UNIVERSITY OF LAGOS

FACULTY OF SCIENCE
DEPARTMENT OF ZOOLOGY
SEMINAR TOPIC: TERRSTRIAL ARTHROPODS AND THEIR ECOLOGICAL
IMPACTS
SUBMITTED BY
ALFRED ADESOJI SYLVESTER
MATRIC NO: 090809013
IN PARTIAL FULFILMENT TO THE REQUIREMENTS FOR THE AWARD OF B.Sc.
ECOTOXICOLOGY AND CONSERVATION
UNDER THE SUPERVISION OF
DOCTOR (MRS) OLALERU

TABLE OF CONTENT
CHAPTER ONE
INTRODUCTION
CHAPTER TWO
2.0 Ecological importance of terrestrial arthropods
CHAPTER THREE
Beneficial impact
Pollinators
Biological control
Burrowers
Decomposers and nutrient cyclers
Detrimental impact
Pests and disease vectors

CHAPTER FOUR
4.0Case studies
CHAPTER FIVE
5.0 Conclusion and recommendation
CHAPTER SIX
References

ABSTRACT
Arthropods are the most diverse phylum of animals on earth. They have been classified
into subphyla Myriapoda, Uniramia and Chelicerata. Terrestrial arthropods are
ecologically important as pollinators of flowering plants, biological control agents of
other insects, decomposers and nutrient recyclers. They are soil burrowers aiding in soil
aeration, and are major components of terrestrial food webs. Other beneficial uses of
terrestrial arthropods include fibre and medicine production. Some terrestrial arthropods
have been implicated as pests and disease vectors to both plant and animals, including
humans. This poses an immense burden on global health, economic and agricultural
productivity leading to the use of pesticides and other control measures. Efforts should be
taken to conserve terrestrial arthropods from the adverse effect of human activities.
Pesticide application should be target-specific to prevent the destruction of beneficial
arthropods.

INTRODUCTION
There are over 800,000 named species in the Phylum Arthropoda,
named from the Greek arthros (= jointed) and poda (= foot), including
the familiar arachnids, crustaceans, and insects, together with a host
of less familiar critters, like centipedes, millipedes and sea spiders. All
arthropods have jointed appendages. This evolutionary innovation is
probably the key to the stunning success of this diverse group. There

are about 1018 (10 billion billion) arthropods alive at any one time.
There are over three times as many species of arthropods as there are
of all other animals on Earth, and there may be millions more that we
haven't even discovered. Arthropods do everything with legs or
modified legs. They walk, they swim, they creep and crawl, they use
legs to sense with (the antennae), to bite and sting with, and even to
chew with. That's one reason arthropods look so alien when we see
them up close. They chew sideways, and it's all done with legs.
The insects are the most numerous of all terrestrial animals both in
terms of species and totalabundance. However, only a small fraction is
pestiferous (Ewuim, 2004). These insects are therefore strategic in the
welfare of man throughtheir activities. The beetles which constitute
about
two-thirds of all known insects, and about one-third of all known animal
species invariably, participate in various activities, resulting' in several
changes in the ecosystems. The beetles like other insects, oftenevolve
and exists as components of communities ofplants and other animals.
Most of the species areterrestrial even though some are aquatic. In
terms offood and feeding habits, many beetles are planteaters; some
are predacious with others beingscavengers, while some of them are
wood-borers. In
terrestrial ecosystems many of these herbivorous forms constitute
serious pests of crops and causingsignificant damage either directly or
even transmitting diseases even though some are beneficial
herbivours.
Variability in climatic variables such as temperature, humidity, rainfall,
litter amount and slope of the terrain and elevation, and soil nutrients
influence the abundance and diversity of the Arthropods (Samson et
al., 1997; Townsend et al., 2008). Arthropod diversity is also dependent
on habitat heterogeneity expressed as differences in litter pool and
food (e.g., palatable leaf litter, fruits, seeds, and herbivore frass),
toxins, (e.g., phenols and tannins) and structural complexity that
creates habitat (e.g. branches and leaf litter depth) (William et al.,
2000; Dominy et al., 2003; Kaspari et al., 2003). In Rwanda, forest
plantations cover 10% of national territory, of which Eucalyptus
plantation covers 65% (Nsabimana et al., 2008). Thought it is know
that forest ecosystems conserve largest biodiversity, arthropod
diversity has never been studied in Rwanda, suggesting a need for
studying litter arthropods under these plantation species. In spite of
the generalized use of Eucalyptus in forestry in many parts of the
World, there is little information on its impact on the invertebrate fauna
(Florence, 1985; Da Silva et al., 2008). In addition, theresults of
previous studies carried out throughout the World, were not conclusive
about the influence Eucalyptus plantations on the abundance of
invertebrates (Chander and Goyal, 1995).

Soil biological and chemical properties as well as habitat conditions

alter drastically when there is a conversion from natural to agricultural
habitat. Curry (1998) and Lee (1991) highlighted frequent tillage and
use of
agrochemicals as having major impact on soil organisms and habitats.
Agricultural activities have positive or negative impact on abundance,
diversity and activity ofsoil fauna mostly following the changes in soil
temperature, moisture as well asquantity and quality of organic matter
(Hendrix and Edwards, 2004). Fields whichare more diverse, stable,
isolated and managed with low intensity have preferencefor ongoing
ecological processes compared with simple and disturbed agricultural
systems. Uncultivated habitats between fields could enhance species
diversity ofmany organism groups, and function as refuges (Lagerlof
and Wallin, 1993 andLagerlof et al., 2002). Spatial variability in
dispersion of soil organisms can be key to understanding the structure
and function of soil biodiversity (Ettema andWardle, 2002).
Soil arthropods are a vital link in the food chain as decomposer and
without
these organisms, nature would have no way of recycling organic
material on its own (Trombetti and Williams, 1999). The process of
decomposition are controlledlargely by soil arthropods in conjunction
with some soil invertebrates like protozoaand worms which also
contribute to the soil community by mixing, loosening andaerating soil
(Evans, 1992). Arthropods also serve as the largest prey base for
smallpredators, thus sustaining other arthropods. Without arthropods
most terrestrialecosystem would rapidly collapse (Iloba and Ekrakene,
2008). Arthropods havebeen able to fill every niche available in the
ecosystem.
The direct ecologicaleffects of these minute arthropods include the
reduction in the mass of organicmatter and microbial tissue. This is as
a result of their ingestion and assimilation ofsuch materials, their
respiration and excretion which is important in influencing oxygencarbon dioxide ratio of the soil and nutrient made available from the
breakdown of faecal pellets (Filser, 1995). Therefore, their secondary
productionturnover is very fundamental because this is the basis on
which the organisms, wayup the food chain are dependent upon. Many
soil arthropods, including Collembola, Oribatida, Isopoda or
Diplopoda,live a sedentary life in soil in close relationship with the
external conditions of theirecological niches. As a consequence, the
structure of the micro arthropodcommunity closely reflects the
environmental factors affecting the soil, includingthe impact of human
activities, and could be considered a bio indicator of soil health
(Knoeppa et al., 2000; Linden et al., 1994 and Sparling, 1997).

Heavyanthropogenic activities alter the equilibrium of the environment

owing to theintroduction of cultivated and inhabited areas, as well as
other human-inducedchange (Arroyo and Iturrondobeitia, 2006).
CLASSIFICATION OF INSECTS

CHAPTER TWO
ECOLOGICAL IMPORTANCE OF TERRESTRIAL ARTHROPODS
FOOD FOR WILDLIFE
Many different kinds of animals eat insects making them essential
parts of the food chain.Examples: amphibians, reptiles, birds, fish,
arthropods and other invertebrates, mammals (primates including
humans, insectivores focus almost exclusively on insectprey
(hedgehogs, shrews, tenrecs, moles).

CHAPTER THREE
BENEFICIAL IMPACTS
POLLINATION
Approximately on third of the worlds crop production depend directly
or indirectly onpollination by insects. The value of pollination in the
world, most of it by insects, has been estimated at about $117 billion
per year (Costanza et al. 1997).
Examples of crops: coffee, tea cacao (chocolate), vanilla citrus fruits,
nuts (almonds), vegetables, other fruit (figs, cranberries, cherries,
mangos), clover to name afew.
Examples of pollinators: The honey bee (Apis mellifera) is responsible
for the production of approximately $14 billion worth of agricultural
products in the United States in 1998, however other pollinates are
important pollinators as well including bumble bee which are usedto
pollinate tomatoes and cranberries; and carpenter bees which pollinate
wild and domestic tree fruits.

Insects that provide ecological services

There is a growing recognition of the value of the services derived from
their ecological functions that insects provide to human well-being
(Kremen and Chaplin-Kramer,2005; Losey and Vaughan,
2006 ,
2008 ). Th ese wide-ranging services include pollination,protection
from pests, decomposition, food for wildlife, and recreation through
activities such as insect observation. Losey and Vaughan ( 2006, 2008)
have calculated values for many of these services, and providean
informative comparison of conservation efforts that focus on ecological
services with those that focus on endangeredstatus (Loseyand
Vaughan,2008). They argue that the commonness and positive
economicimpact of insects that provide ecological services make them

excellent candidates for flagship species. Commonness may make

insects more familiar and less threatening,and thus better able to
overcome the charisma-challenges (Berenbaum,
2008 ) that
generally characterize insects. Additionally, if landowners are aware of
the benefits provided by insects, they may conduct land management
activities that benefit insects, provided that they are aware of
strategies that will accomplish conservation. Here, we focus on
conservation efforts directed toward pollinators (see Figures 3-5 ).
Pollinators have clear economic and even survival importance to
humans. Three quartersof all flowering plants, including most species
of food crops, rely on pollinators for fertilization, and decreases in wild
pollinator populations could have severe impacts on natural and
managed ecosystems. Pollinators are estimated to provide over $57
billion worth of services in the United States (Losey and Vaughan,
2006), and $217 billion worldwide (Gallai et al.,
2008). Many
organizations are promoting pollinator conservation in a variety of
ways that could havebroadconservationeffects. The Xerces Society,
dedicated to invertebrate conservation (Xerces Society, 2008 ), and
state extension agencies (e.g. Krichik, 1998; Riedl et al., 2006 ) have
published a varietyof information sheets for the public that describe
simple strategies for sup portingpollinators. These range from
strategies
for
avoiding
mortality
through
pest
management;supportingpollinators
with
appropriateplantings
in
rural,suburban and urban yards; and providing shelter. Th e North
American Pollinator Protection Campaign (NAPPC) is an alliance of
pollinator researchers, conservation and environmental groups, private
industry, and state and federal agencies in Canada, Mexico and the
United States (The Pollinator Partnership, 2008 ). NAPPC organizes
local, national, and international projects involving pollinator research,
education and awareness, conservation and restoration, special
partnershipinitiatives, and policies.

M.S. Guiney and K.S. Oberhauser / Terrestrial Arthropod Reviews 1 (2008) 111123

In late 2006, the United States National Research Council, at the

request of NAPPC, published a report that recognized the downward
long-term population trends for many North American pollinators and
called for increased efforts to monitor pollinators and improve
understanding of their basic ecology (NRC, 2006 ). While much of the
current data on pollinator decline focuses on the non-native honeybee
( Apis mellifera ), the report also notes declining numbers for several
wild bee species, especially bumblebees, as well as some butterfl ies,
bats, and hummingbirds. The report recommends support for research
aimed at the sustainable management of these populations, as well as
actions that encourage landowners to make habitats more pollinator
friendly By growing native plants and limiting pesticide use that could
affect pollinators.
The efforts of NAPPC and other groups focused on pollinator
conservation have recently paid off at the national level. Th e US Farm
Bill passed in 2008 includes measures that will help shape USDA
programs to protect and enhance the habitat of native and managed
pollinators. These include recommendations for pollinator friendly
plantings in field borders,hedgerowsand shelterbelts; corridors to
connect pollinator habitatpatches; and grants for researchthat will
addressthe impacts of pesticides onpollinators.

Karner blue buttery ( Lycaeides melissa samuelis) Kurt Adolfson, www.bluedotimages.com

The efforts of organizations addressing pollinator conservation have

also included avariety of public outreach eff orts, ranging from
pollinator postage stamps to school curricula. Through these eff orts,
increasing numbers of people are aware of the importanceof pollinators
to our survival.This awareness of the absolute necessity of a
naturalecological process carried out, in the main part, by insects, can
only benefit natural
system conservation.
3.1. Insects as Biological Control Agent.
Biological control means the action of parasitoids, predators, and
pathogens in maintaining other organisms density at a lower
averagethan would occur in their absence. In modern context,when we
are becoming aware of the harmful effects ofunilateral use of chemical
insecticides in various agriculturalfield or ecosystem, the role of insect
as biocontrol agent isimmensely vital. The first dramatic example of
deliberatebiological control was the importation of vedalia
ladybeetle,Rodolia cardinalis (Mulsant) in California, in 1888 to
controlthe cottony-cushion scale insect, Icerya purchasi Maskell
oncitrus [94]. Many insects act as potential predators andparasites of
destructive pests of insect-order Lepidoptera(Butterflies and Moths),
Diptera (Flies), and Orthoptera(Grasshoppers) [27]. Predators are
scattered in about 167 in the field of biocontrol. Thus, further studies
are required to explore successful insect species as biocontrol
agents.families of 14 orders of the class, Insecta [95]. Major

groups of entomophagous parasites belong to the order

Hymenoptera and family Tachinidae of the order Diptera.
The well-known parasitoids acting as potential biocontrol
agents are Ichneumonids, Chalcids, Proctotrupoids, and
Evanoids. The Tachinid flies species of Sturmia and Tachinia
Increase the capillary permeability
Immune response and tissue spread propertiesAntigenic
parasitize the insect pests like paddy armyworm and fruit
moth larvae [27]. Biological control effort against noxious
weed, Parthenium hysterophorus L. through the utilization
of insect species, Zygogramma bicolorata (Coleoptera:
Chrysomelidae) has also been advocated by Gautam [96].
However, only few species are well established and employed In the
field of biocontrol. Thus, further studies are required
to explore successful insect species as biocontrol agents.

Far less attention has been paid to augmentative biocontrol,whereby

native predators, herbivores, or parasites are used to control a target
pest species. Augmentative biocontrol is typically attempted with
insect predators or parasitoids, reared in insectaries, that are released
in massive numbers (inundative biocontrol) to suppress insect pests
(Collier and Van Steenwyk 2004). A major benefit of augmentative
biocontrol is that it does not involve introducing species outside their
native geographic ranges, and thereby avoids problems that
accompany species introductions. Its major limitation arises from the
inability of the native natural enemy to control the pest without being
augmented repeatedly. In other words, if the pest and control agent
coexist naturally, but the pest (by definition) is normally not controlled
by the agent, the effectiveness of the control agent seems in doubt.
Poor control under normal (unaugmented) conditions, however, can be
caused by low natural contact rates between the control agent and the
target. A key challenge for augmentative biocontrol is to alter the
abundance or distribution of the control agent to maximize contact
rates with the target species.
BURROWERS
DECOMPOSITION & NUTRIENT CYCLING

Insects and their arthropod relatives are responsible for much of the
nutrient cycling,conditioning and aeration of the soil. The value of
nutrient cycling in terrestrial ecosystems is estimated to be over $3
trillion per year (Costanza et al. 1997). Insects and related arthropods
make up half the animal biomass in some tropical forests (Fittkau and
Linge 1973).
Examples of reducers and recyclers: dung beetles, flies, carrion
beetles, millipedes, cockroaches, ants termites.
Lindner [97] was the first to suggest the use of house fly larvae to
recycle organic wastes, specifically human waste, to produce protein
and fat as a useful byproduct. Although insect herbivory is common in
terrestrial ecosystems, it has only recently been considered an
important and persistent control on ecosystem processes and has not
been included as a factor in most ecosystem models. Herbivore
alteration of litter inputs may change litter decomposition rates and
influence ecosystem nutrient cycling too [98].

IMPORTANCE TO HUMANS

OTHER BENEFICIAL USES

MEDICINE
Insects have been invaluable as research models todiscover key
scientific principles.
Examples:
genetics: fruit flies, Madagascar hissing cockroaches
bioprospecting sources of new pharmaceuticals
first anesthesia; flies during WWII used to clean wounds
insects pollinate the rosy periwinkle plant which was an important
drug to treat childhood leukemia
sources of bioluminescent chemicals used in medical diagnostics
Potential Actions of Propolis

(The Use of Propolis in Medicine)

3.1. In Medicine

General medicinal uses of propolis include treatment of the

cardiovascular and blood systems (anemia), respiratory apparatus (for
various infections), dental care, dermatology (tissue regeneration,
ulcers, eczema, wound healing particularly burn wounds, mycosis,
mucous membrane infections and lesions), cancer treatment, immune
system support and improvement, digestive tracts (ulcers and
infections), liver protection and support and many others [4].
7. BASIC RESEARCH
How the world works; flight, miniaturization,
biomimetics, genetics,population biology.

neural

networks,

7.1. Anti-Bacterial Effect

In a study by Grange and Davey, they used a Propolis dilution of 1:20

in nutrient agar which completely inhibited the growth of S.aureaus, S.
epidermidis, Enterococcus spp. Corynebacterium spp. B. catarrahlis
and B.cereus. This dilution partially inhibited the growth of P.
aeruginosa and E.coli but had no effect on K.pneumonia. Tube dilution
studies showed that it was bactericidal for B. cereus and the gramnegative cocci at dilutions of 1:160 to 1:320, and that growth of the
H37RV reference strain of Mycobacterium tuberculosis was totally
inhibited at 1:320 and partially inhibited at 1:640 dilution[5]. Koo et al.
in Brazil found antibacterial effect of Propolis on S. mutans, S. sanguis
and A. naeslundaii in addition to the inhibition of glycosultransferase
[6].
Forensic Entomology
Forensic entomology, the use of insects and other arthropodsin
forensic investigations, has gained a lot of importanceduring the past
few decades [88]. Some of the recentliteratures like, The Manual of
Forensic Entomology bySmith [89], The entomological review by
Catts and Goff
edible herbs[60][90] and the installation of the International homepage
ofForensic Entomology [91] may be cited as examples. Mende[92]
listed a number of animals which feed on corpses. Thislist includes
flies, beetles, and other insects. By analyzing thestages of succession
of insects a first rough estimation ofthe postmortem intervals can be
made [89]. Depending onthe biogeographical region and ecological
habitat, differentspecies of necrophagous insects are involved in the
decayof a corpse. For example, examinations on insect-successionfrom
Canada are not applicable in the conditions of Germany[88]. However,
the primary purpose of forensic entomologyin todays context is the
use of insects in determining elapsedtime since death [93]. Forensic

entomology, therefore, holdsa vital position in the arena of forensic

science. The uniquerole played by insects in this field is overwhelming,
whoseplace is nonreplaceable by other organisms.

C. COMMERCIAL PRODUCTS
Silk production, the silk worm (Bombyx mori) is the sole sources of
commercial silk.
Cotton (Gossypium hirsutum) can self pollinate but can cross pollinate
with services of bees: bumble bee (Bombus spp.) Melissodes bees, and
honey bees (Apis mellifera (MfGregor 1976)
Red cochineal dye for textiles and cosmetics is produced by the scale
insect Datylopius coccus
(Dactylopiidae) native to Mexico and Kermes vermilio (Kermidae)
native to Europe.
Honey is produced by bees and harvested
Beeswax: candles, cosmetics, soap
Shellac: the scale insect Laccifer lacca (Lacciferidae), native to India,
is used to makephonograph records and varnish
Salt production: Brine shrimp cleaned salt is better tasting than salt
containing the organism
that the brine shrimp eat. The price of the salt is affected.
4. ECOLOGICAL MONITORING
Insects are excellent indicators of ecosystem health. The numbers of
species and kinds ofspecies of insects can tell scientists if a particular
ecosystem is healthy or disturbed.
5. HUMAN AND LIVESTOCK HEALTH
Arthropods as vectors of disease insects weaken or kill 200 million
people per year. Examples:
Mosquito borne diseases: malaria, yellow fever, dengue, West Nile
virus, some forms of encephalitis
Tick borne diseases: lyme disease, erlichiosis,tularemia, Q fever,
Colorado tick fever, Rocky
Mountain spotted fever, relapsing fever and more
Flea borne diseases: plague, typhus
6. AESTHETICS
A summer nightsound of crickets and the flash of fireflies.
A spring meadow withthe fluttering beauty of butterflies.
A fly-fisherman withdiversity of insects to fed the trout.
Writers, artists, poets do withthe inspiration and imagery of insects.

5. DISEASE AGENTS OR VETORS

Diseases provide natural population control, helping to control
overpopulation oroverexploitation of natural resources.

CHAPTER FOUR
Case Study
The effect of dung beetles in temperate region of New South Wales, Australia.
By Gollan et al., (2011)

Study sites
Our study was conducted in the Upper Hunter Catchment, NSW,
Australia (320_0__S 1510_0__). The lower lying areas of the valley
floor and riparian corridors have been cleared of native vegetation and
developed for timber harvesting, agriculture, mining, and urban land
use (Peake 2003). Large amounts of money and an innumerable
amount of volunteer time and effort have been spent on native tree
planting, with the goal of protecting, conserving and enhancing native
biodiversity (Anonymous 2003). Study sites (total n = 12) within the
catchment included three replicate locations of: (1) unplanted
grassland (previously cleared of all tall native vegetation and
dominated by low growing exotic forbs and pasture grasses); (2) young
revegetation (replanted with tall growing native species between one
and three years prior to this study); (3) old revegetation (replanted
with tall growing native species between seven and ten years prior to
this study), and (4) mature woodland (containing mature tall native
species [greater than 20 years of age] that had been planted at the
revegetated sites). Unplanted grassland served as an endpoint of
disturbance and a starting point of restoration, mature woodland
served as an endpoint of restoration, while replanted sites were
considered intermediate. Mature woodland was not undisturbed
remnant woodland. There is no remaining natural riparian woodland in
the Upper Hunter Valley as all has been cleared and grazed at some
time and suffers from weed invasion and other effects associated with
fragmentation and water regulation of the Hunter River. Nonetheless,
we considered that mature woodland represented an appropriate
target for restoration because it was dominated by large mature to

over-mature native plant species, there were native herbs and grasses
in the understorey while weeds were only patchy, and the ground was
mostly covered in large woody debris and leaf litter. Locations were
chosen so that they were interspersed to avoid a spatially confounded
design, and site management histories were similar. This approach
known as space-for-time substitution or a chronosequence approach
is an alternative to long-term studies, since a long-term trend is
inferred from locations of differing ages (Pickett 1989). Further detail of
all sites can be found in Gollan et al.(2011a, 2011b).
Dung beetle trapping
Dung beetles were sampled using 10 pitfalls (9 cm in diameter,13 cm
deep) at each of 12 sites. All traps were: one-third filled withethylene
glycol as preservative; placed flush to the ground surface;opened
within 24 h of each other; covered by an upturned pot-plantbase held
above the opening with pegs (to protect the preservativefrom
rainwater and debris), and baited with a small amountof pig manure.
The manure was placed in a nylon mesh (aperture2 mm) bag that
was suspended above the opening of eachtrap using a piece of
galvanized wire. Traps were collected after48 h. The trapping method
we used is a conventional techniqueconsidered effective for trapping
dung beetles (Larsen & Forsyth2005).All dung beetles were counted,
identified to species and lodgedat the Australian Museum, Sydney.
These data and species identifications are published elsewhere (Gollan
et al. 2011b).
Rates of dung removal
Rates of dung removal were measured at the same time as pitfalltraps
were operating. To do this, 10 pellets of pig manure wereplaced
randomly approximately 10 m from each trap in each site.Each pellet
was made by rolling and compressing an equal amount of manure into
a cylindrical shape (5 cm long 2 cm diameter) without internal voids
and with a smooth outer surface devoidof cavities and depressions.
Pellets were placed on bare soil andafter 48 h, dung removal was
recorded by inspecting each pelletand estimating the amount (as a
total percentage) that was missing.Shallow, half-sphere depressions
were apparent where dungbeetles had removed pieces (Fig. 1),
otherwise entire pellets wereremoved. The dung used in this trial was
collected from a commercialpiggery 24 h before any survey work
commenced.

Fig. 1. A dung pellet after 48 h showing shallow, scalloped depressions

(circled)
where dung beetles had removed pieces. Dung removal percent was
recordedby estimating the amount that was missing. When deployed,
pellets were 2 cmdiameter5 cm long and smoothed on the outer
surface to remove cavities anddepressions.
Analysis
For dung beetle abundance, species richness and percent
dungremoved, the average of the 10 samples at each site was
calculated.Least-squares regression was used to test the relationship
betweendung beetle abundance and species richness and dung
removal.
The average rates of dung removal across the four habitat
types(unplanted grassland young revegetation, old revegetation
andmature woodland) were assessed visually to determine if
removalrates after revegetation were trending toward those in
maturewoodland.A fully nested ANOVA was used to test for significant
differences(p < 0.05) among replicate locations and among habitat
types. Rawdata were transformed (arcsine) prior to analyses. Factors in
thedesign were treated as follows: (1) habitat type was fixed with
fourlevels
(unplanted
grassland,
young
revegetation,
old
revegetation,and mature woodland); and, (2) location (nested within
habitattype) was random with three levels.
Cost per sample
To provide an indication of the cost per sample, the time taken(in min)
to deploy the 10 dung pellets and the visual estimatesof decay at each

site was recorded. For the costs associated withmeasuring dung beetle
abundance and richness, the time taken tocomplete installation of the
10 pitfall traps, trap collection, and laboratorysorting and identification
was recorded. Total times were divided by 120 (12 sites 10 samples
per site) to indicate an averagetime measure, or transaction cost, per
sample.

Fig. 2. Relationship between the mean percent dung removal, and dung
beetleabundance and species richness. Lines show the linear least
squares regression foreach relationship.
Results
The regression between average percent dung removal anddung
beetle abundance was strongly positive and significant(p < 0.001, r2 =
0.84, n = 12). The relationship between average percentdung removal
and dung beetle richness was also positive andsignificant (p < 0.001,
r2 = 0.69, n = 12; Fig. 2).Across all sites and on a per sample basis,
dung removalranged from 0% (where there were no obvious signs of

removal) to 100% (pellet completely removed). Variance in dung

removalamong sites of the same habitat type was significant (F8,108 =
14.47,p < 0.0001), but there were no significant differences among
habitattypes (F3,8 = 0.39, p = 0.76). Average percent dung removal
rates(s.e.) were highest in unplanted grassland and mature
woodland(303.3% and 33 6.4%, respectively). Dung removal
ratewas least (183.2%) in young revegetation, but slightly
higher(222.9%) in old revegetation (Fig. 3). The trajectory of mean
dungremoval across the restoration gradient showed an increase
fromsites recently revegetated toward those with mature woody
vegetation

Measuring dung removal took an average of 4 min per sample.The

measurement of total dung beetle abundance required justover three
times that of dung removal (average 13 min per sample),while the
transaction cost for dung beetle richness was an average
of 17 min per sample.

Discussion
Even though dung removal was variable among locations of thesame
habitat type, the trajectory in mean dung removal across
therestoration gradient showed an increase from sites recently
revegetatedtoward
those
with
mature
woody
vegetation,
indicatingrecovery of this ecosystem function. Given the strong
correlationsbetween dung removal and dung beetle abundance and
speciesrichness, both numbers of dung beetles and numbers of
differentspecies may be recovering with native revegetation.While
there are many good reasons to include terrestrial arthropodsin
biological monitoring programs, there are also manydisadvantages.
Most disadvantages are due to the requirement ofspecialist knowledge
as well as the time consuming nature of collecting and identifying
terrestrial arthropods, which translates tohigh costs. For our method of
measuring the rate of dung removal,the cost per sample was reduced
by three fold or more. If transferredto a monitoring or survey program,
a greater number of replicates inboth space and time could be
afforded, leading to a greater chanceof detecting a trend or pattern
(i.e. reducing the chance of statisticalerrors). Importantly, our method
of measuring rates of dung removal isnon-lethal. Collecting dung
beetles using baited pitfall traps is notonly lethal to invertebrates, but
also to vertebrates that become trapped. Non-lethal methods align well
with a conservation philosophy. We estimate that thousands of nontarget invertebrateswere killed in our study. Regrettably, 23
vertebrates (15 lizards andeight frogs) were also killed in our pitfall
traps. While none of thespecies killed were rare in the region or
threatened or endangeredunder Australian legislation at the time of
this study, the fact thatany organism was maimed or killed is at odds
with the philosophyof restoration. The fact that the destruction of
organisms inbycatch is inconsistent with the aims of ecological
restoration hasbeen noted by others (e.g. Bowie & Frampton 2004).
While there have been numerous studies investigating the pros and
cons of surveyinginvertebrates using pitfalls traps (e.g. Digweed et al.
1995;Greenslade 1964; Luff 1975; Spence & Niemel 1994; Topping
&Sunderland 1992), none have raised this as an ethical dilemma.
Onecould imagine if methods such as pitfall trapping were employedin
an operational monitoring program (i.e. a program where samplesare
repeated in space and time in order to report on status andtrends in
populations) there is a risk of depleting themperhaps to local
extinction. While techniques are available to limit bycatchsuch as dry
pitfall traps, installing mesh or netting over the opening,or sieving
dung in order to isolate dung beetles, these poseother problems such
as organisms eating each other or excludingorganisms due to size of
mesh. None of these escape the taxonomicimpediment and none
necessarily provide an indication of the levelof the dung removal
occurring in the system.In an effort to minimise stress on animals as

well as surveyingunseen fauna, vertebrate surveys often utilise signs

of activityor presence such as tracks or scats (e.g. Birks et al. 2005;
Glen &Dickman 2003). Arthropod signs have rarely been exploited in
thisway. It is largely impractical, if not impossible, to identify scats
orfootprints of arthropods. But as we and others have shown,
arthropodsdo produce signatures such as nests, entrance burrows
andcatching webs that can be quantified and are useful in survey
work(e.g. Barros 2001; Gollan et al. 2010; Lobry de Bruyn 1993a;
Schlick-Steiner et al. 2006). Signs of invertebrates can be more
accuratein identifying resident species than conventional trapping
methods.Lobry de Bruyn (1993a) showed this by mapping ant
nests,entrances of spiders, scorpions and termites and compared
thesedata with species collected concurrently in pitfall traps.Different
baits where signs of removal are easily quantifiedcould also be utilised,
and these approaches could be useful forother invertebrate taxa. For
example, Lomov et al. (2009) used seedremoval rates to quantify seed
dispersal by ants. However, severalissues need to be addressed with
the choice of material. We usedpig manure because it was readily
available in sufficient quantitiesand was of a texture and consistency
that allowed molding ofa standard-sized pellet with a smooth outer
surface. We are confidentthat dung beetles were responsible for the
removal of the dungbecause the scalloped depressions in the dung
pellets are characteristicof dung beetle activity (Fig. 1). Where entire
pieces wereremoved, we observed that the soil where the dung was
positionedhad been turned over. This is typical of dung beetle species
thatbury dung where it lies.
Some dung beetle species also feed onfruit and fungi, and these could
potentially be used in a similarway as the dung in our study. Dung of
other animals is also potentiallyamenable to the technique trialed here.
However, the abilityto source large quantities, fashioning of
standardised pellets, andensuring that the taxa of interest are
responsible for removal allneed consideration. Both fruit and fungi
were abandoned in ourtrials as a range of invertebrates (e.g. carabid
beetles and ants) andvertebrates (e.g. wombats, kangaroos, and
wallabies) was likely toremove such materials.The type of dung could
confound results if rates of dungremoval were used in an operational
monitoring program. Preferencefor different dung by dung beetles has
been demonstrated (e.g.Whipple & Hoback 2012) and so it would be
important to use thesame type of dung when comparing among sites
or time. It is alsopossible that the same species of livestock fed on
different dietsand exposed to pesticides and herbicides could influence
results.Davis et al. (2004) highlighted the utility of dung beetles as
indicatorsof toxicity because species responses or assemblage
structurediffer between dung containing residues and that lacking

residuesof livestock pesticides. Thus, careful planning is needed to

reduce
the chance of biasing samples.Another issue to address prior to fullscale survey work is thelength of time samples should be left out,
which may be related tothe size of pellets. Here we used 48 h, but this
time interval may notbe suitable for all areas, especially where
abundance is much higher.Baits that have been completely removed
would be of little valueif this method was used as a surrogate of
abundance and richness.
The relationships between rates of dung removal and dung beetle
abundance and richness presented here may not to be
directlytransferable to all ecosystems or regions. It is possible that
areaswith only one or two species of dung beetle could remove
dungthat is comparable to those rates found here. Trials to determine
asuitable length of time, the size of pellets (through trialing a rangeof
different sized pellets) and the strength of relationships amongindices
are necessary in other situations.
A justification for using invertebrates in monitoring programs isthat
their large population size allows repeated collection with littlepotential
for depleting populations. This may only be true for locallycommon
species for which the method is species-specific, avoidingthose that
are rare or have small population sizes. It is well knownthat large
numbers of invertebrate species collected insurveys aresingletons
(one only collected per survey) or unique (Lim et al.2011). While
sampling methods could be responsible for suchspecies frequency
distributions, it is also likely that there are rarespecies with life
histories that are susceptible to significant populationimpacts by
repeated removal. Taxa that are characterized by short-range
endemism, that is, ones that have restricted distributionalranges, poor
dispersal powers, discontinuous habitatsand low fecundity (Harvey
2002), may be particularly susceptibleto impacts by repeated
collection. A non-lethal technique like theone trialed in this study
avoids depleting locally rare populations.
Our method of measuring the ecosystem service provided bydung
beetles also provides a way in which non-specialists andcommunity
groups can study invertebrates. Once the relationshipbetween dung
removal and dung beetle abundance and richnesshas been
established, the need for specialist help is eliminated.
So far, community monitoring initiatives involving invertebrateshave
largely been restricted to a few invertebrate groups such as aquatic
invertebrates, easily identified butterflies, the large andconspicuous
bumblebee (Bombus terrestris) or charismatic flagshipspecies such as
dragonflies (for examples, see Firehock & West1995; Kadoya &
Washitani 2007; Kadoya et al. 2009; Kuehn et al.2008). We are not

suggesting that proxies like the one we trialedwill (or should) replace
the need for specialists and theirskills, but we do encourage
invertebrate researchers to developmethods and techniques that allow
non-specialist involvement in
invertebrate surveys and conservation. Community involvement in
meaningful invertebrate conservation and habitat restoration should
contribute to overcoming some of the perception problems of
invertebrates, given that a large portion of the populationonly
recognise the dirty cockroach and the nuisance fly (Samways2007).
Despite the strengths of our method, there are also drawbacks.One of
these is the reliance on visual assessment, and thus observeraccuracy
and repeatability could be a potential issue if utilised ina full-scale
monitoring program. An effective operational monitoringprogram
requires reliable information, thus it is important tocontrol for intra and
inter-observer variability (Vos et al. 2000). Anoption to control for this
type of error would be to weigh dungpellets before and after survey.
Weighing would also account fordung removal from the interior of the
pellet that is not observableby superficial examination. Indeed, larvae
of some dung beetlespecies dig their way through dung, ingesting it as
they go. Weighing,however, would require more time and field
equipment (e.g.drying ovens and weighing balance), potentially
negating the gainsin transaction costs of the dung removal method.
Providing trainingcould be useful for ensuring observers collect robust
data. Carefultraining and retraining of observers has been shown to
ameliorateobserver-related biases in a variety of field-collected data
(e.g.Thompson & Mapstone 1997).
A second weakness is that measuring rates of dung removal doesnot
provide indication of the dung beetle species involved in dungremoval,
nor dung beetle species present (whether they removedung or not).
Knowing the dung beetle species and their traits suchas country of
origin, may be important in some situations. Turnoverin species
composition, species inventories or deriving an exoticto-native ratio
are examples that would require knowledge of thespecies responsible.
Indeed, knowledge of the country of origin wasimportant in an analysis
of the dung beetles across the restorationgradient in the larger study
that this trial was part of (seeGollan et al. 2011b). Gollan et al. (2011b)
also showed that dungbeetle species composition differed between
unplanted grasslandand mature woodland. The finding that dung
removal was similarin unplanted habitat and mature woodland (Fig. 3)
illustrates thediscrepancies that can occur.
A final point worth reiterating is that species metrics are not
necessarilydirectly related to ecological function. A temperate
foreststudy in South America, for example, found that effects of

fragmentationdid not translate into changes in dung decomposition

butthat fragmentation did affect dung beetle richness and
abundance(Bustamante et al. 2004). Paradoxically, the justification for
usinginvertebrates in ecological research and monitoring is that
theyindicate ecological function, but the relationship between
biodiversityor perhaps more correctly the metrics used to measure it
(e.g.abundance and species richness) and ecosystem function
remainspoorly known and is not always clear (Ormerod 2003).
Contrastingresults have been shown for soil fauna where diversity and
abundancehave correlated positively with decomposition rates in
soils(Loreau 2001; Setala & McClean 2004; Tinnov & Scheu 2005),but
poor relationships have also been obtained (Cox et al. 2001;Griffiths et
al. 2000; Janzen et al. 1995). Lobry de Bruyn (1993b)showed that ants
indicated a similar functional role through soilmodification in areas with
natural vegetation compared to farmlandcompletely cleared of natural
vegetation.
However, there wasa significantly smaller number of ant species found
at sites clearedof vegetation. Such disparities have led some to
conclude thatquantifying ecosystem structure and function offers more
insightsinto ecosystem status than reliance on structural data alone.
Theyshould therefore be complementary (e.g. Lobry de Bruyn
1993a;Lomov et al. 2009) where the cost of monitoring both permits.

CHAPTER FIVE
Conclusion and Recommendation
The ecological importance of terrestrial arthropods
cannot be
overemphasised.
Efforts should therefore be taken to conserve them from the adverse
effects of human activities.
Pest control measures should be target-specific to prevent the
destruction and loss of beneficial arthropods.

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