in fiont of four

pairs of wallcing legs. In mature male spiders, the pedipalps are modified into copulatory

organs.

The "head" part of the prosoma ba r s the eyes and the chelicerae. Most spiders

have eight eyes, which are arranged in specific patterns in the various families. Usually

the eyes lie in two rows, and accordingly they are referred to as anterior lateral eyes

(ALE), anterior median eyes (AME), ponerior lateral eyes (PLE), and posterior median

eyes (PME) (Figure 7).

Figure 5. Cladistic hypothesis for the Infiaorder Araneomorphae (after Coddington and Levi 1991).

ids

Thomisidae Othe? dionychans Amsurobiidae Other amaurobioids Tengetlicke Acanthoctenid ae 20-dae Ctenidae Pisauridae T techaleidae Lyeosidae Psechridae

Araneidae tinyphiidae cyattioiip'dae Synoîaxidae NestMdae TneMiidae

Figure 6. Anatomy of a generalized spider of the Infiaorder Araneo morphae (after

Preston-Mafham 1 99 1 ).

Figure 7. Carapace of a member of the Salticidae showing arrangement of eyes. ale,

anterior lateral eye; orne, anterior median eye; ple, posterior lateral eye; pme, posterior

median eye (after Dondale and Redner 1978).

The chelicerae are the fust appendages of the prosoma. Each chelicera consists of

two parts, a stout basal part and a movable articulated h g . N o d y the fang rests in a

groove of the basal segment like a blade of a pocketknife. When the spider bites, the

fangs move out of their groove and penetrate the prey. At the sarne time poison is

injected through a tiny opening at the tip of the fang. Both sides of the cheliceral groove

are ofien armed with cuticular teeth. These act as buttresses for the movable fmgs and, in

addition, allow the spider to mash a prey item into an unrecognizable mass. Spiders

without such teeth can only suck out theu victirns through small bite holes formed by the

fangs.

The second pair of appendages is the pedipalps. With the exception of an absent

metatarsus, pedipalpal segmentation corresponds to that of the legs. Despite their general

resemblance to legs, the palps are usually not used for locomotion. Instead, they ofien

play a rnanipulative role during prey catching. The most notable modification of the palps

is found in male spiders. Male palps act as copulatory devices by ha sucking up fiesMy

deposited spem on the male's sperm web and then depositing this into the fernale's

copulatory organs.

The mouth opening is bordered lateraiiy by the maxillae, in fiont by the rosmim,

and in the back by the labium. The four mouthparts form the mouth proper, which leads

into a flattened pharynx. The pharynx consias of a movable, hinged fiont (rostnim) and a

back wall (labium) and is lined by cuticular platelets. These contain very fme grooves

covered by srnall teetb which together function as a micro filter. The pharyngeal lumen

c m be widened by the action of several muscle bands. Thus, the pharynx acts as a suction

pump and the spider does not chew its food but instead sucks the contents of its prey

through the holes or macerated sections it makes in the prey's exoskeleton.

Four pairs of legs fan out radially fiom the pliable connection between carapace

and sternum These legs are referred to as legs 1, II, IIi, and IV starting fiorn the anterior

pair. Each ieg has seven segments: a shon coxa, a short trochanter, a long femur, a knee-

like patella, a slender tibia and metatarsus, and nnally a uunis with two or three claws.

The tip of the tarsus bars two bent claws, which are generally senated k e a comb; a

third claw may be present between them. Many hunting spiders possess dense tufts of

hair, the scopulae, directly under the claws. Al1 spiders that have scopulae on their feet

can easily waik on smooth vertical surfaces.

Most spiders bear a sofk, expansible and unsegmented opisthosoma. Only the

Mesothelae, believed to represent an ancient fonn fiom which present-day spiders are

derived, possess a clearly segmented abdomen (Platnick 1 995). The antenor dorsai

surface of the opisthoçorna may possess a darkly coloured, triangular mark that may

stretch to the midway mark toward the spinnerets. This is the heart mark and under it is

found the spider's primitive heart. On the undersurface, again toward the antenor end, is

a pair of book lungs and a single epigastric furrow. Both the male and fernale's

reproductive organs are found beneath this furrow. In fernales however, this h o w is

normally sclerotized fomiing an epigynal plate with a pau of pores, one on either side of

the midline. These openings a10 w the insertion of a male's charged palps and lead

directly to the sac-like spennathecae where semen is stored until oviposition Retrolateral

to the furrow are the book lungs. Primitive spiders have a second pair of book lungs

found toward the posterior end of the abdomen directly in fiont of the spinnerets. A pair

of spiracles and associated tracheae in advanced spiders replaces these posterior book

lungs.

A spider has three pairs of spinnerets on its abdomen, which represent modifîed

appendages. The spinning glands terminate in little spigots on the surfhce of each

spinneret. AU three pairs of spinnerets, anterior, median, and posterior, are extremely

mobile because they are equipped with a well-developed musculature. The antenor

median pair is O ften extremely reduced and many spiders (such as Linyphiidae,

Therîdiidae, and Thomisidae) have only a vestigial bump, which is referred to as the

colulus. In the remaiaing spiders, the colulus is absent altogether. Numerous spiders

possess an additional spinning organ, the cribeilum, a srnall plate located in fiont of the

three pairs of spinnerets. The cnbellar ara is densely covered with many tiny spigots

through which are extrudeci thin silk threads of the "hackle band". These thin silks are

combed out of the cribellu. by rhythmic movements of the calamisaun, a row of comb

shaped hairs situated on the metatarsi of the fourth legs. Al1 wanderiug spiders are

without a cnbellurn and are accordingly terxned ecribellate.

Since the families L ycosidae and Gnaphosidae are the most diverse of the

wandering spiders, a discussion of their anatomy bars greater import. Members of the

L ycosidae are also known as the three-clawed hunters because they possess a third or

median claw at the tip of each leg tarsus. Individuals are elongated and eaher flattened or

somewhat cylindrical. Theu eight eyes are arranged in either two or three transverse

rows, which may be straight, procurved, or recurved according to the genus or species.

These eyes are also grouped in four pairs: the anterior median, anterior lateral posterior

median, and posterior lateral. The lateral eyes possess a peculiarly shaped tapetum, a

layer of light-reflecting cells. In most spiders, this tapetum takes the shape of a canoe but

in the Lycosidae, it has the shape of a grate, or a lattice.

Memben of the Gnaphosidae share many characten with the Clubionidae and

related families of hunters but are distinguished fiom al1 of them by the presence of an

oblique depression on the ventral suface of each palpcoxal lobe. In addition, the

spinnerets are unisegmented and the anterior spinnerets are elongate, well sclerotized,

cylindrica~ and well separated at their base. Many representatives also possess irregularly

shaped secondary eyes and conspicuous cuticular epigynal marg ins.

III. C. Anatomy as it Relates to Classification

The classification of spider fiunilies relies on the structure of the spinnerets,

chelicerae. tanal claws, and the labium. Genital structures however, are used rnainly for

the separation of species and are the only features that aEord any reliable identification.

Consequently, only aduh specirnens may be accurately identified to species. Dondale and

Redner (1990, 1982, 1978) and Pktnick and Dondale (1992) give excellent accounts of

sexual organ anatomy.

The tarnis, pretamis, and the tibia of the d e ' s palpus are modified to form a

copulatory organ cailed the pedipalp. The pedipalp consists of a dorsal shield-like

cymbium and a rounded genital bulb. The pedipalps of male spiders Vary greatly in form

and complexity. In their simplest form, each pedipalp bears on its cymbium a teardrop

shaped genital bulb (Figure 8). The more complex palpal organs are formed of hard parts

and w> ft parts called sclerites and hematodochae. respective1 y; the sclerïtes bear processes

called apophyses (Figure 9). The genital bulb in these spiders consists of a weil-

sclerotized tegulum, within which are found an haorninent organ called the embolus. the

seminal duct and the seminal reservoir. A terminal apophysis is associated with the

embolus and a median apophysis is associated with the tegulum The palpa1 tibia in many

wandering spiders supports a weil-scierotized retrolateral apophysis. However, this

apophysis is lacking in the L ycosidae. Al1 of the variously shaped apophyses are heavily

used to ciassa adult males to species.

Female spiders possess a pair of ovaries in the opisthosoma. n i e lumen of eac h

ovary leads into an oviduct, and the two oviducts unite to form a uterus ( a h called the

vagina). The uterus opens to the outside in the epigastric b o w . Maoy spiders possess a

complexly smictured sclerotized plare just in fiont of the epigashic furrow. This plate,

called the epigynum, extends over the genitai pore and bears the copulatory openings

(Figure 10). AU female members of the wandering spider guild possess an epigynal plate.

This epigynum is heavily used to classify adult fernales to species.

Figure 8. A simple male pedipalpal copulatory structure (after Bnixa and Bnisca 1990).

Figure 9. A complex male pedipalpal copulatory stmcture (&er B w a and Bnisca

1 990).

Figure 10. Cut-away, dorsal view of a fernale spider reproductive system ( d e r B m a

and Bnisca 1 990).

0 / X...'

Cymbium Zbia (= I~RUS)

ktdlbrriai duct

8

III. D. Wandering Spider Ecology

Understanding the role a study organism plays in an ecological context is crucial

to any diversity assessment. nie ecology of wandering spiders should be reviewed before

any attempt is made to examine their diversity on tailings habitats. Topics that need to be

investigated hclude feeding behaviour, reproduction, vagility, habitat preferences, and

survivonhip . Hunting or vagabond spiders employ difTerent strategies fkom web building

spiden. They restrict their use of s ik to the dragline they ofien trail behind them as they

move about, to protect the eggs, or, in some cases, to iine their retreat. Unlike web

builders, they are not restricted to prey that cornes to them but rnay venture forth to

searc h O ut desirable prey species. The y have greater opportunit ies to exercise c ho ice or

preference in respect to prey. Because hunting spiders have no permanent station at

which they rnay be observed, Little work on prey preference has been reponed (Tmbull

1973). They consume and discard their prey at the points of capture, which rnay be

widespread.

A few papes report on the range and movements of hunting spiders (mainly

Lycosidae) (Hailander 1967, McCrone 1965). However, it is difficult to assign motives to

these movements. We may observe a spider's wandering for reasons we cannot assess;

we cm only assume it is searching for prey when we observe it reacting to the presence

of prey.

Regardless of how a wandering spider searc hes, it must somehow approach close

enough to its prey items to seize and to overpower then Ambush is a cornmon strategy.

The genus Misumena Latreille (Thomisidae) sits motionless on flowers and ambushes

visiting insects seeking pollen or nectar. The spider extends its long, forward-directed

raptorial forelegs and awaits the approach of a prey item. Active running is another

common strategy ; the Philodromidae pursue and overtake small prey. The Saiticidae are

the most active of the wandering spiders; they wander over surfaces and foliage searching

with po werfùl eyes. In contras to the web builders, the primary eyes of the Salticidae can

form sharp images and the binocular vision of the anterior median eyes enables the

estimation of distances. The Salt icidae hunt only during &y light and apparent ly are

unable to capture prey in the dark. The Lycosidae and Pisaundae are gewrally rapid

runners, also with good eyesight. They may pursue, pounce, bite, cwh, and digest many

kinds of ground-dwelling invertebrates. Although very active on the soi1 surface and in

the leaf liner of foreas and meadows, they often wait quietly in ambush for their prey.

Their eyes cannot form images that are as clear as those of the Salticidae, but their

eyesight is better developed than that of the web builders. The Gnaphosidae and

Clubionidae are closely related and share similar habits. They are all short-sighted and

hunt mainly at night. Exceptions to the nocturnal activity of most Gnaphosidae are certain

re presentat ives of the genera CaIIiZepis Westring , and Micaria Westring , and po ssibly

others that specialize on ants as prey. Comparable observations on North Amerka

Gnaphosidae are lacking (Tunibull 1973).

One of the c haracterist ic behavioural features of females in the famil y Lycosidae

is the carrying of egg sacs on their spinnerets. The female transports the sac throughout

the incubation period and carefully hunts for it if it becomes dislodged. She also carries

the newly hatched spiderlings in a cluster on her abdomen M e r several days, during

which the young spiders eat nothing, they drop off and take up independent lives.

Juveniles of presumably al1 wandering spiders disperse by a process known as

ballooning. The young spiders climb to the tip of a plant stem or some other convenient

prominence where they stand with abdomen uptihed and eject or comb out strands of

sik, which are cast into air currents. When enough sik is airbome to provide a suscient

l a . the spider releases its hold on the substratum. Glick (1939) recorded airborne spiders

in traps on airplane wings at an altitude of 4,500 m. Schmoller (l97la, b) suggests that

many of the spiders found in the alpine and aeolian zones of high mountains may get

there by drift ing on air currents.

The reasons why juvenile spiders balloon seem clear enough: to break up the

family group, thus avoiduig overcrowding and cannibalism. Why older juveniles and

some srnall-bodied adults balloon is far fiom clear. Although adult wandering spiders

have never been observed to balloon (and are likely quite incapable of doing so because

of their size), a few nonmembers of the guild such as the Linyphiidae and Engonidae are

capable of ballooning at ail life stages. Many of the aeronauts must corne to earth in

inhospitable environments; thus, the risks involved must be high.

Wandering spiders are both good walkets and clhbers, and some cm nui rapidly

for short distances. Their principle method of locomotion is walking and they range over

substantial areas. Hallander ( 1967) fouad that adult males of Pardoscl chekata (Thorell)

(Lycosidae) covered straight-line distances of up to 100 m. Dondale et al. (1970) indicate

that the normal home range for many wandering spider species is considerably smaller

than suggested by the maximal distances of travel figures. They suggest this home range

to be about 200 m2. in both of these papers, females moved much shorter distances than

males and had much smaller ranges. Though a specific home range is suggested by these

data no one has yet filly demonstraîed the existence of a definitive home range for these

animals.

Some work has also been done on the habitat preferences of the wanderinkspider

guild. Since this is a rather large guild, it would be best to examine the two moa diverse

families, the Lycosidae and the Gnaphosidae. Some genera of the Lycosidae occupy

mainly wet habitats such as bogs and swamps (Pirata Sundevail). Othen are

characteristic of grassy meadows or deciduous forests (Schizocosa Chamberlin, and

Trochosa C.L. Koch), and ni11 others are partly (Hogna Simon) or completely

mbtemean (Geolycosu Montgomery). Members of a few genera occupy a diversity of

habitats fiom arctic or alpine tundra, or both, to praùies, salt marshes, sandy kaches, and

dense forests (Pardosa C.L. Koch, Arcrosa C.L. Koch). Activity may continue in winter

under snow (Aitchison l984a b, 1978). According to Grimm (1 985), (as noted by

Platnick and Dondale 1992) moa Gnaphosidae are found in bright dry habitats such as

stony hillsides, grasslands, vineyards, and crevkes in tree trunks. Only a few inhabit wet

fields, meadows, or bogs, and almo st none live in dense shady forests. They Live in plant

liner, in crevices on tree trunks, and among stones.

Because female spidm produce an average of well over 100 eggs, most spiders

mut die before maturity. Not much is kwwn about the causes of mortality, or how

mortality may Vary with changes of spider population density. Spiders have been shown

to be rernarkably resistant to starvation (Tumbull1962), hence, pro bably few spiders

starve, except perhaps immediately after spiderihg dispersal fkom the family cluster. A

number of insect parasites attack and kill spiders and several predatory wasps specialize

in spiders, but these account for only a very small proportion of spider mortality. It is

supposed that new ly independent spider ling s are vulnerable to starvation, that many

spiders die during molts, that many are lost through ballooning mishaps, and that birds

and rodents eat many spiden, but none of these possible reasons for mortality have k e n

quantitatively documented.

Adverse weather is O fien cited as the cause of death of terrestrial arthropods

(Uvarov 193 1 ). Cold winters are said to be particularly hard on northem and temperate-

zone spiders. A few species, however, are extremely cold resistant, as presurned fiom

their Arctic distribution (Leech 1966). Most northem wandering spider species

ovenvinter in their penultimate adult instar. Dondale and Legendre (197 1) report that the

spider Pisaura mirabilis (Walkenaer) (Pisauridae) ovenvinters in a state of diapause.

Leech (1966) and Dondale (1961) report that in colder climates some n o m l l y annual

spiders adopt a biennial or even longer period of development.

TV. SITE DESCRIPTIONS

W . A. Study Locale

The audy area was in the region of Sudbury, Ontario (46'2 1' N, 80'59' W, 259 m

above sea level) (Figure 1 1). The data were collected in an area centering on the 38 1-m

ta11 smokestack of WC0 Ltd. in Copper C E , Ontario.

The study area is influenced by a continental climate. Precipitation averages 84

cm annually, with snowfalls representing 26% of this total. The average mean

temperature ranges fiom - 12OC in January to 20°C in July, and the average number of

fiost fiee days per year is 183 (Freedman and Hutchinson 1980).

IV. B. Study Sites

INCO Ltd. tailings are located West of the smeher in the town of Copper Cliff

(Figure 1 1 ). For engineering and management purposes, the tailings area had ken

divided into a numkr of distinct sites. Fresh tailings were without vegetation whereas

vegetation had been established in older sites. Four sites were sampled for wandering

spiders (Figure 13, based on their age and on the diversity of vegetation. Ali tailings

sites are henceforth denoted A, B, C, and D fiom least to greatest successional age;

similarly, 1,2, 3, and 4 denote the control sites fiom least to greatest successional age.

The geographical coordinates for al1 these sites were recorded to the nearest 15-m (Table

1).

The four control sites were located in various areas within the Sudbury region

(Figure 11). These were chosen based on their sirnilarity to the four tailings sles, as will

be described. A control site was not established for the bare tailings site (Site A) because

naturai equivalents are not found within the region. Consequently, control sites 1,2, and

3 were simiiar in physiognomy to tailings sites B, C, and D, respectively. These control

sites were chosen based on similarities in size, physiognomy, and age to the tailings sites.

These variables were determined with the assistance of the Ontario Ministry of Naturai

Resources Forest Fire Management Centre, who sifted through their database of

Figure 1 1. Localities of study sites in the Sudbury region. 1-4, wntrol sites; T, al1 four

tailings sites. See text for their description.

Figure 12. Location of the four tailings sites. The upper left area is cunently receiving

fkesh tailings while the lower right portion has k e n reclaimed to various extents. A-D.

tailings Sites A to D. See text for their description.

FRESH TAILINGS

Table 1 . The latitude and longitude for snidy sites on the INCO Ltd. tailings area and on

control sites. Replicates represent locations of pitfàil trap clusters.

Habitat Site N W le vat ion' (k15m) (*15m) (ml

Tailings A

B

C

D

ControLs 1

7 -

3

42

~ o t e ' : elevation only determineci for m e replicates ~ o t e ? control Site 4 cardinates determineci using a UTM map

historical fies. It was hoped that these control sites were the naturally recovering

equivalents to the tailings reclamation program A fourth control site, 4, was located on

the Laurentian University campus and represented what the tailings sites would likely

become if left unmanaged. Thk site was not disturbed by forest fies but instead,

represented natural recovery in an area affected by the mining process, as described by

Amiro and Courtin (1 98 1). Habitats such as this likely provided and continue to provide a

source of wandering spider species for the tailings habitats.

The vegetation layers in al1 tailings sites were sweyed as part of a metal analysis

study undertaken by Drs. G. Bagatto and J. D. Shorthouse in the Department of Biology,

Laurentian University. Voucher spec imens are stored in the Laurent ian University

Herbarium, the curator of which is Prof. Keith Witerhalder. A list of plant species was

assembled (Appendk A) and their importance values (IV = [% cover + % fkquencyll2)

were calculated (Appendices C and D). Unfomuüitely, similar tables were not available

for the control sites. It is presented in this thesis as a baseline source for future vegetation

studies.

IV. B. i) INCO Ltd. Tailings Sites

Site A (Figure 13)

WC0 Ltd. officials had narned this site 'A'. This bare, oxidized tailings site had

been in existence for roughly 5 years. No attempts were made to vegetate this site and

consequently, it was alrnost completely devoid of vegetation. Existing vegetation

consisted entirely of sparsely distnbuted clumps of Deschampsiu cuespitosa (L .) Beauv.

(Tufted Hairgrass). Bare tailings constituted over 90% importance, rock constituted

approximately 696, and the remainder was clumps of D. caespitosa (Appendix B).

Site B (Figure 14)

INCO Ltd. officials had also narned this site 'PT. This site was typical of vegetated

tailings within two to five years of establishg grasses such as Pou compressa L.

(Canada Bluegrass) and Festuca rubm L. (Red Fescue). Dead gras and the moss, PohZia

nutans (Headw.) Lindb. constituted the greatest importance (each greater than

Figure 13. MC0 Ltd. tailings Site A. This site is almost completely devoid of vegetation.

Figure 14. MC0 Ltd. tailings Site B. This site is typical of grassland tailings sites after

approximately 5 years pst-liming, -fenilking, and wseeding.

approximately 45% importance value) ( Appendix B) . Agrostis gigantea Roth was present

but was of lesser importance throughout the summer (< 5% importance) (Appendix B).

Early attempts were made to plant Pinus banksiana Lamb. (Jack Pine), but these were

largely unsuccessfbL The site remained as gnissland with very thin litter. Strips of

exposed tailings were often seen between patches of grass.

Site C (Figure 15)

MC0 Ltd. officials had also named this site 'M'. This site was typical of

vegetated tailings afker grasses and legumes had established for approximately 15 years.

Several species of conifers were planted and are notably more successful than on Site B

( Appendix A). Robinia pseudoacacia L. ( B lack Locust ) was planted on this site and was

successful (importance value > 75%) (Appendix C). However, Black Locust was

suffering fkom extensive dieback because of the CO Id clirnate. Numerous species of

volunteer herbs and trees such as Populus tremuloides Michx. (Trembling Aspen) and

Salk sp. (Willow), were well established on this site, parts of which were the most

botanically diverse in the tailings area. Consequently, there was tittle to no exposed

tailings due to a thick litter and humus layer (importance value approximately 45%) and a

moss of the genus Polynichum (importance value approximately 45%) (Appendix B).

S ite D (Figure 1 6 )

INCO Ltd. officials had dso named this site 'CD'. This site was the first site to be

reclaimed (Peters 1984). It was established in the early 1960's and fkst planted with

grasses and legumes followed by various conifers ten years later. Pinus banksiana has

grown most successfully; some stands are over 5 m in height. Conifers dominated parts

of Site D and there was a thick layer of slowly decomposing needle litter under most mes

(importance value approximately 45%) (Appendix B). However, between tree stands,

only a thin to often absent litter layer was present, exposing bare tailtigs to the surface

(importance value approximately 50 %) (Appendix B). Betula papyrifera, Populw

tremuloides and Salix sp. colonized remaining parts of this site. A cornplete list of planted

and volunteer species found in various parts of this site is given in Peters (1984).

Sampling was restricted to the coniferous section within this site.

Figure 15. INCO Ltd. tailings Site C. This site is the most botanically diverse of al1

tailings sites and is approximately 15 years old pst-liming, -fertiliwig, and -seeding.

Figure 16. INCO Ltd. tailings Site D. This site is approximately 30 years old and consists

largely of monocultural stands of Pinus banksiana Lamb. (Jack Pine).

Because of its proximity to areas of fiesh taihgs deposition (Figure 12) fkom

which wind m y transport dried and oxidized tailings dust and because it has suffered

fiom the rupturing of tailings pipes, the reclamation success of this site has been

periodically hindered.

IV. B. ii) Control Sites

Site 1 (Figure 17)

This site was located in the town of Val Caron, Ontario about 15 km north of

Sudbury. This site suffered a 15 ha fire in 1992 making it 4 years old at the time of

sampling. The collection area was shifted approximately 500 m north on May 30, 1996

because of vandalism. This second site, by the bea approximation, was once a potato

field but had gone fallow for upwards of 5 years. The dense ground vegetation within the

field consisted of grasses, sedges, and many European plants.

Site 2 (Figure 18)

This site was located about 12 km northwest of the TNCO Ltd. smelter in the town

of Azilda, Ontario. This site suffered a 10 ha fire in 1988 making it approximately 8 years

oM at the tirne of sampling. This site consisted of a dense overstory of Betula papyri$iera

and Ainus sp. (Alder) with very little understory

Site 3 (Figure 19)

This site was located approximately 20 km north of Sudbury, in the town of

Hanmer, Ontario. It suffered a 60.7 ha fire in 1977, making it 1 9 years old post-fire at the

time of collection, and was quite similar in appearance to the vegetated tailings Site D.

The dominant vegetation consisted of Pinus banhiana with very little understory and

large tracts of exposed soil.

Site 4 (Figure 20)

This site was located on the east side of the Laurentian University campus within

the Birch Transition Zone as describeci by Amiro and Courtin (1981). The dominant

overstory veg eîaîion was Bet ula papyriferu and Populus tremuloides. The dominant

understory vegetation was Vaccinium angustfolium Ait. (Lo wbus h Blueberry). Litter

could be found in depressions and there was a large arnount of exposed, hilly rock.

Figure 1 7. Control Site 1 . This site is an approximately 5 year-old fallow potato field

with rnany herbs and grasses.

Figure 18. Control Site 2. This site is approximately 8 years old pst-fire.

Figure 19. Control Site 3. This site is approxirnately 19 yean old post-fie and is

rernarkably similar to tailings Site D.

Figure 20. Control Site 4. This site is siîuated within the Birch Transition Zone as

described by Arniro and Courtin ( 1 98 1) and is located on the Laurentian University

campus.

V. GENERAL MATERIALS AND METHODS

Pitfàil traps are the standard means of sampling Uwcts and spiders that run about

the surface of the ground (Southwood 1978). Despite conflicting discussion about the

interpretation of ecological information obtained by pitfall trapping, such traps remain the

best available means of sampling wandering spiders (Koponen 1 994, Uetz and Unzicker

1976, Uetz 1975). Trapping over extended periods also serves to equalize the effects of

weather.

The behaviour of spiders rnay affect their propensity to falling and remaining in a

pitfall trap. As demonstrated by Topping (1993), three linyphiid species Vary

dramatically with respect to pitfall trap encounter rates, percent entry, time within a trap,

and overail swival tirne. It is obvious that the pitfall trap is not an efficient design for

the capture of this and other like families. However, the wandering spider guild is

generally more active and does not build capture webs as do the hyphiids. A second

technique w d by arachnologists for population estimates is quadrat sampling. This

technique could have been used in this nudy. However, a continuous, unbiased sarnpling

method is desirable when investigating the population of cursorial spiders. Population

e s t h t e s based on quadrat sampling are heavily influenced by the presence of the

investigator since many spiders escape capture by running away as he/she approaches.

Uetz and Unzicker (1976) have shown that pitfall trapping gives a closer estimate of the

total number of species in a community, and is especially useful in studies of cursorial

species diversity.

The traps in this study consisted of cylindrical, white, plastic containers about 1 L

in volume that were sunk into the ground with the open end flush with the soi1 surfiice. A

small container at the bottom held the liquid into which surface dwellers fell and

drowned. The fiuid preservative consisted of soapy, salty water to prevent both

invertebrate attraction and repulsion and al10 wed for unbiased collection. The soap served

to break the surface tension and the sah served as preservative. Preservatives, which have

been used in other studies, include ethylene glycol and ethawl but these liquids influence

specimen collection and rnay attract vertektes. A funne1 above the bottom container

prevented active insects and spiders fiom crawling out. Plastic lids were suspendeci above

each trap, which served as rain and Litter guards. Traps that were occluded with debris or

were empty of fluid at the tirne of collection were excluded fkom the data set. This was a

notable problem for traps in Site A due to wind-blown taiiings and to exceedingly dry

condit ions.

Prior to installing the traps, three 20-m X 20-m plots within each tailings and

control site were visually established. Four pitfall traps were randomly installed in each

plot based on 1 -m X 1 -m grids using a random numbtrs table. In stmmmy, eac h tailing s

and control site contained 12 pitfall traps equally apportioned in 3 plots. Al1 traps were

installed on May 13, 1996. The traps were emptied and the liquid recharged every

Monday and 'Ihursday fiom May 16 to August 29, 1996. The total pitfall trapping effort

was 26 16 hours per site. Al1 trap inhabitants were returned to Laurentian University

where the spiders were removed, identified to species where possible, and tdied. Only

adult males and females were included in the analyses because it is currently impossible

to idente an immature specimen to the species level. It is possible to identfi gender in

some cases (Arnaya and Klawinski 1 996) but this does not aid diversity studies. The adult

spiders fiom each pitfall trap per collection date were stored in separate vials containing

70V0 ethanol. Becaw of the large numbers of spiders obtained and the tirne needed to

identm each to species, the working &ta set was limited to four collection dates in each

of May, June, July, and August. One Monday and one Thursday collection were retained

both at the beginning and at the end of each month. The total working pitfall trap hours

was thus reduced to 1344 holns per collection site. None of the tmps fiom this reduced

collection were occluded with debris or were empty of preservative thus, all traps were

included in the data set. The reniainhg portion of the coilect ion continues to be stored as

described above.

VI. FAUNISTIC ASSESSMENT OF WANDERING SPIDERS

ON iNCO LTD. TAILINGS AND CONTROL SITES

VI. A. Introduction

Spiders are one of the most common and ubiquitous groups of animals: they are

found over the entire life-supporthg landmasses of the world. Where any form of

terrestrial life exists? it is safe to assume there will be spiders living close by. Spiders

exist in the most northern islands of the Arctic (Leech 1 966), the hottest and most arid of

deserts (Cloudsley-Thomson 1 962), at the highest altitudes of any living organism

(Schrnoller 1970, 1971a,b), and the wettest of flood plains (Sudd 1972). In d l terrestrial

environrnents spiders occupy Wtually every conceivable habitat.

In Canada and Alaska, the most diverse f d l i e s wit hin the wandering spider

guild are the Lycosidae (1 4 genera, 1 07 species), the Gnaphosidae (16 generq 1 O0

species), the Clubionidae (8 genera, 66 species), the Thomisidae (7 genera, 63 species),

and the Philodromidae (5 genera, 47 species). The grand total of members within these

families in Canada and Alaska is 50 genera and 383 species (Platnick and Dondale 1992;

Dondale and Redner 1990, 1982, 1978). Although spiders are very comrnon in al1 parts of

Canada, it is surprishg that little about their distribution or habitat preferences have been

recordeci. Exception are The Insects and Arachnids of Canada CO-authored by Platnick

and Dondale (1992) and Dondale and Redner (1990, 1982, 1978) and Pirata published by

Bélanger and Hutchinson (1 992). However, these publications contain only brief

descriptions of habitat occurrences in conjunction with maps pinpointhg scattered

sightings. Further insight into Sudbury's potential wandering spider fauna cm be deduced

fiorn that of three collections made within 450 km of the city limits. Kmta (1943)

assembled a list of spider species in the Lake Nipissing and Lake Temagami regions,

Martin (1965) collected spiders in Sault Ste. Marie, Ontario, and Freitag et al. (1982)

made collections in Wawa, Ontario.

In this section, a species list has been cornpiled and an atternpt will be made to

determine, in very g e n d terms, the habitat preferences of some of the most abundant

species found on INCO Ltd. taillligs and on their respective control sites. It is hoped that

this list will alert others of wandering spider range extensions, will elaborate on our

inadequate knowledge of their prefened habitats, and may reveal important differences

and similarities between tailings ecosysterns and their surroundhg ecosystems.

W. B. Methods

The broad geographical ranges of the wandering spiders collected on MC0 Ltd.

tailings and control sites was examined using lists and maps compüed by Platnick and

Dondale (1 992) and Dondale and Redner (1 990, 1982, 1978). Where possible, faunal lias

assembled by Kurata (1 943), Martin (1 965), and Freitag et al. (1982) were also used.

It would be a daunting task to determine the habitat preferences of al1 species in

any collection. In very broad terms, the habitat preferences of 10 species were determined

based on their abundance and fiequenc y in this study. The selection criteria were at least

a 2.5% relative abundance (approximateiy 150 specimens) and at least a 75% fiequency

(Le. collected in 6 of the 8 study sites).

Dr. Robin Leech of the Northem Alberta Institute of Technology in Edmonton,

Alberta verified a reference collection consisting of both adult males and females of each

species (where available). The collection is stored in the Insect Museum at Lawntian

University in Sudbury, Ontario, tbe curator of which is B. Joseph D. Shorthouse.

VI. C. Results and Discussion

A total of 5075 specirnens, 7 families, 29 genera, and 74 species (Table 2) were

coilected on the tailings and control sites. The species fiequency within each of the

wandering spider families were as fo llows: L ycosidae, 22 species; Gnaphosidae, 2 1

species; Clubionidae, 1 4 species; Thomisidae, 10 species; Philodrornidae, 4 species;

Hahniidae, 2 species; and Pisauridae, 1 species.

Pardosa C.L. Koch (Lycosidae) predominated in abundance (2616 specimens), in

species richness (8 species), and in site fiequency (collectively in 100% of sites) (Table

2). Four Pardosa species attained a relative site fkquency of 50% or greater. P h s a is

one of the largest of spider genera, cornprishg approximately 1 00 species in North

America, of whic h 46 are represented in Canada and Alaska (Dondale and Redner 1990).

VI Table 2. Totals and frequencies for each of the 74 species of wandering spiders on INCO Ltd. tailings sites and control sites. A blank

ceil indicates a specimen was not collected. Species in bold type indicate selection for discussion.

Taxon

- - - - - -- - -. - - - - - --

'Tailings Sites Control Sites

A B C D 1 2 3 4 Y0 Tot al Oh of Tot al Frequency Frequency

Clubion idae Agroeca prateruis Emetton Casriune ira cirtgirlatu (C. 1,. Koch) Castiarteira descripra (Hen tz) Casfiarieira gertsclti Kaston Custiarieir~ longipulpu ( Hen tz) Clirbiona bryaritae Gertsch Clirbiona chippewa Ger tsch Uirbiona johnsoni Gert sch Cltrbiotta kastoni Gertsch Cltrbiona opeongo Edwards C'lirbiona pikei Gertsch Clirbiorta trivialis C. L. Koch Phr~~rotinprs bnrealis (Em ert on) Scohella prrgnatct (Emert on)

Gnaphosidae CalIiIepis plrrto Banks Drnssodes neglechrs ( K e y ser l ing) Drassyllics depressirs (Emerton) Drussyllics rliger (Banks) Drassyllirs socitrs C harn ber1 in Gltaphosa nttrscorirnr ( L . Koch) Gnapltosa parvula Banks Hupludrcrssirs biconttcs (Erncrton) Hqdodrnssirs himialis (Emerton) Huplodrassrrs sig~lfer (C. L. Koch) Micaria gerrsclti Barrows & lvie

Taxon

'Tailings Sites Control Sites

A B C D 1 2 3 4 % Total % of Total Frequency Frequency

Mictrrici lurtgipc~s Em er t on Micaria lortgispim Emert on Micaria pldicaria (Sundevall) Micariu riggsi Gertsch Micaria rossictr Th or el l Sergiolirs decoratirs Kaston Zelotes exigtcoides Platn ick & Shada b Zclofes fratris Chamberlin Zelotes hentzi Barrows Zelotes pirritams Cham ber lin

Hahn iidae I-lahriiu cinera Emerton Neoanfisteu magna (Keyserling)

Lycosidae Alopecosrr actileata (C lerck) R lopecosa kochii (Keyserl ing) Arctosa wrbicrrndo (Keyserling) Hogna frondicola (Emerton) Pardosa distincfa (Blackwall) Pardosa firsctda (Thor el 1) P ~rdusa hyperborea (Ihorel 1) Pardosn lapidicina Emerton Pardm morlica (Blackwall) Pardosa nruesfa Banks Pardosa sarat ilis ( Hen t z) Pardosa xerantpelir~a ( Ke yser 1 ing) Pircr fa canau'errsis Donda le & Redner Pirafu cantralli Wallace & Exline Piruta insirluris Emerton Pirata minufus Emerton Schizocosa a v i h ( Wal kenaer ) Schizmosu coninitrnis (Emerton) Sc hizocosa crassipalputa Roewer

* VI

Taxon

Tailings Sites Control Sites

A B C D 1 2 3 4 Y0 Total % of Total Frequency requaicy

Sc hitacosa r~tccoo ki (Mon tgom cry) Schizocusa saltatrlr (Hen b) Trochosa terricola Tborell

Philodromidae Pliiludrorrtlc~. ylucidics Banks Thart nt ils forni icintcs (Clerck) Thmintirs strintrrs C . L. Koch Tibelltrs rrraritirriirs (Menge)

Pisauridae Dolorrtedes s?riuitrs G ie be l

Thom isidae Ozyptila gertsclri Kuratn Xystictîs attrpi~ffutcrs Tumbull et al. Xysticrrs discursur~s Keyserling Xystictrs elegarts Keyser l ing Xystictrs ellipticrrs 'Tum bu Il et al. Xystictrs enrertorri Keyserling Xysticrirs/éro (Hen tz) Xj~sficrrs lrrctans (C. L. Koch ) Xystici~s rmnrttnrtertsis Key ser l ing Xvsticrrs péllm O. Pickard-Cam bridge

The most commody trapped species were Pardosu rnoestu Banks (36.33%),

Trochosa terricola Thore Il (9.79%), Pardosa distincts (Blackwall) (7.2 1 %), Pardosa

modica (Blackwall) (5.60%), ZeZotes fratris Chamber lin (4.45%), Schizocosa saltatrix

(Hentz) (4.02%), Neoantism magna (Keyserling) (3.82%), Pirata minutus Emerton

(3.29%), Hognu fiondicola (Emerton) (2. MN), and Gnaphosa pantula Banks (2.68%).

Thirty-six of the 74 species recorded were represented by fewer than ten specimens per

species.

On a broad geographical d e . some interesting points were apparent. Five

species were deemed outside their known distribution ranges, 1 7 were Wtely within their

distribution ranges but have not been recorded fiom the Sudbury region, and the

rernaining 52 species were within their known distribution ranges (Platnick and Dondale

1 992; Dondale and Redner 1990, 1 982, 1978) (Table 3). The five species collected

outside their known distribution ranges include 1 ) Custianeira gertschi Kaston (kno wn

only to the New England States, New York, and Southem Ontario), 2) Clubionapikei

Gertsch (known only f?om Florida to southem Ontano and Maine), 3) HapIodmsus

bicornis (Emerton) (distributed in the southem parts of Canadian provinces and in the

northern United States), 4) Micaria rossico Thorell (as fkr east as central Minnesota), and

5) Xysticus montanensis Ke yserling (as fiu east as Saskatchewan). Three of these species

were represented by oniy one individual whereas H bicornis and M. rossica were

represented by 7 and 1 1 individuals, respectively (Table 2). This is an indication that

these two species may be establishing or have established residency in the Sudbury

region and are not simply incidentals. Distribution ranges difkent from published

records may not be very unusual, especially if one considers the larger than twenty-year

gap in collection dates. AU wandering spiders are assumed capable of ballooning in the

immature stage. They readily catch upward air currents and drift to new habitats where

they may settle and prosper. It would seem reasonable that the Canadian spider fauna will

tend to increase its distribution in an easterly direction, following the prevailing winds.

Indeed, approximately half of these five species likely originated fkom Western Canada.

However, one must be cautious with assumptions about origin since locality records are

confounded by coilectors' efforts, or lack thereof. One must dways keep in mind that

populations are not uniformly distributed over a region.

Table 3. Wandering spider species found on INCO Ltd. tailings and control sites in

conjunction with their known ranges (Platnick and Dondale 1992; Dondale and Redner

1990, 1982, 1978). '+' indicates within geographical range, O-' indicates outside

geographical range and 'likely' indicates likely within geographical range.

Taxon Range

C lu bio nidae Agroeca pratensis E merton Cosiianeira cingulafa (C. L. Koch) Cas~iuneira deseripa (Hentz) Castianeira gertschi Kaston Castianeira ZongipaZpu (Hentz) Clubiona bryantae Gertsch Clubiona chippewa Gertsch Chbionajohnsoni Gertsch Clubiona kastoni Gertsch Clubiona opeongo Edwards Clubiona pikei Gertsch Clubiona trivialis C. L. Koch Phnrrotimpus borealis (Emerton) Scotinella pugnaîa (Emerton)

Gnaphosidae Caflilepis pluto Banks h s o d e s neglectus (Keyserling) Drassyllus depressus (Emerton) Drassyllus niger (Banks) Drassyllus socius Chamberlin Gnaphosa muscorn (L. Koch) Gnaphosa parvula Banks Haplodrassus bicumur (Emerton) Haplodrassus hiemulis (Emerton) H~plodrassus signifr (C. L. Koch) Miconci gertschi Barrows & Ivie Micaria longipes Emerîon Micaria longispina Emerton Micaria pulicaria (S undevall) Micaria riggsi Gertsch Micaria rossica Thorell Sergiolus decoratus Kaston Zelotes exiguoides P latnic k & S hada b Zelotes fiatris Chamber lin Zelotes hentzi Barrows Zelotes puritanus Chamberlin

+ + + - + + + + + + - + + +

+ + t

+ + + + - + +

likely likely likely

+ + -

likeiy + +

likely +

Taxon Range

Hahniidae Hahnia cinera Emerton Neoantistea magna (Keyserling)

Lycosidae A Iopecosa aculeata (C lerck) Alopecosa kochii (Keyserling) Arcfosa nrbicunda (Keyserling) HognafiondicoIu (Emerîon) Pardosa disrincta (Blackwall) Pardosa fmcula (Thoreil) Pardosu hyperborea (Thore 11) Pardosa lapidicina Emerton Purdosa rnodica (Blackwall) P ardosa moesta Banks Pardosa suxatiZis (Hentz) Pardosa xerampelina (Xeyserling) Pima canadensis Dondale & Redner Pirata cantrulli Wallace & Exline Pirata insularis Emerton Pirata minutus Emerton Schimcosa mida (Walkenaer) Schizocosa cornmunis (Emerton) Schizocosa crassipcZpata Roewer Schizocosa mccooki (Montgomery) Schizocosa saltatrix (Hentz) Trochosa terricola Thorell

Philodromidae Philodromus placidus Banks Thanatus fornicinus (Clerck) Thanatus striatus C. L. Koch Tibe1Zu.s muritirnus (Menge)

Pisauriche Dolomedes striatus Giebel

Thomisidae Oryptila gertschi Kurata Xysticus ampullutus Tumbull et al. Xys t im discursans Keyserling Xysticus elegans Keyserling Xysticus e2lipticu.s Turnbull et al. Xysticus emertoni Keyserling Xysticus ferox (Hentz) Xysticus Zuctans (C. L. Koch) Xysticus montanensis Keyser ling Xyst iw pelZax O. Pickard-Cambridge

likely likely

+ + + + f

+ + + + +

likely +

likely + + + + +

likely + + +

+ +

likel y likel y

likel y

likel y + + +

likely + +

likel y -

likely

The boudaries of a species's range are not fixed but fluctuating due to

cornpetit ion, predation, climat ic changes, and many O ther factors. These distributions

rnay also be defhed temporally at seasonal, annual, or greater sales. This could perhaps

explain the variation in the published number of species found or assurned preseat within

450 km of the Sudbury region (Table 4) and on a potenth155 additional species to the 74

collected in this study (Table 5).

North American distributions and regional sightings of the ten most abundant and

most fiequent species in this collection were ewnined to determine if their presence in

the Sudbury region is substantiated and to serve as a preambir to inferences about their

habitat preferences. Two members of the Gnaphosidae, one mcmber of the Hahniidae,

and seven members of the Lycosidae were chosen (Table 2). Becaw no two areas are

identical there is a potentially infinite variety of ecological associations. We can see

strong patterns in cornrnunities, which often reflect important diflerences in

environmental conditions, but the apparently straightforward task of classiQing this

variation proves extremely complex. The attempt to classify ten wandering spider species

in this CO Uection by their habitat preferences is also complex. What little informat ion that

is available rnay be enough to substantiate an image of the wandering spider guild on

INCO Ltd. tailings in Copper Cliff and on sites within the region of Sudbury.

Gnaphosri p m Z a (Gnaphosidae) has been collected fiom Alaska to

Newfo undland, south to Colorado and West Virg inia (Platnick and Dondale 1 992)

(Figure 21). At the regional level the closest record for this species was Sault Ste. Marie,

Ontario (Martin 1965). Gnaphosa panda has a widespread distribution with a sighting

in the Sudbury region therefore, its presence in the Sudbury reg ion is substantiated.

Gnaphosa pamZa was most abundant on tailings Site C (59 individuals)

collected throughout summer 1996 (Table 2). It was also abundant in Site B yet was not

collected on Site A. Based on site descriptions (see Section IV), this species seems to

pre fer bo tanicall y diverse habitats (with both deciduous and coniferous trees) with thick

liner layers but may also be fou& occasionally in grassland habitats. Gnaphosa pantula

does not seem to prefer open sand such as kaches and sand pits. Platnick and Dondaie

Table 4. Published numbers of wandering spider species within 450 km of the Sudbury

region either collected or not collected in the present study. The 1st row of authon

compiled Canadian distribution maps fiom which presence/absence in the Sudbury region

was inferred.

Authors

Collected in the Not collected in

Reg ion present study the present study

Freitag et al. ( 1 982) Wawa, ON 9

Martin ( 1 965) Sault Ste. Marie, ON 14

Dondale and Redner

(1 990, 1982,1978) and Various regions

Platnick and Dondale

Table 5. Species of wandering spiders not found in the present study but predicted to

occur based on known geographical ranges (Platnick and Dondale 1992; Dondale and

Redner 1990, 1982,1978).

Taxon Habitat Preference(s)

Clubionidae Agroeca ornata Banks Clubiona aboti L. Koch Clubiona bishopi Edwards Clubiona canadensis Emerton Clubionafircata Emerton Clubiona kulczynskii Lessert Clubiona maritirna L. Koch Clubiona moesta Banks Clubiona mkta Emerton Clubiona norvegico Strand Clubiona obesa Hentz Clabiona riparia L. Koch Clubionoides excepta (L. Koch) Phrurotimpur alurius (Hentz) Phrurorimpus cerfus Gertsch

Gnap ho sidae Drassyllus eremitus Chamberiin Cesiona bilineata (Hentz) Gnaphosa borea Kulczynski Gnaphosa brumulis Thoreli Gnaphosa microps Holm HepyZlus ecclesiasticus Hentz Micaria aenea ThoreU ûrodrasstls canadensis Platnic k & Shada b Sergio lus ocellarus ( Walkenaer) Sergiolus montunus (Emerton)

Lycosidae Arctosa emertoni Gertsch Arcrosa raptor (Kulcynski) Geolycosa domfiex (Hancock) Gladicosa gulosa (Walkenaer) Geolycosa wrightii (Emerton) Hogm helluo (Walkenaer) Pardosa mackenziana (Keyserling) Pardosa milvina (Hentz) Pirata montanus Emerton Pirata piraticus (Clerck)

W idespread Widespread Deciduous forests. bogs Widespread Bogs, meadows Widespread Widespread Coniferous forests Deciduous forests Bogs. rnarshes Deciduous forests Marshes, lakeshores Deciduous fo rests Deciduous forests Deciduous forests

W idespread Widespread Marshes Coderous forests Meadows Widespread Coniferous forests Coniferous forests Widespread Widespread

Widespread Bogs, meadows Sandy areas Deciduous forests Sandy areas Marshes, bogs Coniferous forests Widespread Widespread Marshes

Taxon Habitat Preference(s)

Philodro rnidae Philodromus cespitum (Walkenaer) Philodromus exilis Banks Philodromus irnbecillus Keyserling Philudromus oneida Levi Philodromus pernix Blackwall Philodrumus p a e i s Keyserling Philodromllr rufus quartus Dondale & Redner Philodromus Mus vibrans Dondale TibelIus oblongus (Walkenaer)

Pisauridae Dolomedes tenebrosus Hentz Pisaurina mira ( Walkenaer)

Thomisidae Coriarachne utahensis (Gertsc h) Misumena vatia (Clerck) Ozyptila distans Dondaie & Redner Oslplila sincera canadensis Dondale & Redner Xysticus britcheri Gertsch Xysticus canadensis Gertsch Xysticus luctuosus (Blackwall) Xysticus punctatus Key ser ling Xysticus trigunatus Keyserling

Widespread Widespread Widespread Coniferous forests CoDiferous forests Forests Coniferous forests W idespread Gr asslands

Widespread, aquatic Wide spread

Widespread Widespread Widespread W idespread Forests Unknown Forests Coniferous forests Grasslands

(1992) suggest that G. purvula rnay be found under Stones, boards, and beach debris, and

in meadows and bogs. This largely implies that it has urban habits and may be found in

sandy areas. Unless there were peculiarities about Site A, impeding the establishment of

this species, Platnick and Dondale's (1992) description contradicts the present analysis.

The fact that G. parvula was found on Site B is somewhat corroborated by their report.

Zelores fratris (Gnaphosidae) has been found fiom Alaska to Newfoundland,

south to California, Arizona, New Mexico, and North Carolina (Platnick and Dondale

1992) (Figure 22). A sighting of Z. fiatris ffom the Sudbury region does not exist in the

literature but its broad distribution justifies its presence in the Sudbury region.

Zelotesfratris was most abundant on tailings Site C (6 1 individuals) (Table 2). It

was relatively abundant on Site D (43 individuals) and control Site 4 (45 individuals), but

only one individual was found on Site A. This suggests that 2. fiutris prefers botanically

diverse habitats with a thick litter layer but may also be found in treed habitats (both

deciduous and coniferous) with open tracts of land or rock. However, Z fratris does not

seem to inhabit sandy areas. Platnick and Dondale ( 1 992) indicate that it may be found in

many habitat types including deciduous and coniferous tree-stands, orchards, meadows,

sagebrush, marshes (salt and fkesh), and sand dunes. The majority of these could be

considered similar to Sites C and D. Absence of Z fiatris on Site A implies peculiarities

about this site that ioipeded its establishment.

Neoantistea magna (Hahniidae) has k e n collected fkom al1 Canada to Alaska It

has also been found in the New England States south to Florida and West through the

northem States to northern California (Kaston 1978). Neither a d s b u t i o n map nor a

sighting in the Sudbury reg ion exists. Consequent ly , the presence of N magna in the

Sudbury reg ion is not well su bstant iated. However, the marked abundance of this species

in the present collection (1 94 specirnens from 6 of 8 study sites), illustrates that

populations are well established.

Neoantistea magna was abundant on ali control sites, mon notably Site 2 (79

individuals), but was not as abundant on the taüings sites (Table 2). In fact, N. rnagnci

was not collected on either Sites A or C. This species seems to have a broad habitat

preference 60 m abandoned agriculNal fields to closed deciduous and coniferous foreas

with thick litter to treed habitats with large tracts of exposed soil. To the author's

knowledge, the habitat preferences of N. magna have never been recorded. It is

interesthg however that it was abundant on control sites yet not on tailings sites of

similar age and physiognomy. This suggests that there were elements to the tailings sites

that hindered it s establishment.

Hognafiondicolu (Lycosidae) has k e n collected fiom the Yukon Territory to

Newfoundland, south to California and Alabama (Dondale and Redner 1 990) (Figure 23).

At a regional scale, H fiondicola hâr been collected in both Wawa, Ontario (Freitag et

al. 1982) and in the Ternagami, Ontario and Lake Nipissing regions (Kurata 1943). Thus,

the presence of this species in the curent study is justified.

Hognafrondicola was most abundant on control Site 4 (50 individuals) and on

tailings Site C (27 individuals) and Site D (25 individuals). This pattern suggests that H

fiondicola prefers both coniferous and deciduous tree habitats but avoids open, grassy

and sandy habitats. Dondale and Redner (1990) simply state that this species is usually

found running on leaf litter in forests or meadows.

Pardosa distincta (Lycosidae) occun ~ o m British Columbia to Nova Scotia,

south to Arizona and to Connecticut (Dondale and Redner 1990) (Figure 24). Pardosa

distincta has also been collected in the Temagarni, Ontario and Lake Nipissing regions

(Kurata 1943) thus there is a high likelihood that it occurs in the Sudbury region.

Pardosu distincfa seemed to prefer the tailings habitats as opposed to the control

habitats. SpecScally, the majority of specirnens were collected &om Site C (1 1 1

individuals) and fiom Site B (99 individuals) (Table 2). This @lies that the conditions

in the tailings sites were more favourable to P. distinct0 than those in the control sites.

Pardosa distincra appears to have a broad range of habitat preferences tiom open, grassy

sites to sites with a nch Litter bed and an overstory of both coniferous and deciduous

trees. This species seems to avoid sandy areas because only one individual was CO llected

fkom S t e A. Dondale and Redner (1 990) also state that this species is fomd less

fiequently on sand dunes, kaches, and quarries. However, they ais0 state that P.

dimtincta occurs less fiequently in deciduous and coniferous woods than in fields and

pastues.

Figure 2 1 . Collection localit ies of Gnaphosa p a d a Banks assembled by Platnic k and

Dondale ( 1 992).

Figure 22. Collection locdities of ZeZotesfiahis Chamberlin assembled by Platnick and

Dondale ( 1 992).

Figure 23. Collection localit ies of Hogna fiondicola (Emerton) assembled by Dondale

and Redner ( 1 990).

Pardosu modica (Lycosidae) has been collected fiom the southem regions of the

Yukon Temtory to Nova Scotia and south to Connecticut (Dondale and Redner 1990)

(Figure 25). Pardosa modica has also been found in the Lake Nipissing, Ontario regions

(Kurata 1943) suppoaing its presence in the Sudbury region.

Pardosa modico was moa abundant on tailhgs Sites B (1 29 individuals) and C

(124 individuals) yet was innequently found on the control sites (Table 2). No specimens

were coilected bom Site 4. These hdings imply that P. modica hdiscriminately selects

both open grassy habitats and bushy habitats. Dondale and Redner ( 1 990) suggest P.

modico prefers swamps, salt rnarshes, and meado ws. Bushy conifwous and deciduous

stands should be added to theu list.

Pardosa moesta (Lycosidae) has k e n found fiorn Alaska to Newfoundland, south

to Utah, Colorado. and Tennessee (Dondale and Redner 1990) (Figure 26). Pardosa

moesta has k e n coilected in the Lake Temagami (Kurata 1943), and Wawa regions

(Freitag et al. 1982). Its presence in the Sudbury region is fitting.

Pardosa moesfa was found in extreme abundance on control Site 1 (1 399

individuals), greater than one third of this study's entke colection. It was also very

abundant in tailings Site C (339 individuals) (Table 2). It seems that P. moesta prefers

habitats exemplified by this abandoned agriculturai field with a diverse assemblage of

grasses and herbs. In addition, it seems to avoid both open, sandy habitats (Site A) and

rocky habitats (Site 4). Dondale and Redner's (1990) findings support this extreme

abundance in Site 1 by stating that large catches fan ofien be made in meadows,

hayfields, marshes, bogs, and even on urban lawns. They also state that additional

habitats include deciduous and conif'erous forests, which this analysis does w t support.

The fact that P. noesta was not very abundant on Site B, the grasslands tailings habitat.

indicates that there were important physical or biological limiting factors.

Pirata mimtus (L ycosidae) has k e n found fiom Saskatchewan to Newfoundland,

south to Utah and North Carolina (Dondale and Redner 1 990) (Figure 27). Closer to

Sudbury, P. minutus has been coilected in the Lake Temagami region (Kurata 1943). Its

presence in the current collection is also fitting.

Figure 24. Co ilection localit ies of Pardosa distincts (B lackwall) assembled by Dondale

and Redner (1 990).

Figure 25. Collection localities of Pardosa modica (Blackwall) assembled by Dondale


Figure 26. Collection localities of Pardosa rnoesta Banks assembled by Dondale and

Redner ( 1 990).

Pirutu ninutus was abundant on control Site 1 (65 individuais) and tailings Site C

(50 individuals) (Table 2). It was not abundant on early successional tailings sites or on

late successional control sites. Consequent ly, a habitat preference is dificult to

determine.

In very general ternis however, it seems that P. minutus prefers open spaces as

opposed to heavily wooded habitats. Dondale and Redner (1990) have determined that

this species prefers meadows, hayfïelds, marshes, swamps, and bogs. Additional

collecting would likely be needed in the present study sites to perceiving a pattern to its

habitat pre ferences.

Schirocoso sultatrix (Lycosidae) occurs in a narrower range, centering on Lake

Huron. However, a scattering of sightings have been made in other parts of Ontario,

Quebec, and Nova Scotia (Figure 28). It has also been found in Colorado, New Mexico,

and northem Mexico to northern Florida (Dondale and Redner 1990). Schizocosa

saltatrhx has been collected in the Lake Nipissing region (Kurata 1943), supporthg its

presence in the Sudbury region

Schizocosa saltatrk was absent fiom Sites D and 3 but was very abundant in Site

4 (132 individuals) (Table 2). It seems that S. saltatrix may prefer mixed forests with

exposed, rocky substrates but avoids sandy or grassy and open habitats. Dondale and

Redner (1990) briefly state that S. saitatrix individuals inhabit deciduous forests. If this

was an acceptable requirement, this species should have been collected fiom Site 3 and

should likely have been more abundant in Site C.

Trochosa terricola (Lycosidae) is widespread from Alaska to Newfo~ndland~

south to northem California, Arizona, and south-central Texas (Dondale and Redner

1990) (Figure 29). Trochosa terricola is quite common in western and central Europe and

likely occurs throughout the temperate and b o r d parts of the Holarctic region (Brady

1 979). Its presence in the Sudbury region is supported by Martin (1 965) in Sault Ste.

Marie and Freitag et al. (1982) in Wawa.

Figure 27. Collection localities of Pirata minurus Emerton assembled by Dondale and

Redner ( 1 990).

Figure 28. Collection local it ies of Schizocosa saltanix (Hentz) assembled by Dondale


Figure 29. Collection localit ies of Trochosa terricola Thorell assembled by Dondale and

Redner (1 990).

Trochosa terricola was collected in aii study sites but was most abundant on

control Site 1 (2 1 7 individuals) and tailings Site C (1 1 1 individuals) (Table 2). Because it

was found at ail sites, this irnplies that T. terricola has a very broad range of habitat

requirements. Dondale and Redner (1 990) &hm that individuals of this species are

generally found under logs or stones in somewhat shady fields and at edges of woods.

Brady (1 979) also states that T. temicola inhabits this type of habitat. They are found

under logs and stones, where they presumably mo lt and constnict egg sacs. Since the

abundance of this species was relatively the same arnong al1 audy sites, linle can be

inferred about the tdings sites in relation to the control sites. Instead, this simply

demonstrates the ubiquity of this species.

Of these 10 species, Zelotes frciris and H o p fiondicola would Likely be good

candidates for indicators of tailings ecosystem health. Their abundance on the tailings

sites roughly rnatched their abundance in respective control sites, assumed to be

approximations of the equilibrium state. However, in order to appreciate hlly these

species as indicators, more data, especially microrneterologicai data, are needed.

This chapter has revealed that only s c a t knowledge of wandering spider habitat

preferences exists and that it is possible to detennine generalized preferences with a

simple table of relative species abundance and fiequency. One of the most abundant

species in this study, Neoantisrea magna, which is also widely distributed throughout

Canada, has never been described in terms of its preferred habitat(s). Knowledge of other

species's preferences is also insignificant. Many of the habitat preferences of the ten most

abundant species were unexpec tedly found to contradict previously published

descriptions. Coasequently , these reports should be re-examined. This section has also

demonstrated that a species list may facilitate the estimation of ecosystem heahh

provided many sites are included in the analysis. Varying abundances of Gnaphosa

parvula, Zelotesfrans, and Pardosu moestu, between sites revealed that there m u t be

important fùnctional differences between INCO Ltd. reclaimed tailings sites and similar,

yet naturally recovering, outlying sites.

VII. WANDERING SPIDER a AND B DIVERSITY AND

COMMUNITY MODELING ON INCO LTD. TAILiNGS AND

CONTROL SITES

VIL A. Introduction

Two major dificulties in sampling a large group of animals are the uncertainty of

çpecies abundance and species richness. Species abundance is the number of individuals

per species and species richness ( a h called a diversity) measures the nurnber of species

within an are* giving equal weight to each species. When can a collecter be certain that

al1 potential species are represented in a collection fiom a particular habitat or geographic

reg ion without jeopardizing the integrit y of the cornmunit y by over-sampling ? Botanists

have long been faced with the problem of over-sarnpling and have developed th

"collecter's curve" or the "species-area curve" (Oosting 1956) whereby collecting

regimes are deemed suficient if the rate at which new species are found is no more than

5% per 10% increase in sample size. Assuming that the removal rate is far less than the

abundance of individuals (as is the case in mon invertebrate censuses), the opposite

problem, that of species underestirnation, becomes an issue. One can never assume that

all of the species in a collection are the maximum that can be found. Species may be

locally "raref', the trapping technique itselfrnay bias their trapping frequency, the

trapping season may not coincide wit h their period of activity, and many other variables

contribute to artifcially low species richness. To cope with this problem, there has k e n

the development of a number of extrapolation methods to estimate the 'hue" nwnber of

species (S) in a community. Ail of these estimators are, to varying degrees, based on the

number of singletons (species represented by a single individual in a sarnple) and

doubletons (species represented by two individuals in a sample). Some of these species

nchness estimators are called ACE (Abundance-based C o v e q e Estimator) (Chadzon et

02. in press), ICE (Incidence-based Coverage Estimator) (Chadzon et al. in press), Chao

(Chao 1984), chao2 (Chao 1987), ~ack' and hck2 (First- and Second-order J a c W e

richness estimators) (Smith and van Belle 1984), and MMMean (Michaelis-Menton

richness estirnator) (Raaijrnakers 1987). ACE was employed in this study to more

accurately compare the health of INCO Ltd. tailings sites to that of control sites based

solely on their numben of wandering spider species.

The most widely used measures of diversity are the information theory indices.

These indices are based on the rationale that the diversity, or information, in a natural

system can be measured in a sirnilar way to the information contained in a code or

message. With a satisfactory measure of diversity, it should be possible to compare the

diversity of dEerent areas. However, diversity measures make the assurnption that

cornmunit ies or ecosysterns are no t site-specific (Haw ksworth and Kalin- Arro yo 1 995).

The main application of these indices is to determine the diversity of subject organisms

within entire regions and are therefore rather limited tools at the level of locality. This

may undennine the extent to which divenity measures derived fiom particular sites can

be used as a basis for generalization However, the essence of present-&y biodiversity

research involves the cornparison between different habitats and ecosystems.

Shannon and Wiener independently derived an information index known as the

Shannon, or the Shannon- Wiener index. It is sometimes incorrectly refened to as the

Shannon- Weaver index (Krebs 1985). Although the Shannon- Wiener index has gained

populanty in diversity studies, it remains a troublesome index to interpret. This dficulty

is attributed to its dependence on three factors: N (the total number of individuals), S and

the relative abundance of the S species. This latter dependency contributes to species

evenness (sometimes known as equitabiiity), a measure of how equally abundant are the

individuals among the species. 1 t is assumed, by convention, that the diversity of a

community increases as the abundances of species become evenly distxibuted (Magurran

1988). Because of its evenness component, the Shannon-Wiener index is highly sensitive

to the abundance of the most abundant species. On the other hand, weii-proportioned

abundances may drasticaily increaw a community's diversity meanire regardless of the

number of species. To c h @ ecological interpretation, a separate measure of evenness is

ofien calcdated.

The Shannon-Wiener index assumes that individuals are randomly sampled fiom

an "indefinitely large" population (Pielou 1 975). The index also assumes that ail species

are present in the sample. If a suficiently large number of sarnples are collecte4 Taylor

(1 978) points out that the corresponding Shannon- Wiener indices will be wnnally

distributed. This property makes it possible to use parametric statistics to compare the

diversity of different habitats. However, satisQing the assurnptions of the index can often

be difficult. If the assumptions cannot be adequately satisfied, an alternative jack-knife

procedure both irnproves the measure and provides confidence lirnits. Quenouille (1956)

orig inally proposed the jack- kni fe procedure which was subsequent ly modi fied by Tukey

(1958). Unfortunately, jack-knifmg limits the use of the Shannon-Wiener index as a

measure of a divenity, or the species richness of localities.

The Shannon-Wiener diversity indices and their associated evenness indices were

jack-knifed in this study because CO llect ion dates, with concomitant seasondit y to species

diversity, could not be regarded as Shannon-Wiener samples. An alternative to jack-

knifing the indices, and hence providing an opportunity for a more quantitative analysis,

would have been to pool the collections fiom individual pitfall traps over the course of

the entire collecting season. This would have provided 1 2 samples per site while

sa t i smg one of the Shannon-Wiener assumptions. However, the second assumption

about the comrnunities, that of an infmitely large and homogeneous assortment of

species, would not be adequately satisfied. The pitfidl traps were installed in three groups

of four permanent traps per site (see General Materials and Methods). Lmportant

microclimatic dserences between individual p itfdl trap locations (e. g . shade, mo isture,

and temperature) leading to heterogeneous densities of species would confound

cornmunity cornparisons between sites. For example, Edgar ( 1 97 1 ) found that Pardosa

lugubris ( Wakenaer) fernales are dispro po rtionately more abundant in c learing s than in

shaded areas possibly due to a preference for h.igher temperatures while sunning their egg

sacs. Pooling all the collections of many pitfall traps from within a site necessarily

moderates any microclimatic e ffects on abundance andor species-specific densit ies.

If the diversity of two sites is to be adequately compared, the use of f3 diversity

measures becomes essential. These measures are typically used in terms of t r a c t s or

environmental gradients but there is no reason why they should be limited in such a

rnanner (Magurran 1 988). Standard ecological techniques of ordination and classification

(Pielou 1984) can be w d to investîgate the degree of association or similarity of sites.

However, the easiest way to measure B divmity, or species-turnover, of pairs of sites is

by the use of simiIarity coefficients.

Many invertebrate diversity studies prernatureiy end discussions once species

richness or species similarities have been described. Additional, and far more rneaningful

information, can be obtained by examinhg the underlying species abundance

distribut ions. A spec ies abundance mode 1 characterizes the structure of a community . Any data set containing information on the number of species and on their relative

abundance will reveal that not ail species are equally common. A few species will

invariably be very abundant, some will have medium abundance, whereas most will be

represented by a few individuals. This early observation has led to the development of

species abundance models that are strongly advocated by many workers including May

(1 98 1, 1975) and Southwood (1978) as providing the only sound basis for the

examination of species divenity. Species abundance models use al1 the information

gathered in a cornrnunity and is the most cornpiete mathematical description of the data.

Although one or more families of distributions will fiequent ly describe species

abundance data (Pielou 1979, they are usually examined in relation to four main models.

These are the geometric modei, the log series modei, the lognormal model and the broken

stick model. The common form of the lognormal model has a truncation at one end to

deal with what Pielou (1975) demibes as the ' ~ e i l line"; the portion of the curve

represent ing the rare and consequent ly unsarnpled species.

When plotted as a rank/abundance graph (Figure 30) with the ordinate

represent h g the rank of species fiom greatest to least abundance, the four models can be

seen to represent a progression fkom low evenness to high evenness. The geometric

model begins the progression at one end, where a few species are dominant with the

rernainder uncommoa The progression continues through to the log series and lognormal

models. where species of intermediate abundance become more common, and finally

ends in the conditions represented by the bro ken stick model where species are equally

abundant. Al1 of these models are well described by Magurran (1 988).

The amgement of models can also be considered in terms of resource

partitioning where the abundance of a species is in some way equivalent to the portion of

niche space it has appropriated (or occupied). Whittaker (1972, 1970, and 1965) has

Figure 30. Hypothetical rank abundance plots i1lustrati.g the typical shape of the four

main species abundance models: geometric series, log series, lognormal, and bro ken

stick. The abundance of each species is p lotted on a logarithmic scale against the

species's rank, in order fiom the most abundant to the least abundant species (after

Magurran 1988).

demonstrated that the geometric model is found prirnarily in harsh environments or in the

very early stages of succession. The geometric model would be predicted to occur in a

situation in which species arrived at an unsatunited habitat at regular intervals of tirne,

and occupied hctions of rernaining niche space (Magurran 1988). As succession

proceeds, or as conditions ameliorate, species abundance patterns grade into the log series

model. A log series pattern would result if the arriva1 of successive taxa were random

rather than regular (May 1975, Bosweii and Patil 1971). The small number of abundant

species and the large proportion of '?are" species predicted by the log series model imply

that, like the geometric model, it will be most applicable in situations where one or a few

factors domhte the environment of a cornrnunity. The majority of communities studied

by ecologists display a lognonnal model of species abundance (Sugihara 1980). The

lognormal model has ken shown to indicate a large, mature and varied natural

community (May 1 986) but may also arise as a response to the statistical properties of

large numbers. The Central Limit Theorem States that when a large number of factors act

to determine the amount of a variable, random variation in those factors will result in that

variable king normally distributed. Sugihara (1 980) has proposed a biological

explanat ion for a canonical lognormal model of species abundance, which may also be

applied to other forms of the lognonnal model, such as the truncated lognormal (Pielou

1975). One must envisage the communai (multidimensional) niche space of a taxon king

sequentially split by the constituent species. The portion of niche space that each species

occupies is proportional to its reiat ive abundance and the probabilty of any fiagrnent of

niche king subdivided is independent of its size. Many other biological explmations for

the lognormal mode1 have been proposed such as Pielou's (1975) sequential breakage

model. The broken stick model (somet imes called the random niche boundary

hypothesis) was fis. proposed by MacArthur (1957) and marks a biologically redistic

end to the continuum of species abundance models. He envisaged the subdivision of

niche space within a community to a stick broken randomly and simuhaneously into S

pieces. Udike Sugihara's lognormal model the broken stick model is concemed with just

one resource and bespeaks a much more equitable state than that of the geometric mode&

log series model and lognomial model.

The purpose of this chapter is to compare a and P diversities and species

abundance distributions of wandering spiders collected fiom reclaimed tailings sites to

that of their control site counterparts, assumed to be a reflection of the "healthy" or

equilibrium state. The A bundance- based Coverage Estimator and the Shannon- Wiener

diversit y index will be used to examine each site's a diversity, or species richness, and

the Morisita-Hom and Bray-Curtis Similarit y indices will be used to examine P divenity,

or species turn-over. It is hypothesized that the succession through time of wandering

spider richness, diversity and evenness on the tailings sites is similar to that of the control

sites. It is also hypothesized that the communities of wandering spiders on the tailings

sites are similar (as deemed by similarity indices) to commwiities on control sites. The

connol sites, fiom youngest to oldest, are hypothesized to support increasingly varied and

diverse communities of spiders, which is reflected in their species abundance rodels.

The successional States of the tailings sites are believed to foilow a sirnilar pattern The

testing of these hypotheses will reveal if the tailings sites are developing biota in

h o n y with their surroundings.

W. B. Materials and Methods

The richness estimator employed in this study is a modification of the Chao and

Lee estimaton discussed by Colwell and Coddington (1995), called ACE. This is a

parametrie estimation of species richness, or a diversity, as opposed to the Incidence-

bsed Coverage Estirnator (1CE)- a nonpafametric estimator. Chao and Lee (1992)

developed a new class of estimators based on the statistical concept of "sarnple

coverage". These estimators have since been revised to cope with overestimation of

species richness when sample numbers are low. The revised ACE can now handle species

with 10 or fewer individuals per sample. The concept of sarnple coverage c m be thought

of as the sum of the probabilities of encounter for the species observed, taking into

account species present but not observed. Another way to grasp the concept is to imagine

a unit line k i n g broken into S segments, with the length of each segment representing the

tme proportion formed by one of the S species found in the full sampling universe.

Sampling coverage then is the sum of the segments that beloag to all species actually in

the sample (Figure 3 1).

Al1 species

Species

I I obsewed

Coverage

Figure 3 1. For the Abundance-based (ACE) and Incidence-based (ICE) coverage

estimators. coverage is represented here as the sum of the probabilities of encounter for

the species observed, taking into account species present but not observed (after Colwell

1 997).

Each of the tailings and control site wandering spider data sets were bandled

using Estirnates 5.0.1, a statistical estimation of species richness software package

developed by Colwell(1997). This package computes many richness estirnators such as

Chao, ~hao', ACE, ICE, and others with a broad selection of input data formats. The data

were input with samples in rows and species in columns; the samples king pooled

monthly abundances. Only ACE was employed in this study. The accuracy of the ACE is

enhanced since the software computes successively pooled estimates based on

randomizations of sample order. Estimates selects a single sample at random, computes

the richness estimators based on that sample, selects a second sample, re-computes the

estimators using the pooled data fiom both samples, and so on until al1 the samples in the

matrix are included. Samples were randomly selected 4 times, once for each of four

pooled monthiy samples. Colwell and Coddington (1 994) explain the strategy of

randomization and estimator evaluation in more detail. The Estirnates software also

computes standard deviations for many of these estimates fkom which standard estimates

of the mean may be easily calculated. Because the samples chosen were pooled monthly

collections, SE indicates the scale of monthly fluctuations in the proportion of singletons

and doubletons.

The a diversity of wandering spiders for each of the four revegetated INCO Ltd.

tailings sites and their control sites was determined using the Shannon-Wiener diversity

index (H'). It is calculated fiom the equation:

Where p, is the proportion of iialividuals found in the ith species.

In a simple, the true value of p, is estimated as nJN (the maximum likelihood estimator,

Pielou 1 969). A substantial source of error when utilizing this index cornes fkom a failure

to include all species fiom the community in the sample (Peet 1974). This error increases

as the proportion of species represented in the sample declines. For this reason, collection

dates cannot be regarded as samples; as with any invertebrate swey, there is always a

marked seasonalit y to species composition This precludes the use of parametric statist ics

to compare the diversity of wandering spiders between collection sites. However, an

elegant tool rnay at least be adopted to improve the estimate and to attach confidence

limits. The method makes no assumption about the underlying distribution (Magurnui

1 988). The j ack-Wig technique involves recalculating overall diversity while rnissing

out each wnple in turn. This creates a series of jack-knife estirnates, one for each of the

removal procedures, whic h are then CO nverted to pseudovalues, VPi, using the fo llowing

equat io n:

VP; = (nV) - [(n- 1 )(V Ji)]

Where n = the number of samples, VJi = the jack-knife estimate and V =

the overall diversity index (in this case, H').

The mean of the VPis is the bea estimate (VP) of the diversity index and the confidence

limits are calculated in the usual way :

Standard emr of VP = Standard deviation of VP~S/ .J~

The jack-knifmg procedure is rather tedious to apply so pooled rnonthly species

abundances were used as sarnples. In this way, only four sarnples were successively

removed while recalculat ing overall diversit y. Because pit fa1 1 traps were randoml y

installed in each of the collection sites using a random numbers table and a delineated

giid it is also assumed that individuals were randomly sampled.

Because the Shannon-Wiener diversity index can often be difficult to hterpret, its

associated evenness index (E) was calculated for each of the revegetated tailings and

control sites. The maximum diversity (H,) which could possibly occur would be found

in a situation where al1 species were equally abundant, i.e. H' = H, = in S. The ratio of

observed diversity to maximum diversity can therefore be taken as a measure of evenness

(E) using the following equation (Pielou 1 969):

E is constrained between O and 1 .O with I .O representing a situation in which al1

species are equally abundant. Al1 evenness values were also jack-knifed using the same

aforementioned procedure since this index also assumes that ail species in the cornmunity

are accounted for in the sample.

The variance of H' and E indices and a correspondhg degree of fkedom may be

calculated for each site (see Magurran 1988) but this only allows for t-tests to compare

the diversity and evenness of taxa between one pair of collection sites at a tirne. If an

eight-way cornparison rnatrix were to be aîtempted, too many of the site pairs would be

deemed significantly similar due to the increased probability of a Type II error, or the

acceptance error. The only alternative to compare the diversity of sites is to calculate P diversity indices designed expressly for this purpose.

A bewildering nurnber of similarity coefficients exist but one of the moa

cornrno nly applied and sat is factory measures is a modified Morisit a-Hom quantitative

index ( Wolda 1 983). This measure takes into account the number of species and theû

abundances to calculate a percent similanty between a pair of samples. It is not strongly

hfiuenced by sample size or species richness (Magurran 1 988, Wo lda 1 98 1, Huhta

1979). However, it û presently limited to pair-wûe comparisons. This allows only for the

creation of large and dificult to interpret matrices; more sample comparisons translate

into larger, perplexing matrices. Hierarchical clustering of sites with respect to their

Morisita-Hom indices would require extensive computer programming knowiedge as the

calculations would be too lengthy and tedious to perform by hand. However, a "virtual",

internet-based computer program exists which calculates clusters for biological data

ushg the Bray-Curtis Similarity measure. This index is also quantitative because it takes

into account the number of species between pairs of sites but is not as numerically sound

as the Morisita-Hom index (Uagurran 1988, Huhta 1979). The clustering technique

al10 ws for the formation of a dendogram whereby site sirnilarities are quickly perceived.

Typically, the cornmunities of subject organisms fiom two sites are considered similar if

their sirnilarity measure is greater than 80% (Pielou 1984).

The Morisita-Hom index of community sirnilarity is calculated as foilows:

2 (anibni) C ~ H =

(da + db) aN -bN

Where aN = total number of individuals in site A and = number of

individuals in the ith species in A.

The Morisita-Hom indices were calculated for each of the study sites in a pair-

wise fashion using the statistical software package EstimateS 5 .O. 1 developed by Co lwell

(1 997). The data were input with samples in rows and species in columns; the samples

king pooled rnonthly abundances.

The Bray-Curtis quantitative, hierarchical clustering technique, was employed

using unweighted aahmetic average distances between samples and clusters. It is

calculated as fo 110 ws:

Where ni i and nzi are the numben of the ith species in the two sarnples.

The 'vinual" intemet-based software w d was developed by Brzustowski (1997) and a

dendogram was created using ~ r e e ~ i e w ' (Page 1997).

According to Huhta ( 1979), the Bray-Curtis measure should not be attempted

without initial data transformation because it is known to be especially inconsistent. On

the other hand, transformation of data fkom the same comrnunity prior to applying the

Morisita-Hom index results in the loss of it showing unity . As such, the n a t d logarithm

of the wandering spider species abundances, [In(x+l)], were w d for the Bray-Curtis

measure whereas raw abundance values were used for the Morisita-Horn index. Huhta

(1979) deduced that the Morisita-Horn index may fail to reveal successional trends but it

is used here because it is known to be mathematically faultless.

To determine which of the four main abundance models best fit the species

distributions within each of the tailings and control sites, the methods used can be found

in M a g m (1988). Each of these models in each of the tailings and control sites were

tested for their fit using the Chi-square goodnesssf-fit test, making certain to group

theoretical values such that they sum to five or more as required by the Chi-square test.

The cornmon practice is to examine which of the four models has the best fit. The

unfortuate aspect to any goodness-of-fit test is that they are limited to the negative

mode. The primary use of the Chi-square test in this study is to reject theoreticai

distributions as potential models for the given set of data. Acceptance of a mode1 at the

95% significance level means relatively litt le, since mathemat icall y speaking , there is an

infjnite cc llection of functions that will fit the data as well or better than the one king

used. An important credit to the Chi-square test is that it d e s cornparisons possible

between competing theoretical models as to how well they fit field abundance &ta.

VIL C. Results

VU. C. 1. Species Richness

VII. C. la. Control Sites

Wandering spider species richness on control sites is presented in terrns of

singletons, doubletons, and others (Figure 32). This figure represents al1 the species of

wandering spiders collected over the course of sixteen collection dates fiom the

beginning of May 1996 to the end of Augua 1996.

The four-year-old pst-f ie control site in the t o m of Val Caron, Site 1, supported

37 species of spiders of which 6 were doubletons and 9 were singletons.

The eight-year-old pst-fie control site in the town of Azilda, Site 2, supported

the greatest number of wandering spider species compared to al1 other control sites.

Among the 42 species trapped here, 3 were doubletons and 10 were singletons.

The nineteen-year-old ps t - f ie control site in Hanmer, Site 3, supported the

fewest species compared to other control sites. Of the 32 species trapped, 3 were

doubletons and 17 were singletons. The collection fiom this site contained the greatest

number of singletons.

The control site on the Laurentian University campus, within the Birch Transition

ecotone, Site 4, supported 34 species of spiders of which 7 were doubletons and 3 were

singletons. The collection fiom this site contaiwd the fewest singletons.

The overall pattern of species richness with respect to control sites can be seen in

the first four columns of Figure 32. The nchness of species rose to a peak of 42 in Site 2

and fell to 32 and 34 in Sites 3 and 4, respectively.

W. C . 1 b. Tailings Sites

Wandering spider species richness on INCO Ltd. tailings sites is presented in

t e m of singletons, doubletons, and others in Figure 32. Similar to the control sites, this

figure represents ail the species of wandering spiders coilected over the course of sixteen

collection dates fiom the beginning of May 1996 to the end of August 1996.

Figure 32. Control site ( 1 to 4) and tailings site (A to D) species richness expressed as

singletons (spec ies represented by a single individual), dou bletons (species represented

by two individuals), and others (species represented by three or more individuals).

Species Richness

Bare tailings, Site A, supported 16 species of wandering spiders of which 3 were

doubletons and 10 were singletons.

The five- year-old, grassland tailing s, Site B, supported 3 5 species of wandering

spiders of which 8 were doubletons and 5 were singletons. Of al1 the tailings sites, this

site supported the fewest number of singletons.

The fifteen-year-old, rnixed wood tailings site, Site C, supported the greatest

number of spider species compared to al l the other tailings sites. Of the 38 species

trappeci, 2 were doubletons and 14 were singletons. This represents the greatest number

of singletons cornpared to all other tailings sites.

Wandering spider spec ies ric hness of the thrty- year-O Id tailings site, Site D, fell

between that of Sites A and B. Thirty wandering spider species were trapped on this site,

5 of which were doubletons and 9 were singletons.

The overall pattern of species nchness with respect to tailings site reclamation can

be seen in the 1st four colurnns of Figure 32. The richness of species rose to a peak of 38

at fifteen years pst-revegetation and fell to 30 at thirty years post-revegetation.

VII. C. 2. Abundance-based Coverage Estimations (ACE)

W. C. 2a. Control Sites

The ACE of the true species richness for wandering spiders fiom al1 four control

sites was as follows: Site 1,45.7 f 1 . 1 species; Site 2,48.8 * 2.2 species; Site 3,65.1 * 7.3 species; and Site 4,36.0 -t 1.6 species (Figure 33). This represents a dserence

between O bserved and estimated number of species as fo ilows: Site 1, 8.7 f 1.1 additional

species; Site 2, 5.8 f 2.2 additional species; Site 3,33.1 k 7.3 additional species; and Site

4, 1 .O k 1.6 additional species. By far the widest SE for control sites richness estimates, k

7.3 species, was that obtained for Site 3. The remaining SE richwss estimates were

exceptionally m o w .

W. C. 2b. Tailings Sites

The ACE of the true species richness for wandering spiders fiom each of the

tailings sites was as follows: Site A, 25.9 * 3.8 species; Site B, 40.7 3.3 species; Site C.

57.0 * 2.9 species; and Site D, 42.7 I 3.8 species (Figure 33). Site C, which supported the

greatest number of singletons of al1 the tailings sites, resulted in the greatest increase in

ACE. Thex estimates represent a merence between observed and estimated number of

species as follows: Site A, 10.9 + 3.8 additional species; Site B, 3.7 f 3.3 additional

species; Site C, 19.0 t 2.9 additional species; and Site D, 12.7 f 3.8 additional species.

The SE for each of these estimates were consistently narrow.

VII. C. 3. Shannon-Wiener Diversity (H ') and Evenness (E)

W. C. 3a. Control Sites

The jack-knifed Shannon-Wiener diversity indices for the wandering spiders on

the control sites were dissimilar (Figure 34). Site 1 supported the least diverse CO rnrnunit y

of spiders with an index of 1.208 * 0.199 (* SE, n = 4). The remaining three control sites

supported sirnilar diversities of wandering spiders with indices as follows (where t SE, n

= 4): Site 2, 3.148 k 0.069; Site 3, 3.202 f 0.159; and Site 4, 3.016 f 0.226. This last

value was the least precise jack-knifed Shannon- Wiener divenity estimate of aU control

sites.

The jack-knifed evenness indices for the wandering spiders on the control sites

forrned a similar pattern as the diversity indices (Figure 3 5). Site 1 supported the least

evenly distributed assemblage of spiders with an index of 0.348 k 0.01 8 (t SE, n = 4).

The remaining three control sites supported spider assemblages with similar evenness

indices. These were (where t SE, n = 4): Site 2,0.791 f 0.006; Site 3,0.792 k 0.01 5; and

Site 4,0.774 k 0.017. AU SE values were remarkably precise.

VIL C. 3b. Tailings Sites

The jack-knifed Shannon- Wiener diversity indices for the wandering spiders on

the taiiings sites were more uniform than those for the wntrol sites (Figure 34). These

indices were as follows (where + SE, n = 4): Site 4 3.047 + 0.136; Site B, 2.780 f .282;

Site C, 2.500 f 0.105; and Site D, 2.946 I .102. The widest error rnargin was that for Sire

Figure 33. Control site (1 to 4) and tailings site (A to D) Abundance-based Coverage

Estimations (ACE) of species richness. Enor bars indicate standard emrs about the mean

estirnate.

Abundance-based Coverage Estimator (ACE)

lu W P cn 0) 4 00 O O O O O O O O O

B. 0.282 units. The remaining standard errors for the tailings sites had magnitudes less

than 0.136 units.

The jack-knifed evenness indices for the wandering spider communities on the

tailings sites were not as consistent as their associated diversity indices (Figure 35). The

spiders collected kom Site A had a notably greater evenness index, 0.898 t 0.01 7 (k SE,

n = 4). The remaining three evenness indices were as follows (where f SE, n = 4): Site 2,

0.673 2 0.026; Site 3,0.636 t 0.010; and Site 4,0.788 i 0.010. These error matgins were

also remarkably precise.

W. C. 4. Monsita-Hom and Bray-Curtis Community Similarity

None of the site pairs reached the elevated Morisita-Horn similarity of 8O%,

which would deem them similar (Table 6). Although a pattern to these indices was

diffiicult to perceive, the similarity between tailings sites and both their adjacent sites and

control sites rnay at least be mentioned.

Of al1 tailings sites, Site A was most similar to Site D at Cm" = 54%. Of al1

control sites, Site A was most simiiar to Site 2 at Cm* = 51%.

Of al1 tailings sites, Site B was most similar to Site C at CmH = 53%. Of al1 control

sites, Site B was most similar to Site 3 at C m ~ = 36%.

Of al1 tailings sites, Site C was most similar to Site D at CmH = 60%. Of al1

control sites, Site C was moa similar to Site 1 at = 74%, the pair-wise sixnilarity

closest to 80%.

Of dl tailings sites, Site D was most similar to C (already reported). Of all control

sites, Site D was most similar to Site 3 at cmH = 63%.

Afier the natural log transformation had been made of al1 the cursorial spider

species abundances over the entire field season sites were clustered with respect to their

Bray-Curtis similarity. The resuiting dendogram (Figure 36) revealed that the comrnunity

of spiders coilected nom the reclaimed tailings Sites C and D were over 60% similar. The

communities of spiders on control Sites 2 and 4 had comparable similarity. Of al1 the

control sites, only the Site 1 community of spiders approached similarity to any of the

tailings sites. Interestingly, this cornmunity was roughly 50% similar to a cluster formed

Figure 34. Jack-knifed Shannon-Wiener Divenity indices (Hg) for all control sites (1-4)

and INCO Ltd. tailings sites (A-D). Error bars are the SE jack-knifed average estimate

where each estimate is based on pooled monthly samples.

Shannon-Wiener Diversity (Hm)

Figure 35. Jack-knifed Shannon- Wiener Evemess indices (E) for al1 control sites (1 -4)

and INCO Ltd. tailings sites (A-D). Error bars are the SE jack-knifed average estimate

where eac h estimate is based on pooled monthly samples.

Shannon-Wiener Evenness (E)

Table 6 . Morisita-Hom (CmH) percent cornmunity similarity indices for

al1 INCO Ltd. tailings sites (A-D) and control sites (1-4).

Figure 36. Bray-Curtis dendogram illustniting the percent similarity of I W O Ltd.

tailings sites (A-D) to control sites (1-4) and to each other. Pairings were constnicted

based on unweighted arithmet ic average distance between samples and clust ers.

Sites

by Sites C and D. The dendogram also illustrated that the community of spiders on Site A

was only 10% similar to a cluaer formed by al1 other sites.

W. C. 5. Abundance ModeIs

VIX. C. 5a. Control Sites

Each of the control site wandering spider species abundances were ploned in

Figure 37. Ali of these c w e s were very similar in shape to one another, and appeared to

fit the lognormal model. There were however, a few subtle diffierences in the pitch of

their descent. The initial descent of both Site 2 and 4 reached a plateau between the 2"

and 41h most abundant species but a plateau was not present in the very steep c w e for

Site 1,

The model of best fit for Site 1 was the tmcated lognormal with X' = 0.76, df =

5. P = 0.98 (Table 7). None of the remaining 3 models approached a signScant fit.

The truncated lognormal was also the model that best fit the abundance

distribution for Site 2 with x2 = 0.95, df= 4, P = 0.92 (Table 7). The geometric

distribution significantly fit the observed abundance distribution only to a marginal extent

(x' = 44.84, df = 3 1, P = 0.05).

For Site 3, the tmcated lognormal was the model of bea fit with X' = 0.07, df =

2, P = 0.96 (Table 7). The log series also significantly fit the data (X2 = 4.2 1, df = 2, P =

0.12) but was not as close a fit as the tmcated lognormal.

The tnuicated lognormal model best fit the wandering spider species abundance

distribution of Site 4 with x2 = 0.06, df = 5 , P = 0.40 (Table 7). This was the only model

that significantly fit the data for Site 4.

W. C. 5 b. Tailings Sites

Each of the tailings site wandering spider species abundances were plotted in

Figure 3 8. Site A clearly had a very simple abundance distribution, beginning at 3

individuals for the most abundant species and ending at one individual for the 1 5"

species. The rernaining species abundance distributions appeared to fit the lognormal

Figure 37. Control site (1 to 4) species abundance distributions. The abundance of each

species is plotted on a logarithmic scde against the species's in order fiom the mon

abundant to lean abundant species.

Abundance

Table 7. The results of Chi-square goodness-of-fit tests for each of the four main abundance models on the observeci cursorial spider species a bundances for control sites ( 1 to 4). X' Chi-square value; P. P-value: df, degrees of fieedom. Numbers in bold indicate the model of best fit, ** a highly significant fit and * a significant fit.

Control Sites

Geometric 5701.99 ~0.01 29 44.84* 0.05 31 Log Series 16.97 ~0.01 2 17.91 ~0.01 3 Tuncated Log Normal 0.76** 0.98 5 0.95** 0.92 4 Broken Stick 50.75 ~0.01 5 16.25 0.01 5

Geometric 41.96 <0.01 17 93.56 <0.01 26 Log Series 4.21** 0.12 2 19.63 ~0.01 2 Tuncated Log Normal 0.07** 0.96 2 0.06** 0.40 5 Broken Stick 7.22 0.03 2 11.19 0.02 4

abundance model. However, the plateaux between species ranked 2"d to 4' and 5' to 7'

in the cuve for Site C hinted at the broken stick model. Al1 of these curves had

exceptiondy long tail ends.

The model of b a t fit for Site A was the geometric model with X' = 1.67, df = 2, P

= 0.98 (Table 8). The other three models fit the abundance distribution for this site as

follows: the log series model (X2 = 0.74, df = 1, P = 0.39), the tnincated lognomial model

(x' = 1.5 1, df = 1, P = 0.22), and the broken stick mode1 (x' = 3.08, df = 1 , P = 0.08).

The truncated lognormal model best fit the abundance distribution for Site B with

X' = 2.60, df= 4, P = 0.65 (Table 8). One other model with a iess signifiant fit was the

log series model O(' = 2.60, df = 4, P = 0.65).

None of the four main models fit the w a n d e ~ g spider species abundance

distribution for Site C (Table 8). Only the log series model approached signifïcance (x' =

16.15, d f = 2, P < 0.01).

The model of best fit for Site D was the geometnc model with x2 = 21.87. df =

20, P = 0.35 (Table 8). The only other models that significantly fit the abundance

distribution for this site was the tnincated lognomial model (x' = 5.83, df = 3, P = 0.12).

Figure 38. Tailings site (A to D) species abundance distributions. The abundance of each

species is plotted on a logarithmic sa le against the species's rank, in order from the most

abundant to least abundant species.

Abundance

Table 8. The results of Chi-square goodness-of-fit tests for each of the four main abundance models on the observed wandering spider species abundances for INCO Ltd. tailings sites (A to D). X' , Chi-square value; P , P-value; df: degrees of fieedom Numbers in bold indicate the mode1 of best fit, ** a highly significant fit and * a significant fit.

Tailings Sites

Model X' P df X' P df

G eometric 1-67" 0.98 2 278.3 1 qO.01 36

Log Series 0.74" 0.39 1 4.02** 0.40 4 Tuncated Log Normal 1.5 1 ** 0.22 1 2.60** 0.65 4 Broken Stick 3.08** 0.08 1 33.96 ~ 0 . 0 1 5

Geometric 548.03 4 . 0 1 29 21.87** 0.35 20 Log Seriw 16.15 ~ 0 . 0 1 2 19.47 <0.01 2 Tuncated Log Normal 29.09 <0.0 1 3 5.83** 0.12 3 Broken Stick 37.71 <0.01 4 12.01 0.01 3

VII. D. Discussion

Al1 four control sites supported sirnilar numbers of wandering spider species

suggesting that they share roughly equivalent healthy States. The tailings sites on the

other hand, had variable numbers of species; most notably, Site A supported 22 fewer

species than Site C.

Ahhough each study site was sarnpled in an identicai rnanner, a cornparison of

wandering spider species richwss between sites is not as straightfonvard as might £ka be

presumed. Keeping track of the numbers of singletons and doubletons CO Uected fkom

each site, instead of simply reporting the numbers of species revealed valuable

information (Figure 32). Probability dictates that if many singletons and doubletons are

found in a CO llection, there are likely more species to be fourad. Variable habitat space

and quality among and within sites necessarily dictates variable sampling regimes if the

goal is to collect ail potential species thoroughiy . Species ric hness extrapolation

techniques such as the Abundance-based Coverage Estimator (ACE) allow for a closer

estimate of the true number of species within a collection area regardles of the sampling

intensity or the particular characteristics of the site. thus allowing for more accurate site

cornparisons.

The control sites in this study were assumed to represent the equilibrium or

heakhy state. ïhis assumption will be discussed in detail later. The ACE for the control

sites climbed from 45.7 + 1.1 species (Site 1) to 48.8 + 2.2 species (Site 2), peaked at

65.1 & 7.3 species (Site 3), then fell to 36.0 f 1.6 species (Site 4) fiom youngest to olden

site (Figure 33). Although Site 4 was not selected with the same criteria as the other

control sites (see Generai Methods), it is assumed to be older and more mature. This

successional pattern, with an older, more mature stage supporthg fewer species than a

younger, earlier successional stage, is common (van der Merwe et al. 1996, Frank and

Nentwig 1995, Bultman 1980, Neumann 1973, Huhta 1971, Williams 1962, and Lowrie

1948). Huhta (1971) explained this successiod pattern in relation to changing

microclimatic conditions including solar radiation, moisture, spatial structure, as well as

in relation to changing biological conditions including nutrition, predation and

cornpetit ion.

Tailings Sites A, B, C, to D followed the same pattem in eaimated wandering

spider species richness as observed fiom control Sites 1, 2, 3 to 4. Estimated species

richness climbed kom 25.9 f 3.8 species (Site A) to 40.7 2 3.3 species (Site B), peaked at

57.0 f 2.9 (Site C), then fell to 42.7 t 3.8 (Site D) (Figure 33). The bare, unreclaimed

tailings Site A, with an eairnated 25.9 f 3.8 species, illustrates the potential maximum

capacity of wanderuig spiders on u~nanaged sites. A cornparison of interest to the INCO

Ltd. reclamation program is that between Sites D and 3, which shared very similar ages

and physiognornies: the difference in the tme number of species between these sites

ranged fiom 1 1.3 to 33.5, with Site 3 supporting the greater nurnber of species. This

suggests that there are important limiting biological and/or physical factors on the oldea

reclaimed tailings site. Although the overall patterns of true wandering spider species

richness were similar between comrol and tailings site, the control sites supported greater

numbers of species. This suggests that the control sites may be heaithier than the tailings

sites in providing greater oppomullty for spider establishment.

The standard error of the mean eairnated nurnber of species in each site (Figure

33) was calculated with pooled monthly collections acting as samples. A wide margin of

error is hence an indication of both pronounced seasonal changes in the proportion of

singletons and an irnprecise estirnate for the true nurnber of species. The greatest error

margin was that calculated for Site 3 at 65.1 k 7.3 species and indeed, the pooled

collections during the months of May and Augua contained more singletons than during

Jme and July (not reported). The standard emrs of the mean estimated number of

species for the remaining sites were narrow, indicating precise and more reliable

estimates in addition to weU distributed singletons over the course of the summer. This

may suggea that spider arrivai and establishment on these tailings and control sites

occurs at rektive ly constant rates throughout the summer.

The jack-knifed Shannon-Wiener diversity and evenness indices for the

wandering spider cornmunities on both the taiiings and control sites did not fonn a

pattern similar to their species richness estimates. One highly abundant species relative to

the abundances of ail other species in a community may greatly depress that cornrnudy 's

diversity even though maay constituent species may be present. This was the case for

control Site 1. It supported a large number of species (Figure 32) yet a relatively low

Shannon- Wiener diversity index (Figure 34). The extreme abundance of Pardosa moesta

(1399 individuals, see Table 2), far surpassed the 2d most abundance species, Trochosa

terricoh (2 1 7 individuals), and other species. This explains the relatively low Shannon-

Wiener evenness index (E = 0.348 * 0.01 8, Figure 35). On the other hand, well-

pro port ioned abundances may drast ical ly increase a CO mmunity ' s Shannon- W iener

diversity regardless of the number of species. This was the case for tailings Site A.

Because the 25 individuals collected fiom this site were evenly distributeci among 16

species (with E = 0.898 * 0.017, Figure 39), the resultant Shannon-Wiener diversity of

this community was the greatest of ail tailings sites (Figure 34). The remaining reckimed

tailings sites and their control sites ali had a similar diversity of wandering spiders, partly

attributed to sirnilar evenness measures.

The results provided by the Shannon-Wiener diversity and evenness indices

superficially indicate that the health of tailings communities is superior to that of

communities removed fiom the effects of smelting and tailings deposition However. if

the goal of the restoration project is to create habitats in concert with their surroundings,

perhaps the findings should be more comparable. In other words, perhaps the grasslands

tailings site should be supponing a community of wandering spiders where one or two

species dominate and the remaining are locally 'rare'. as was the case for the grassland

control site, Site 1. Spider diversity studies in agricultural settings indicate that species

richness and overall abundance decreases fiom the edge of fields, where microclimatic

conditions are favourable, to the centre of fields where the conditions are more extreme

(Frank and Nentwig 1995; Alderweireldt 1994, 1 989; Bishop and Reichert 1990). In

addition, Alderweireldt (1989) found that only one or two species of Lycosidae dominate

the centre of maize and Italian rye grass fields in Belgium. Duffey (1 962) also found that

habitats with the greatest vegetation diversity support a larger number of spider species

whereas the most compact vegetation structure, such as Festuca turf, has the highea

number of individuais per unit area. Although diversity indices were not calculated in

these studies, these results mggest that Shannon-Wiener diversity and evenness values

decrease fiom a field's edge to its centre. The index values in the present study indicate

that the grasslands tailings site may contain a stable and varied cornrnunîty of spiders,

when an unstable and transitional cornmunity may be expected.

The Monsita-Hom index of community similarity is a quantitative masure that

takes into account both the number of species and their abundance and calculates a

percent similarity between a pair of sites. Because the measure is presently limited in this

way, a matrix of similarities is al1 that can be constmcted. This rnakes analysis of

cornmunit y similarit y rather taxing . In very general te-, the cornmunit ies of wandering

spiders on later successional reclaimed tailings sites seemed moa similar to control sites

(Table 6), whereas earlier successional tailings sites did not seem to share this similarity.

This suggests that only reclaimed tailings in the relatively later stages of succession are

able to support resident populations of spiders similar to more natural conditions. In

addition to this, the community of spiders on Sites A and B appeared to be more similar

to adjacent tailings Sites C and D than to their control sites (Table 6). Al1 of these

considerations rnay allude to a general avenue of establishment: wandering spiders may

becorne success fully founded on later successional reclaimed tailings subsequent ly

establishing themselves on early successional tailings sites.

The Bray-Curt is S imilarity Coefficient al10 ws for a visual interpretation of spider

community similarities. The dendogram constnicted f?om clusters of sites clearly

illustrated that the later successional tailings communities (Sites C and D) were moa

similar to an early successional control site (Site 1). This agrees well with the

mathemat icall y superior Mo risitaoHom similar it y mesures. This suggests that either

these communities are under stress (and hence reverthg to an early successional stage) or

may simply mean that approximately thirty years are required before reclaimed tailhg

sites are able to support communities sirnilar to that of more natural, outlying areas.

Coyle (1% l), in his study of the effects of clear-cutting on mature forests in Norîh

Carolina found that the communities of spiders in clear-cut forests were moa sirnilar to

each other using the Bray-Cmtis measure than they were to the uncut forest. It is

expected then that late successional communities should be different fkom early

successional communities. The k e n tdings site supported a cornrnunity of wandering

spiders only 8% similar to a cluster formed of all other commUILities, strongly suggeaing

that there are many factors W i n g wandering spider establishment.

If the discussion of tailings ecosystem health was limited to wandering spider

species richness, the conclusion would indicate superficially that tailings sites are as

healthy or are very similar to the equilibrium state as those represented by their control

site counterparts. Ho wever, if abundance models are invest igated, increasing ly

meaningful conclusions about tailings ecosystem health may be drawn.

Al1 of the abundance distributions of wandering spiders collected fiom the control

sites best fit the tnincated lognormal model (Table 7). Accordhg to the biological

explanation of this modeL the control sites each contain large, varied and mature

communities of wandering spiders and the available resources are more evenly

distributed among the taxa than would be assumed under either the geometnc or log

series models (Magurran 1988). It was anticipated however, that the control sites species

abundance distribut ions would exhibit a successional pattem fkom the geomeîric model

(Site l), to the log series model (Site 2 and perhaps Site 3), through to the truncated

lognormal model (perhaps Site 3 and Site 4). In other words, it was hypothesized that

wandering spider spec ies abundances would re flect increasing eco s ystem cornplexit y

from Site 1 to 4. These sites are either inappropriate standards to gauge INCO Ltd.

tailings ecosystem health, or the abundance models are insufficiently precise tools, or the

interpretation of assumed abundance patterns requires reexamination.

In order to ascertain a baseline health for each of the tailings sites accurately, the

ideal approach would be to locate at least three control sites per tailings site. However,

the volume of specimens and the collecting time necessary to maintain such an endeavour

wouid serîously jeopardize the b i t ion of this project . Consequent ly, these control sites

must remain as a . approximation, albeit imperfect, to the tailings sites. The species

abundance models codd dm be criticized as king indequate or flawed. Dewdney

(1997), while proposing his own "logistic" m d e l recently criticized the truncated

lognormal model as statistically invalid. Regardless of the strengths or weaknesses of the

truncated lognormal model it continues to enjoy a rich bed of supponing literature with

concomitant biological interpretations. Dewdney (1997) himself points out that the

truncation procedure merely approximates his potentially superior logistic model.

AssumSig that these control sites and the four species abundance models are mfncient

for the purpose of this study, the remaining quandary is the observed species abundance

pattern.

Spiders are known to behave spontaneously to environmental change yet are

quick to colonize sites once conditions have ameliorated (Collins et al. 1996, Haskins and

Shaddy 1986, Coyle 198 1, Huhta 1971). It is not surprishg then to h d that the

populations of wandering spiders on the control sites are best represented by the

truncated lognormal model. The post-fie control sites have presumably recovered to a

sufficient state for wandering spider re-colonization and the control site not afTected by

fne is presumably diverse enough to support a large and varied community. On the other

hand, a population not represented by the truncated lognormal model is an indication of a

very recent disturbance or an indication that only a few environmental andlor biological

factors are contributing to the observed distribution pattern.

The model of best fit for the wandering spiders on tailings Sites A was the

geometric model (x' = 1.67, df = 2, P = 0.98). Because all of the models significantly fit

the abundance distribution of wandering spiders in Site A, it may be that the very low

numbers of species and their abundances restrict any one model king a superior fit than

another. The situation rnay be similar to fïtting a line of best fit through highly scattered

data points; an infinite nurnber of lines are possible with no one line king a more a

accurate representation than another. Site A was a hanh environment because of the

abrasive nature of swirling, dry tailing s dut, elevated temperatures throughout the

daylight hours, and, without the moderathg effects of mil, iitter, and vegetation, likely

suEers fi0 m unseasonab1 y cool temperatures at night . Consequent ly, the geometric model

may aptly demonstrate a simple systern dominated by a few biological and physical

factors.

The community of wandering spiders on the grassland tailings Site B was best

modeled by the truncated lognormal model (X2 = 2.60, df = 4, P = 0.65), which indicates

a varied and complex nanual systea It was anticipated however, that Site B would be

best represented by either the geometric model or the log series model as these models

are typical of early successional habitats.

The community of wandering spiders on the rnixed wood tailings Site C was not

modeled by any of the four species abundance models. This is largely due to the elevated

abundance of species ranked 2nd to 4" and 5" to 7", forming two plateaux in the

abundance plot (Figure 38). Because a model cannot be fitted to the data, a biological

explanation for this distribution is not available. This odd distribution pattern may be an

indication of two or more overlapping communities, forrning an overall abundance

pattern that eludes modeling; or a fifih model, positioned between two of the four main

rnodels might form a better fit. The logistic mode1 (Dewdney 1997) did significantly fit

this site's wandering spider species abundance data (not reported), whic h po tentially

supports the author's daim for its superiority.

It was surprishg to see that the wandering spider community on the oldea tailings

site, presurned to be a varied, diverse, and complex site, was best modeled by the

geometnc mode1 (x2 = 2 1.87, df = 20, P = 0.3 5). In light of the A Abundance-based

Coverage Estimation of the true number of species however, the geometric model may fit

the species abundance distribution justly. Because the numbers of species and their

abundances were far greater than that observed for Site A, there is also little chance that

the abundance distribution has k e n incorrectly classified. Similar to the conditions in

Site A, Site D rnay also be dominated by a few factors or has suffered a recent

disturbance. Both possibilities are likely. This site may be dominated by a few fkcton due

in large part to the thin and ofien absent coniferous litter layer. It is also positioned

adjacent to areas of active tailings deposition fiom which tailings dust may fieely blow

and it bas suffered also fiom accidental tailings pipe rupture.

The species abundance model cornparison of W e s t to the MC0 Ltd.

reclamatioa program, between Site D and control Site 3, also revealed that there may be

important environmental and/or biological factors limiting the development of the oldest

reclairned tailings site. Site 3 was best modeled by the truncated lognormal model,

indicating a later successional state, and tailings Site D was best modeled by the

geometric series, indicating an early successional state. Detennining what the limiting

factors are is outside the scope of this data set and requires additional research.

It is not an easy task to assess the health of an ecosystem. The approach taken in

this study is w t cornmon and is one of the fïrst of its kind to be applied to tailings

ecosystems. The bewfits of using a diverse group of animals to examine the

sustainability of an ecosystem include the abilÎty to take advantage of mathematical tools

that are weil documented in the literature. A recurring result of the application of a few of

these tools was the rnarked differences beiween the community of spiders on the oldest

reclaimed taiiings site, Site D, to that of outlying control sites. Should al1 the INCO Ltd.

reclaimed tailings sites reach this aate after thirty years, the reclaniation procedure may

need improvement.

VIII. GENERAL DISCUSSION

INCO Ltd. officials at Copper Ciiff, Ontario are hced with a complex ecological

problem It is hoped that someday, their tailings will be reconstructed as self-sustainhg

ecosysterns in harmony with their surroundings. Ho wever, Little work has been directed at

how to assess the re-greening process and little to no effort has been directed at

monitoring either the health, or how quickly these systems are returning to a more natural

state.

Analyzing the diversity and succession of communities is one of the moa

mathematically difficuh tasks with which an ecologist can be faced. When land

management decisions hinge on the interpretation O f results, the dificuit y is exacerbated.

Historically, diversity and succession were exarnined by compiling species lists and

attempting to ider diversity and/or species-turnover. This approach is still valuable

because individual species in a data set may be exarnined in terms of their reg ional and

local distributions and in terms of their habitat preferences. It was soon realized however,

that much information could be gleaned fiom divenity data sets if more sophisticated

tools were applied. Unfominately, the present diffcuity in examining diversity and

succession in their purely mathematical senses is choosing the best tool among a

consternating variety of measures and indices. When an index was deemed inadequate,

ecolog ists develo ped their own measure to bea represent their data (Magurran 1 988).

This has ied to a large number of indices and measures; Mme are mathematically sound

but most are specific to particula. data sets. There is always a temptation to try out many

indices and models on a set of diversity data. In most cases, it is most economical and

informative to restrict the aoalysis to one or a few of the more commonly adopted

measures. S implifying an andysis in such a manner provides the most concise re ference

for policy-maken and land managers. The tools utilized in this study are only a handfùl

of measures that exist in the literature. They are al useful because of either the

information they provide, such as the Abundance-based Coverage Estimator (ACE), the

species abundance distriiut ion models, and the cornmunity similanty measures, or simply

kcause of their popuhnty, such as the Shannon-Wiener diversity and evenness indices.

Each of these tools provided distinct insights into the health of INCO Ltd. tailings

ecosystems by examining w a n d e ~ g spider communities at three leveis. The presence of

this cornmunity of spiders infers the existence of diverse food webs, which in turn hints at

ecosystem sustainability.

A thorough checklist of Manitoba spiders compiled by Aitchison-Bene11 and

Dondale (1 990) included 147 wandering spiders. Other checklists in closer proxirnity to

the Sudbury region coilectively included 129 species (Platnick and Dondale 1992;

Dondale and Redner 1990, 1982, 1978; Freitag et al. 1982; Martin 1965; and Kurata

1943). The total number of species of wandering spiders actually collected in the present

study was 74. Wandering spider geographical ranges (Dondale and Redner 1990, 1982,

1 978; and Platnick and Dondale 1 992) dlustrated t hat most species in the present study

reside in the region. Exceptions to these reports cm possibly be attributed to an

expansion of geographical ranges but more likely ascribed to incomplete records. The

checklist assembled for this thesis (Table 2) provides an invaluable and very current

addition to Canadian wandering spider biogeography . An assessrnent of habitat

preferences was attempted for ten of the most abundant species. This revealed that

previously published habitat preferences might be insufficient. Although these remhs

irnprove the understanding of habitat requirements for a few species, this analysis is by

no means complete. The abundance of these spiders mut be examined in many more

habitats in many more geographical reg ions.

The ACE measure and the Shannon-Wiener diversity and evenness indices

developed images of wandering spider a diversity, or species richness at a local leveL

The Morisita-Hom md Bray-Curtis community sirnilarity measures developed images of

wandering spider P diversity, or species tum-over. Finally, at a much larger scale, the

abstract and theoretical species abundance distribution models provided insight into the

tailings and control site ecosystems.

Large merences between obse~ed and potential numbers of species c m be

attributed to working at limited biogeographical scales and/or to the sampiing regixne. In

order to make accurate inferences about the health of tailings sites, an estimate of the

total possible number of species in the control sites and the tailings sites became

necessary. The Abundance-based Coverage Estimator (ACE) (Chadzon et al. in press)

was utilized to estimate the 'true' number of species. This tool takes into account

singletons and doubletons in collections, that is, species represented by one and two

individuals, respectively. Each of the tailings and control sites experienced an

approximately two-fold increase fiom the actual to the estimated number of species,

while rernaining below the 129 expected number of species determined by published

checklists. Although the successional patterns in wandering spider species richness were

similar between control and tailings sites, control sites collectively supported a greater

estimated number of species. This suggests that thete rnay be important environmental

andor biological limiting factors in tailings ecosystems.

A popular movement in diversity research is to encapsulate al1 the information

about species richness and abundance into a single çcalar number, thus simplifying

computation. The Shannon-Wiener diversity index (H') is the most commonly appïied

"information" measures. Ironically, the interpretation of this index is in stark contrast to

its computational sirnplicity. ïhis index was used in the present study merely because it

is so widely (and indiscriminately) applied to diversity data. Because the assumptions

governing the use of H' could not be satisfactorily met, parametric statistics could not be

executed. In their place, the jack-knifmg procedure was used to both irnprove their

estimation and to assign confidence intervals. Unexpectedly, the barren tailings site

supported the greatest diversity of wandering spiders on INCO Ltd. tailings (Figure 34).

This, however, is largeIy a result of an elevated evenness index (Figure 35). The

grasslands control site (Site 1) supported the least diverse community of all sites (Figure

34) due to the extreme abundance of Pardosa moesta. Diversity studies in similar

grassland and agricultural settings have demonstrated that the dominance of a few species

in these habitats is common (Alderweireldt 1989, DufFey 1962). Consequently, the H' for

the community on the grasslands tailings site (Site B) should possibly have approximated

the H' for the comrnunity of spiders on Site 1.

Because these jack-knifed Shannon-Wiener indices were limited to a diversity

comparisons, two sirnilarity coefficients were implemented. ûne coefficient, the

Morisita-Hom index, is mathemat ically perfect (Ma- 1 988) but is present ly limited

to pair-wise cornparisons between sites without the ability to construct a dendogram. A

second coefficient, the Bray-Curtis measure, is commonly applied because such a

dendogram may be constructed. Both of these maures reveaied that only the later

successional reclaimed tailings spider communities were most similar to control site

communities (Table 6 and Figure 36). More specifically, a cluster fonned by the spider

communities on tailings Sites C and D was most similar to the community on control Site

1 (Figure 36). This provided fbrther evidence that the later successional tailings habitats,

moa notably Site D, rnay be approaching an ecological state similar to outlying habitats

but that some fundamental differences remain.

A more powefil tool than a and p diveaity measures, which encapsulates aii the

infonnat ion about a community 's numerical characterist ics, is species abundance

distribution model-Ming. Just as there are many species richness estimators, there are

also many models fiom which to choose. However, distributions are rnost cornrnonly

analyzed using four main rnodels (Magurran 1988). These are, in order of increasing

abundance uniformity: the geometric model the log series modeL the lognormal model,

and the bro ken stick model (Figure 30). These models also describe a gradient fkom early

to late succession (Magurran 1988). AH of the control site communities of spiders best fit

a tmcated version of the lognormal distribution indicating large, varied, mature, and

natural communities (Table 7). This was not anticipated since it was assumed that at least

the grasslands control site (Site 1) would support a mmrnunity in the early stages of

succession. Ho wever, spider populations are kno wn to behave spontaneously as a

response to an environmental change (Stork and Samways 1995) so perhaps these control

sites were more ecologically stable than the tailings sites. Without long term monitoring

on either the tailing s or control sites, these çtatements remah speculat ive. The wandering

spider communities on both the k e n tailings site (Site A) and the oldest, forested

tdings site (Site D) best fit the geometric model (Table 8) indicating that these sites are

early successional. This suggests that either Site D is stressed and bas reverted to an

earlier successional state or it bas stabilized in its present condition, a condition

uncharacteristic of a s tn ic tdy similar site (Site 3) (see Site Descriptions). From a land

management perspective, fùrther rehabilitation may be needed to reclaim this site.

Strangely, the community of spiders on the most botanically diverse tailings site did not

fît any of the four main species abundance models (Table 8). It is therefore impossible to

make Uiferences about the successional state of this habitat. A greater sampling effort

might have benefited the model-ming analysis for Site C but unfortunately, this would

have obscured community comparisons. In order to make sound cornparisons. the

sampling efforts at each site must be identical.

Up unt il this point , predominant ly mathemat ical CO nsiderat io ns have ken

developed to assist the cornparison between control and tailings sites; little thought has

k e n directed at the biology of the wandering spider guild and how this might aid site

comparisons. This discussion has been saved until now because it relies heavily on

literature and conjecture but rnainly because it will provide direction for future studies.

Several eco log ical parameters determine the species ric hness and abundance of

wandering spiders. S o m of these paramet ers include temperature preferenda, humidil y

preferenda, and litter complexity.

Perhaps the moa fiequent ly cited exp lanat ion for invertebrate distribut ions is their

temperature preferenda. As with insects, spider development, growth, and reproduction

are iinked to temperature and, as a result, species prefer specific temperature ranges.

Developmental processes are essentially coupled to metabolism: increasing temperatures

n o d y increase development and growth rates and shorten developmental phases,

respectively (Pulz 1 987). Thermal influences on growth are more dificult to assess

because of interfering frictors such as humidity and food (Workman 1978). Reproduction

rnay be affected in terrns of the copulation behaviour (Davis 1989), copulation duration

(Costa and Sotelo 1 984), pro pensit y to ovipo sit, the oviposit ion intervais, and the nurnber

of egg sacs (Li and Iac kson 1 996). Drastically different surface temperatures between

tailings and control sites leading to variable development, growth, and reproduction may

account for some variation in cursorial spider richness andlor abundance. Dondale and

Binns (1 977) found that the population densities of spiders in an Ontario meadow are

large1 y linked to cumulative seasonal temperature, although they rnake no mention of

physiological links. However, Workman (1978) found that flucniating temperatures had a

significantly decreasing effect on growth rate and instar duration. Ail of these

physiological effects may account for variable temperature preferenda such as those

found for two species of Lycosidae by Sevacherian and Lowrie (1972). Wandering

spiders may simply avoid the barren tailings site with presumably elevated temperatures

during the &y and depressed temperatures at night.

Spiders are resistant to desiccation, largely due to cuticular lipids. However, there

are several bodily regions where water is fieely lost (Pulz 1987) and spiders must replace

this water via soi1 capillary drinking (Pany 1954) or by other methods. It stands to reason

that water availability rnay limit the local distribution patterns of wandering spiders.

Tailiogs sites with notably dry surfaces were Sites A, B, and, C. Site A was without

vegetation, Site B supponed sparsely sown grasses, and Site C was largely without a liner

lay er . Temperature, humidity and wind couectively affect the propensity for immature

spiderlings to kiloon (Richter 1 970). Ballooning is imrnediately resumed if microc limate

conditions are not favourable upon landing. If the ballooning rate could be quantifiecl,

this might also provide an avenue to compare tailings to control site habitats.

Perhaps the single moa important parameter affecthg distribution patterns of

wandering spiders is the liner layer, or lack thereof. More specifically, Uetz (1979) and

Bultrnan and Uetz (1 984) found that wandering spiders are particularly afTected by the

complexity of the litter. Deciduous liner, with greater interstitial spaces, supports a

greater diversity of wandering spiders than coniferous litter. Litter complexity in these

studies was measured using an index devised by Uetz ( 1974). These hdings demonstrate

how valuable wandering spiders are in assessing the humus cap of iNCO Ltd. reclaimed

tailings. Litter is also indirectly tied to plant popuiations and communities (Facelli and

Pickett 1 99 1 ), complet h g the relationship between wandering spiders, litter, and plant

communities. In addition to the importance of interstitial spaces, litter also provides a

sexual display arena whereby male wolf spiders stridulate to announce their presence and

to attract a mate (Stratton and Uetz 1983). Liner that is more complex allows for greater

sound amplification. Liner wmplexit y also affecteci the microhabitat selection, activit y

patterns, and density of Schirocosa ocreata ( Walkenaer) (L ycosidae) (Cady 1 9 84).

The abundance and species richness of the wandering spider guild have k e n

successfully used to examine the health and succession of INCO Ltd. reclaimed tailings

ecosystems. Although reclaimed tailings ecosystems are recoverhg in a manner

approximating control sites of similar ages, sizes, and physiognomies, there remain

distinct differences in their wandering spider cornmunities; these communities king

indicators of ecosystem health and vigor. The determination of these differences would

requue continuous monitoring with the additional collection of micrometeorological data.

The latter would greatly assist in determinhg the functional differences arnong individual

reclaimed tailings sites and between tailings sites and suitable control sites.

Restoration ecology is a major growing field of research and management that is

expected to continue to expand in importance. Restoration, however, is a last resort and is

far more expensive than maintainhg an intact ecosystem in the fia place. There must be

a critical assessrnent and monitoring of an area to see whether it is simply being "re-

greened" or whether a fùnctional ecosystem is indeed k i n g reconstituted. This audy

represents an important step toward assessing and monitoring the INCO Ltd. tailings

reclamation megaproject fiom a purely biological perspective. More studies are

increasingly necessary as INCO Ltd. nears the decommissioning phase of its history.

Appendix A. Plant species on RJCO Ltd. tailings sites.

Species Site type Scientific name Common name

Grass

Shmb Shmb Grass Grass Grass Moss

Tree Tree Tree Tree Shmb Shmb Shmb Herb Her b Herb Herb Sedge Grass Grass Grass Grass Moss Moss

Lichen Lichen

Tree Tree Herb Grass Grass Grass Grass Grass Moss

Deschampsia caespitosu ( L . ) Beauv.

Pinus banksiana Lamb. Betula papyrfera Marsh. Festuca mbra L. Poa compressa L. Agrostis gigantea Roth Pohlia nutam (Headw.) Lindb.

Pinus banksiana Lamb. Pinus resinosa Ait. Populus tremuloides Mic hx. Ro binia pseudoacacia L. Pinus resinosa Ait. Popuius tremuloides Michx Robinia pseudoacacia L. Lotus cornidatus L. Rumex acetoseila L. Hieracium aurantiacum L. Cirsium anense (L.) Scop. Carex scoparia Schk. Poo compressa L. Agrostis gigantea Roth Bromus inemis Leyss. Festuca rubra L. Polytrichum sp. Pohlia nutans (Hedw.) L ind b. Cladonia rei Scbaerer Cladonia cristatella Tuck.

Pinus banksiana Lamb. Picea gIuuca (Moench) Voss Epilobiun angus~~~olium L. Deschampsia caespitosa (L.) Beauv. Brornus inennis Leyss. Pou compressa L. Poa pratemis L. Festuca rubra L. Pohlia nutans (Hedw.) Lindb.

Jack Pine White Birch Red Fescue Canada B luegrass Redtop

Jack Pine Red Pine Trembling Aspen Black Locust Red Pine Trembling Aspen Black Locust Birdsfoot Trefoil Sheep Sorrel Orange Hawkweed Canada Thistle Broom Sedge Canada Bluegrass Redtop Brome Grass Red Fescue

Jack Pine White Spnice Fireweed Tufked hairgrass Brome grass Canada BIuegrass Kentucky B luegrass Red Fescue

Appendix B.

Importance values (IV %) of ground cover categories within two transects on ail MC0

Ltd. tailings sites surveyed between May and August 1996.

Tailings Site A

Survey Date Transect Ground Cover Scienti fic Name Importance Value Num ber Category IV (%)

May 23, 1996 1

June 24,1996

July 26, 1996

August 22, i 996

Bare Soil Rock Tufied Hairgrass

Bare Soil Tufted Hairgrass

Bare Soil Rock Tufleci Hairgrass

Bare Soil Tufleci Hairgrass

Bare Soil Rock Tufted Hairgrass

Bare Soil Tuftecl Hairgrass

Bare Soil Rock Tufied Hairgrass

Deschampsia caespitosa

Deschpsia caespitosa

Deschpsia cuespitosa

Desc hampsia cuespitosa

93 .O7 6.93

Deschampsia caespimsa O. 10

Bare Soil Tufted Hairgrass

Tailings Site B

Survey Date Transect Ground Cover Scientific Name Importance Value Nurn ber Category IV (%)

May 23, 1996 t Dead Grass 40.6 Canada Bluegrass (live) Poo compressa 2 1 .O Red Fescue Fesruça rubru 16.7 Moss Pohlia nutans 14.5 Bare Soil 7.3

June 3. 1996 2 Moss Pohlia nutam 56.7 Red Fescue Festuca rubra 29.2 Bare Soil 10.0 Redtop (live) Agrosris gigantea 4.2

June 27. 1996 1 Litter (Dead Grass) 47.02 Bare Soi1 19.4 Canada Bluegrass Poa compressa t 5.67 Red Fescue Festuca rubra 10.45 Moss Pohlia nutans 4.38 Redtop Agrostis gigamea 2.99

2 Moss Pohlia mtam 54.84 Red Fescue Fesruca rubra 29.84 Bare Soi1 12.90 Canada BI uegrass Poa compressa 2.42 Redtop Agrosiis giganfea 0.16

July 25. 1996 1 Litter @ead Grass) 50.38 Red Fescue Fesruca rtrbra 13.74 Canada Bluegrass Poa compressa 13.74 Bare Soi1 9.16 Moss Pohliu mtans 6.87 Redtop rigrosris gigantea 6.1 1

2 Moss Pohlia M C I ~ Red Fescue Festuca mbta Bare Soil Canada Bluegrass Poa compressa Liner (Dead Grass)

August 23, 1996 1 Litter (Dead ûrass) 46.1 O Moss Pohiia nutans 3 1.28 Canada Bluegrass Poa compressa 15.60 Red Fescue Festuca mbra 8.5 1 Bare Soi1 6.74 Redtop Agrostis gigantec 1.42

2 Moss PoWia nutans 57.97 Red Fescue Fesîuca rubra 17.39 Litter (Dead Grass) 9.42 Bare Soi1 8.70 Canada Bluegrass Poa compressa 6.52 W t o p Agrostis giganfea 0.07

Tailings Site C

SurveyDate Transect Ground Cover Scientific Narne Importance Value Num ber Category IV (%)

May 23, 1996 1 Moss (mix)

Canada Bluegrass Moss Redtop (live) Litter (Trembling Aspen leaves) Moss Birdsfoot Trefoil (dead) Birdsfoot Trefoil (live) Brome Grass Trem bl ing Aspen (< l m) Lichen Lichen

May 27, 1996 2 Litter (Black Locust leaves, Jack Phe needles) Moss SedgdGrass (dead)

Broom Sedge Sheep Sorrel Redtop (live) Black Locust (< 1 m) Red Pine (< 1 m) Birdsfoot Trefoil

June 24, 1996 1 Moss Canada Biuegrass Litter (Jack Pine, Aspen) Moss Birdsfoot Trefoil (live) Redtop Birdsfoot Trefoil (dead) Dead Grass Orange Hawkweed Brome Grass Broom Sedge Sheep Sorrel

2 Litter (Black Locust, Jack Pine) Moss Broom Sedge Sheep Sorrel Redtop Moss Birdsfoot Trefoil Canada BLuegrass

Pohlia nutans/ Po!vtrichum sp. Poo compressa Polytrichum sp. Agrostis gigantea

Pohlia rtutans Lotus cornicularis Lotus corniculmus Brornus inennis Populus tremuioi&s Cladonia mi Cladonia cristatella

Pohlia rtutrmr

Carex scoparia Rumex acetosefia Agrostis gigantea Ro binia pseucbacacia Pinus resinosa Lotus corniculatur

Polyrtichum sp. Poo compressa

Pohiia nutans Lotw cornicuIatf(s Agrost is gïgantea Lotus cornlcuIatur

Hieracium auraniiacum Brumus inennis car= scop~~la R u m a aceroselia

Pohlia nutans Carex scopana Rmex: mosel la Agrosris grgantea Polyrnchum sp. Lom corniculcrlzls Pou compressa -

Red Fescue ~esnrca &a

Brome Grass Trem bl h g Aspen (C 1 m)

July 26, 1996 1 M m Litter (Jack Pine, Poplar) Canada Bluegrass Birdsfoot Trefoil (live) Moss Redtop Orange Hawkweed Broom Sedge Brome Grass Canada Thistle Sheep SorreI Trem bling Aspen (< 1 m)

3 - Litter (Locust, Jack Pine, Dead Grass) Broom Sedge Moss Sheep Sorrel Redt op Birdsfoot Trefoil Moss Canada BI uegrass

August 23, 1996 I Moss Litter (Dead Grass, Jack Pine, Poplar) Birdsfoot Trefoi 1 Canada Biuegrass Moss Redtop Broom Sedge Orange bwkweed Brome Grass Trembling Aspen ( 4 m) Canada Thistle

3 - Lirter (Black Locust, Jack Pine, Dead Grass) Moss Sheep Sorrel Broom Sedge Canada Bluegrass Redtop Birdsfoot Trefoil Moss Bare Soi1 Brome Grass Black Locust (<lm)

Bromus i m i s Popdus tmmuloi&s

Polytrichum sp.

Poa compressu Lotus comiculatw Pohlia nutans Apsris giguntea Hieracium aaanîtacurn Carex scoparia Bromus inennis Cirsium mense Rumex acetosella Populus iremuloides

Carex scoparia Pohl ia mtm Ruma acefoseIh Agrosris giganrea Lotus cornicuIalu~ Polytrichum sp. Poa compressa

Polytrichum sp,

Luru c o m i c u l ~ Pou compressa Pohlia nutm Agrostis gigmea Carex scuparia Hieraciwn aurantiucum Bromus inermis Populw rremuloi&s Cirsium arvetrse

Pohlia m t m Rtunex ocetosellit Carex scoparia Pou cumpmssa Agrosris grgantea Lotus conicuIatus Polytrichum sp.

B r o m inemis Robinia ~seu&acacia

Tailings Site D

Survey Date Transect Ground Cover Scientific Name [niportance Value Nurn ber Category IV (%)

May 27, 1996 I

June 3,1996 2

June 27, 1996 1

July 25, 1996 1

Bare Soil Brome Grass (dead) Brome Grass (live) Litter (Jack pine needles) Kentucky Bluegrass Moss Red Fescue Canada Bluegrass

Bare Soil Moss Litter (Jack Pine needles) Kentucky B luegrass Brome Grass

Liner (Dead Grass, Jack Pine) Bare Soil Brome Grass Canada Bluegrass Kentucky Bluegrass Moss Fireweed

Bare Soil Litter (Dead Grass, Jack Pine ) Brome Gtass Moss Canada Bluegrass Kentucky Bluegrass Red Fescue

Litter (Dead Grass, Jack Pine) Bare Soil Brome Grass Canada Bluegrass Kentuch Bluegrass Moss Fireweed Tufied Hairgrass

Bare Soil Liner (Dead Grass, Jack Pine ) Brome Grass Canada Bluegrass Moss Kentucky Bluegrass

Bromus inermis Bromus inerntis

Poa praremis Pohlia nurm Festuca rubra Poa compressa

Pohlia m f m

Poa prafemis Bromus inermis

Brornus inermis Poa compressa Poo pratensis Pohlia mtm Epilobium angusti$oliurn

Brmus inennis Pohl ia mirans Poa compressa Poa pratemis Festucu rubra

Brornus inermis Poa compressa Poa prafensis Pohlia mtans Epilobium angui~rfolium Lkschampsia caespitusu

B r o m inermis Pua compressa Pohlia nutans Poa prafensis

August 23,1996 I Litter (Dead Grass. Jack Pine) Bare Soil Brome Grass Bromus inemis Canada Bluegrass Poa compressa Kentucky Bluegrass Poa pratensis Moss Pohlia mtm Fireweed Epilobium ungustifolium Tufted Hairgrass Deschumpsia caespitosa

2 Bare Soi1 Litîer (Dead Grass, Jack Pine ) Canada Bluegrass Poa compressa Brome Grass Bromu. i n m i s Moss Pohlia mitans

Appendix C.

Tree layer importance values (IV %) within two transects on MC0 Ltd. tailings Sites C

and D. Trees were not found on tailings Sites A and B.

Tree Species

Sites C C D D

Transect 1 Transect 2 Transect 1 Transect 2 (IV %) (IV %) (IV %) (IV %)

Pinus banksiana 85 Pinus resinosa 15 Picea g h c a Robinia pseudoacacia Populus tremtdoides

S h b layer importance values (IV %) within two transects on INCO Ltd. tailings Sites C

and D. Shmbs were not found on tailings Sites A and D.

Plant Species

Sites B B C C

Transect 1 Transect 2 Transect 1 Transect 2 (IV %) (IV %) (IV %) (IV %)

Pinus banksiana 50 Pinus resinosa Betulrr papyrifera 50 Ro binia pseudoacacia Populus tremuloides

IX. LITERATURE CITED

Adams. J. 1985. The definition and interpretation of gudd structure in ecological

cornmunities. J. Anbn Ecol. 54: 43-59.

Aitchison, C. W. l984b. The phenology of winter-active spiders. J. Arachnol. 12: 249-27 1 .

Aitchison, C. W. 1 984a. Lo w temperature feeding by winter-active spiders. L Arachnol.

12: 297-305.

Aitchison, C.W. 1978. Spiders active under snow in southem Canada. Syrnp. zool. Soc.

Lond. 42: 139- 148.

Aitchison-Beneii, C.W. and C.D. Dondale. 1990. A checklist of Manitoba spiders

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