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Current knowledge of spiders in South African
agroecosystems (Arachnida, Araneae)
A. S. Dippenaar-Schoeman
a b
, A. M. Van den Berg
a
, C. R. Haddad
c
& R. Lyle
a
a
ARC-Plant Prot ect ion Research Inst it ut e, Privat e Bag X134, Queenswood, Pret oria,
0121, Sout h Af rica
b
Depart ment of Zoology and Ent omology, Universit y of Pret oria, Pret oria, 001, Sout h
Af rica
c
Depart ment of Zoology and Ent omology, Universit y of t he Free St at e, PO Box 339,
Bloemf ont ein, 9300, Sout h Af rica
Version of record f irst published: 16 Jan 2013.
To cite this article: A. S. Dippenaar-Schoeman , A. M. Van den Berg , C. R. Haddad & R. Lyle (2013): Current knowledge
of spiders in Sout h Af rican agroecosyst ems (Arachnida, Araneae), Transact ions of t he Royal Societ y of Sout h Af rica,
DOI: 10. 1080/ 0035919X. 2012. 755136
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Transactions of the Royal Society of South Africa
http://dx.doi.org/10.1080/0035919X.2012.755136
REVIEW
Current knowledge of spiders in South African
agroecosystems (Arachnida, Araneae)
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A.S. Dippenaar-Schoeman1,2, A.M. Van den Berg1, C.R. Haddad3 & R. Lyle1
1
ARC-Plant Protection Research Institute, Private Bag X134, Queenswood, Pretoria, 0121 South Africa; 2Department of Zoology and
Entomology, University of Pretoria, Pretoria 001, South Africa; 3Department of Zoology and Entomology, University of the Free State,
PO Box 339, Bloemfontein, 9300 South Africa
*Author for correspondence. e-mail: DippenaarA@arc.agric.za
Spiders are one of the most abundant predator groups found in agroecosystems and they have special
adaptations towards a predatory way of life. The aim of this paper is to review our present knowledge of
spider diversity in different agroecosystems of South Africa, as well as their potential prey. This paper
provides a measure of what has been achieved in research on spiders in South African agroecosystems,
and identifies directions for future research. A checklist of spiders found in these systems is provided,
based on published surveys and data from the South African National Survey of Arachnida (SANSA)
database, with information on the guilds that they occupy. Thus far, 51 families with 238 genera and 413
species have been recorded from crops in South Africa. Five agrobiont species have been listed that might
play an important role as natural control agents of pests: Ostearius melanopygius (O.P.- Cambridge, 1879)
(Linyphiidae); Pardosa crassipalpis Purcell, 1903 (Lycosidae); Cheiracanthium furculatum Karsch, 1879
(Miturgidae); Heliophanus pistaciae Wesołowska, 2003 (Salticidae) and Misumenops rubrodecoratus Millot,
1941 (Thomisidae).
Keywords: Agrobiont species, agroecosystems, guilds, SANSA, Salticidae, Thomisidae
INTRODUCTION
Signatories of the Convention of Biological Diversity (CBD)
are obligated to develop a strategic plan for the conservation
and sustainable use of their biodiversity. In 1997, the South
African National Survey of Arachnida (SANSA) was launched
in accordance with the country’s obligations to the CBD
(Dippenaar-Schoeman et al., 2010). Through SANSA, essential
information has been gathered to address issues concerning
the conservation and sustainable use of the arachnid fauna in
South Africa. SANSA is an umbrella project that was implemented at a national level together with researchers and
institutions countrywide, with a common goal to document
and unify information on South African arachnids.
The role of spiders as possible natural control agents of insects
and mites needs to be evaluated, specifically for use in pest
control strategies in agroecosystems (Nyffeler & Benz, 1987;
Greenstone, 1999). Their inclusion in biodiversity inventories is
clearly desirable, as they represent approximately 2.8% of
global animal biodiversity with c. 43 500 described species
(Zhang, 2011; Platnick, 2012), all of which are functionally
significant as predators of other invertebrates. As part of
SANSA, baseline information on spiders in agroecosystems
sampled since 1972 was collated to address their possible
sustainable use. While considerable effort has been put into
gathering baseline data through surveys in agroecosystems in
South Africa, there is still considerable scope for further
experimental work on the biological control potential of the
dominant agrobiont spiders in each agroecosystem. While
predacious mites and insects have received most of the
ISSN 0035-919X Print / 2154-0098 Online
# 2013 Royal Society of South Africa
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attention in South African biological control programmes, the
role of spiders appears to have been largely ignored, despite
their importance as generalist predators (Smith-Meyer, 1996).
Reviews on the role of spiders in agroecosystems (Luczak,
1979; Riechert & Lockley, 1984; Nyffeler & Benz, 1987;
Nyffeler et al., 1994a,b; Green, 1996; Riechert & Lawrence,
1997; Sunderland, 1999; Symondson et al., 2002; Maloney et al.,
2003; Royauté & Buddle, 2012) indicate an increasing interest
in, and recognition of, spiders as natural control agents of
insects and mites in field crops and orchards. Spiders are one
of the most ubiquitous predator groups in South African
agroecosystems (Van den Berg & Dippenaar-Schoeman, 1991),
and inventories have provided valuable baseline information
on the abundant species occurring on various commercial
crops, including orchards. These studies have also provided
an indication of species that can be considered to be
agrobionts, i.e. species that reach high levels of abundance
in agroecosystems (Samu & Szinetár, 2002), and which are
likely to play an important role in the control of pest species.
Spiders have special adaptations towards a predatory way
of life. Their distensible abdomens enable them to consume a
large amount of prey within a relatively short period of time,
while their rate of predation may increase greatly for short
periods when food is locally abundant (Turnbull, 1965). They
have an exceedingly high resistance to starvation, which
enables them to survive and maintain normal reproduction
during periods of low prey availability (Anderson, 1974). This
is accomplished by an ability to decrease their metabolic rates
and also foraging activity.
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Transactions of the Royal Society of South Africa
Spiders are present throughout the year (Dippenaar-Schoeman, 1979), and during their lifespan, which varies from nine
months to 25 years, all instars feed actively as predators. Food
consumption decreases only before and during ecdysis,
although a general decrease in food consumption during the
adult male stage is quite common (Haynes & Sisojevic, 1966).
Spiders are usually polyphagous and feed on a variety of
available prey. Predation is not limited to adult insects only
but includes the egg and larval or nymphal stages as well
(Nyffeler et al., 1990). Spiders are some of the first predators to
colonize newly planted crops, and research has shown the
importance of early season assemblages in limiting pest
numbers while their densities are low (Van den Berg &
Dippenaar-Schoeman, 1991; Maloney et al., 2003).
Different species of spiders may be found in different
microhabitats. A large group of wandering spiders live on
the ground, either under stones or in burrows or cracks in the
soil, and they feed mainly on ground-living insects and larvae
hibernating in the soil. The plant-dwelling spiders are present
on the stems, foliage, bark or flowers of plants and many of
them are nocturnal hunters, actively moving around in search
of prey. A number of web-building spiders spin their webs
between the leaves and branches, under the bark and even
over the leaf surfaces. They will prey on any flying or crawling
insects that come into contact with the silk threads of their
webs.
As predators, spiders have a two-fold effect. Not only do
they feed directly on their prey, but their presence also causes
indirect mortality. For example, the presence of foraging
spiders can disturb insect larvae, which then drop from the
plant and die, or are exposed to predation by grounddwelling predators (Mansour et al., 1981). Further, the reduced
time that phytophagous pests spend on plants following
disturbance may reduce the damage that they cause to the
crop plants (Riechert, 1999). The webs spun over the leaves
also seem to make them less suitable for oviposition and
feeding by pests (Van den Berg et al., 1992). Spiders have the
added benefit of effecting wasteful or superfluous killing,
where more prey are killed than are actually fed on,
particularly at high prey densities. Thus, more prey are killed
than are actually required by the spider, resulting in a greater
degree of pest control (Sunderland, 1999).
The aim of this paper is to review our present knowledge of
spider biodiversity in different agroecosystems in South
Africa, and the prey that spiders consume in crop systems.
Any field of endeavour requires retrospection after a period of
activity, which provides a measure of what has been achieved
and identifies directions for future research. A checklist of
spiders found in South African agroecosystems is provided,
with information on the guilds that they occupy, and
agrobiont species that might play a role as natural control
agents are discussed.
MATERIAL AND METHODS
Surveys
In South Africa, the first arachnid sampling in an agroecosystem was undertaken in 1972 to investigate the role that
spiders play in strawberry fields in the biological control of
spider mites (Dippenaar-Schoeman, 1976, 1979). This was
followed by extensive surveys of spiders in conventionally
cultivated cotton fields (Dippenaar-Schoeman et al., 1999; Van
den Berg & Dippenaar-Schoeman, 1991; Van den Berg et al.,
1990), as well as in fields of genetically modified Bt-cotton
Vol. 00(00): 118, 2013
(Mellet, 2005; Mellet et al., 2006). A number of surveys in
orchards in the Mpumalanga Lowveld resulted in papers on
spiders on citrus (Dippenaar-Schoeman, 1998; Van den Berg
et al., 1992), macadamia (Dippenaar-Schoeman et al., 2001a, b)
and avocado (Dippenaar-Schoeman et al., 2005), while further
surveys were conducted in pistachio orchards in the arid
Northern Cape (Haddad, 2003). Records of spiders sampled in
a variety of crops as part of unpublished studies are also
included in the species list (Appendix 1). A considerable
proportion of records have only been identified to genus level,
but only those species identified to species level are included
in the list. Genus level records are only included if no species in
that genus have been identified to species level (Appendix 1).
Database
Data on spider species in agroecosystems in South Africa
were obtained from existing datasets for this region compiled
for the First Spider Atlas of South Africa (Dippenaar-Schoeman
et al., 2010). Records were obtained from the primary data of
specimens housed in the National Collection of Arachnida
(NCA) at the ARC-Plant Protection Research Institute (ARCPPRI), Pretoria (50 000 records), as well as a digital photographic database containing images of species recorded on
crops by the public.
Guilds
A guild is a group of species that potentially compete for
jointly exploited limited resources (Polis & McCormick, 1986).
For the present study, two main guilds were recognized,
namely wandering spiders (W) and web-builders (WB), with
further subdivisions based on their preferred microhabitat or
type of web constructed (Foord et al., 2011) (Appendix 1).
Web-building spiders construct webs to capture their prey.
The majority of web-builders spin webs on the plants, either
on, between or around leaves and flowers, between branches,
or between trees, and usually catch flying insects. Several
families (e.g. Linyphiidae and Agelenidae) construct webs
close to the soil surface or in low-growing vegetation, such as
orchard ground covers.
Wandering spiders actively hunt their arthropod prey.
Many wandering spiders are nocturnal and construct tubular
retreats of silk in which they spend periods of inactivity
during the day. Some families (e.g. Salticidae, Thomisidae and
some Lycosidae) are diurnally active and rest at night, either
in silk retreats or hanging from draglines from vegetation.
Resting retreats are spun in many different microhabitats,
including beneath young leaves, in curled or dead leaves,
within inflorescences, under bark, in soft sand, or in grasses
and weeds in the ground cover vegetation.
Agrobiont species
Agrobiont species are defined as species that reach a high
degree of dominance in agroecosystems, that may usually
only be found in high numbers in natural habitats that are
frequently disturbed (Samu & Szinetár, 2002). Furthermore,
they defined agrobionts as species with an average dominance of greater than 1% in arable fields that occurred in at
least 75% of the fields sampled within a particular geographical region, in that case Hungarian cereal fields. Particular
agrobiont species are usually characteristic of agroecosystems
of fairly similar structures, e.g. cereals and alfalfa (Samu et al.,
2011). The agrobionts associated with arable lands, orchards
and rice paddies, for example, are inherently different due to
differences in the structure of the vegetation and the
A.S. Dippenaar-Schoeman et al.: Knowledge of Spider Diversity in Agroecosystems
contrasting management strategies applied in each (Samu &
Szinetár, 2002). We regarded agrobiont species as those
representing more than 1% of all the individuals in a sampled
assemblage from a particular crop. It is also important to
consider how widespread the occurrence of a species in the
considered crop(s) is. Species that are below this limit, but still
common in fields, are called ‘‘agrophile’’ species after Luczak
(1979).
RESULTS AND DISCUSSION
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Crops
Avocado
South Africa is the world’s fourth-largest producer of
avocado (Persea americana) (United States Department of
Agriculture, 2005). Locally, avocado trees and fruit are
attacked by at least 30 species of insects and mites (Van den
Berg et al., 1999a). Spiders inhabiting avocado orchards in
Israel were studied by Mansour et al. (1985), who assessed
their role as natural enemies of the giant looper, Boarmia
selenaria (Lepidoptera: Geometridae). In South Africa, surveys
were carried out in two avocado orchards in the Mpumalanga
Lowveld over a one-year period using canopy fogging as the
collecting method (Dippenaar-Schoeman et al., 2005). A total
of 3715 specimens representing 26 families, 68 genera and 90
species were collected. The Salticidae comprised 31.0% of the
total spiders collected, followed by Thomisidae (23.9%) and
Tetragnathidae (11.8%). The most species-rich families were
the Araneidae (20 spp.), Salticidae (14 spp.) and Thomisidae
(12 spp.). The thomisid Oxytate argenteooculata was the most
abundant species and represented 22.2% of all spiders
collected, followed by two salticids, Thyene coccineovittata
(11.5%) and T. natali (11.0%), and a tetragnathid, Tetragnatha
subsquamata (8.4%). Of the spiders collected, 77% were
wanderers and 23% were web-builders.
Citrus
Large numbers of spiders are present in citrus orchards
(Citrus spp.), as has been reported in South Africa (CatIing,
1970; Van den Berg et al., 1987), as well as in other countries
(Shulov, 1938; Carroll, 1980; Mansour et al., 1982; Mansour &
Whitcomb, 1986). Many of the pests and potential pests that
occur on citrus in South Africa are under effective biological
control (Bedford, 1978). However, the natural enemies of a
few others, like the citrus psylla, Trioza erytreae (Hemiptera:
Triozidae), are unable to keep these pests below the economic
threshold. Several spider species have been observed to prey
on citrus psylla (Van der Merwe, 1923; Catling & Annecke,
1968; Catling, 1970), and Van den Berg et al. (1987) expressed
the opinion that spiders are possibly the most important
predators of this pest.
During a two year survey in Mpumalanga Province, South
Africa, a total of 3054 spiders representing 21 families were
sampled using beating and pittraps in an unsprayed citrus
orchard (Van den Berg et al., 1992). Salticidae was the most
abundant family (34.4%) followed by the Theridiidae (21.9%),
Thomisidae (11.9%), Araneidae (7.9%) and Clubionidae
(7.0%). Eighteen species of spiders were observed to prey on
citrus psylla, while six species trapped nymphs and adults in
their retreats or webs. The spider species most commonly
found on the soil was Pardosa crassipalpis (Lycosidae), while
the plant-dwellers are represented by Cheiracanthium
furculatum (Miturgidae), Enoplognatha molesta (Theridiidae),
3
Eperigone fradeorum (Linyphiidae), and Misumenops rubrodecorata (Thomisidae). The wandering spiders constituted
61.5% of the spider fauna collected and the web-builders
38.5%. Data indicate that while spiders are unable to keep
citrus psylla populations at acceptably low levels, they may
contribute to reducing their numbers (Van den Berg et al.,
1992).
Cotton
Cotton (Gossypium spp.) is an important agricultural product
in South Africa, with commercial as well as developing
farmers involved in its cultivation. The wide-scale introduction of Bt cotton has contributed significantly to improving
yields and profitability of cotton production (Limpopo Province Freight Transport Data Bank, 2012). Several pests that
attack cotton have a wide range of natural enemies, of which
spiders are one prominent group (Van den Berg & DippenaarSchoeman, 1991).
Spiders are common and occur in high numbers in cotton
fields, preying on a variety of cotton pests. Plant-dwelling and
ground-dwelling spiders were collected from 1979 to 1997 in
five cotton-growing areas in South Africa using pittraps and
the whole-plant bag technique (Dippenaar-Schoeman et al.,
1999). Thirty-one families, represented by 92 genera and 127
species, were recorded during these surveys. Thomisidae
were the most species-rich family with 21 species, followed
by Araneidae (18 spp.) and Theridiidae (11 spp.). The most
abundant spider species were Pardosa crassipalpis, Enoplognatha
molesta, Mermessus fradeorum (Linyphiidae) and Misumenops
rubrodecorata. Wandering spiders constituted 61.5% and webbuilders 38.5% of all spiders collected. In cotton fields under
different management strategies in Mpumalanga (Mellet et al.,
2006), a survey of the ground-dwelling spiders showed a
dominance of Lycosidae (62.5%), Theridiidae (20.0%) and
Linyphiidae (9.1%). Neither Bt-cotton nor chemically treated
cotton showed long-term significant impacts on grounddwelling spiders. Spiders are also abundant on cotton in the
USA and have been widely studied, with more than 300
species recorded (Whitcomb & Bell, 1964; Leigh & Hunter,
1969; Young & Lockley, 1985; Nyffeler et al., 1987a, b, 1989,
1992a, b).
Macadamia
South Africa is a large producer of macadamia (Macadamia
integrifolia) nuts (Groenewald, 1999). About 60 insect and two
mite species are known to attack macadamia trees and their
fruit (Van den Berg et al., 2000). In South Africa, pentatomid
and coreid stinkbugs are the most important pests in
macadamia orchards (Van den Berg et al., 1999b). Infestation
results in young nuts dropping and older nuts developing
lesions.
Arboreal spiders were collected over a 12-month period by
canopy fogging in three macadamia orchards in the Mpumalanga Lowveld of South Africa. The spiders were sampled
every 23 weeks, with 10 trees sampled in each orchard per
sample date using dichlorvos as a knock-down spray (Dippenaar-Schoeman et al., 2001a, b). A total of 2778 specimens
representing 21 families, 57 genera and 80 species were
recorded. The Salticidae was very dominant, representing
72.7% of the total spiders collected, followed by the Sparassidae (6.9%), Hersiliidae (3.9%) and Araneidae (3.3%). The
most diverse families were the Salticidae (17 spp.), Araneidae
(16 spp.) and Thomisidae (11 spp.). Wandering spiders
dominated the fauna, representing 95.8% of the total
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specimens collected, compared to 4.2% that were webbuilders. The salticid Thyene coccineovittata was the most
abundant species and represented 29.7% of all the spiders
collected, followed by T. natalli with 14.2%, Viciria alba with
8.6% and Tusitala guineensis with 8.3%. These four species
were present in all three orchards throughout the year.
Maize
Maize (Zea mays) is the most important grain crop in South
Africa, being both the major fodder grain and staple food of
the majority of the South African population. After more than
a decade of production, the cultivation of GM maize in South
Africa has increased dramatically. During the 2010/2011
production season, GM maize contributed to 80% or 1.9
million ha of the total commercial area planted to maize
(Department of Agriculture, Forestry and Fisheries (DAFF),
2011b). Maize is attacked by several pest species, of which the
most serious is the maize stalk borer, Busseola fusca (Lepidoptera: Noctuidae), which can cause enormous crop losses.
Other very important maize pests in South Africa are the
noctuid cutworms (Euxoa and Agrotis species) (Du Plessis,
2003).
Although spiders have been studied on maize in South
Africa, as well as overseas, very little is known of the South
African fauna. Midega et al. (2008) sampled ground-dwelling
spiders in South African and Kenyan maize using a combination of pitfall traps and soil samples. They collected a total of
284 spiders in South Africa, with the Lycosidae the most
abundant family.
A second, unpublished survey was undertaken on the farm
Buiteplaas in the Delmas district in Mpumalanga Province,
South Africa during the 20042005 and 20052006 summer
growing seasons. Three treatments were evaluated, namely Bt
maize, conventionally sprayed maize, and unsprayed maize.
Pitfall traps were used to sample the spiders (M. van
Jaarsveld, personal communication, 2006). Fourteen families
were collected, representing 32 genera and 38 species. The
Linyphiidae, represented by four species, occurred in the
highest numbers, with the Lycosidae, represented only by
Pardosa crassipalpis, second.
Pine plantations
In many areas of Africa, the planting of exotic trees has
superseded the area covered by indigenous forests. This has
an effect on the vegetation structure at ground level, which
should have an effect on the community structure of the
ground-living spider fauna. Since spiders are important
components of forest floor ecosystems as invertebrate predators (Moulder & Reichle, 1972), this could have significant
secondary effects on other invertebrate groups and plants as
well. Plantations exceed a million hectares in South Africa
alone. As only a fraction of South Africa’s area (0.2%) is
covered by tall evergreen indigenous forest (Huntley, 1984),
their conservation and that of their cryptofauna is of utmost
importance and should receive high priority for conservation
efforts (Dippenaar-Schoeman et al., 2006).
Two spider surveys in pine plantations have been undertaken in South Africa to date. The first was done in a pine
plantation at Sabie in the Mpumalanga Province, South Africa
(Van den Berg & Dippenaar-Schoeman, 1988). Of the 1484
spiders collected, 38.5% belonged to Miturgidae, 13% to the
Lycosidae, 10% Tetragnathidae and 8% Salticidae. A second
survey was undertaken at Ngome Forest, situated on the
escarpment of northern KwaZulu-Natal, South Africa (Van
der Merwe et al., 1996). This survey of ground-living spiders
was conducted over a one-year period using 180 pitfall traps
(36 per habitat). The survey covered five different habitat
types: grassland, open indigenous forest, dense indigenous
forest, ecotone and pine plantation. A total of 9360 spiders,
represented by 136 species, were trapped. Pine had the lowest
spider species richness while grassland had the highest
species richness. Due to the large variation in species richness
within habitat types, the results from this study did not
convincingly support the notion that pine plantations have a
lower ground-living spider diversity than indigenous habitats.
However, cluster analysis of the sampling grids showed that
different habitat types supported different spider assemblages
(Van der Merwe, 1994).
Pistachio
Pistachio (Pistacia vera) was established as a new crop in
South Africa during the 1990s as part of an investment by the
Industrial Development Corporation in the Prieska district in
the Northern Cape Province. As part of a larger biomonitoring
programme in these orchards, intended to identify local and
introduced pests and their potential biological control agents,
spiders were selected as a target group representing the
generalist predators forming part of the natural enemy
complex in the orchards. Surveys were undertaken at ground
level using pitfall traps and hand collecting, from the ground
covers by sweep-netting, and from the tree canopies using
canopy fogging, in three orchards of contrasting size and age.
The pitfalls yielded 1692 spiders, representing 16 families
and 49 species (Haddad & Dippenaar-Schoeman, 2006). Active
searching yielded 645 spiders, representing 16 families and 63
species. Four families (Linyphiidae, Gnaphosidae, Lycosidae
and Salticidae) dominated the epigeic fauna, collected by both
sampling methods, but their abundance varied considerably
between the two techniques. The sheet-web spider Ostearius
melanopygius (Linyphiidae) dominated the fauna sampled by
both techniques in all three orchards, and can be considered
an agrobiont.
In the ground covers, 1760 spiders representing 55 species
were collected in the three orchards. Two species, Peucetia
viridis (Oxyopidae) and Heliophanus pistaciae (Salticidae),
dominated the spider fauna, accounting for 29.3% and
23.4% of the total, respectively (Haddad et al., 2004a). In total,
5843 spiders were collected from the tree canopies, representing 18 families and 88 species (Haddad et al., 2005). Three
species dominated the spider fauna: H. pistaciae (53.4%),
Cheiracanthium furculatum (12.7%) and Neoscona subfusca (Araneidae, 6.4%).
Based on observations in the field, H. pistaciae and C.
furculatum prey on at least seven orders of arthropods, while
the web-building N. subfusca captured five different orders of
arthropods in their webs. All three species preyed on thrips
(Thysanoptera: Phlaeothripidae), seed bugs (Hemiptera: Lygaeidae), aphids (Homoptera: Aphididae) and leafhoppers
(Homoptera: Cicadellidae), all of which have potential to
cause damage to pistachio leaves and nuts (Haddad, 2003). In
laboratory and field experiments, H. pistaciae was found to
have a restricted role as a predator of Nysius natalensis
(Hemiptera: Lygaeidae), showing a preference for Drosophila
melanogaster (Diptera: Drosophilidae) in laboratory feeding
trials (Haddad et al., 2004b). Nysius natalensis was shown to
cause direct damage to pistachio nuts through the puncture
wounds caused by its feeding, as well as through the
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dissemination of fungal pathogens to the nuts that result in
further damage to the kernels (Swart, 2002).
Based on these results, it is clear that the older orchards had
greater species richness and abundance of spiders in all three
strata compared to the more recently established orchards,
largely due to natural succession. Furthermore, orchard
establishment appears to have a distinct negative effect on
spider species richness and abundance when compared to the
assemblages from surrounding natural habitats, which had a
much greater abundance and species richness (Haddad et al.,
2008), and may ultimately reach abundance and richness
levels comparable to natural habitats. Surprisingly, the
ground-dwelling orchard fauna in South Africa is very similar
to that found in European orchards, but that from the
surrounding natural habitat is very different to the assemblages found in natural habitats in Europe (Haddad et al.,
2008).
Strawberries
Strawberry (Fragaria ananassa) production has increased
greatly in South Africa during the last decade. The common
red spider mite, Tetranychus cinnabarinus (Acari: Tetranychidae) causes serious damage to strawberries in South Africa.
During surveys, Coates (1972) found that Dermaptera and
Araneae were the most important predators of spider mites in
strawberry beds. In continuation of his work, a bio-ecological
study of spiders in strawberry beds was undertaken from 1972
to 1974 to determine the species present and elucidate their
role as predators of mites (Dippenaar-Schoeman, 1976).
Weekly counts of spiders were made and 28 genera representing 14 families were collected by hand. Of the 5059
spiders recorded during the two seasons, 70.3% belonged to
the Lycosidae, and 28.0% to the Araneidae, Thomisidae,
Clubionidae, Salticidae and Linyphiidae, combined. Spider
densities ranged from 0.56/m2 during July 1972 to 13.52/m2 in
February 1973 (Dippenaar-Schoeman, 1979). The most abundant species collected was Pardosa crassipalpis, which accounted for over 80% of all Lycosidae. The life cycle and
some ecological aspects of this species, which is a predator of
T. cinnabarinus, were studied (Dippenaar-Schoeman, 1977).
When it comes to the control of mites by spiders, the three
seasons (wintersummer) during which the planting, growing
and ripening of the fruit takes place, are of prime importance,
because the mite population usually begins to increase during
the winter months. The spider families that develop their full
potential with regard to numbers during this period are
considered to be the more important groups in the natural
control of these mites. In this regard, the Lycosidae is of major
importance, while the Linyphiidae, Thomisidae, and to a
lesser extent Clubionidae, Salticidae and Araneidae, may also
exert some influence as predators.
Tomatoes
Tomatoes (Solanum lycopersicum) are the second most important vegetable crop grown in South Africa, after potatoes.
They contributed approximately 25% to the gross value of
production in 2011. The average household in South Africa
consumes between five and ten tomatoes per week (DAFF,
2011a). Tomatoes are attacked by several insect and mite pest
species (Myburgh, 1988).
Several surveys have been undertaken in the tomatoproducing areas of Limpopo, North West and Gauteng
Provinces in search of biological control agents of tomato
pests (Krüger & Dippenaar-Schoeman, 2000). In addition,
5
tomato plants in greenhouses at the ARC-PPRI research
station were screened regularly for insect and mite pests
and their natural enemies. Insects, mites and spiders were
collected using random plant inspection, as many pests tend
to aggregate. Specimens were hand collected or captured
through beating of plants and sweep nets. During these
surveys, 356 spiders were sampled in total, representing 16
families, 50 genera and 62 species. Araneidae (34%) was the
most species-rich family and also collected in the largest
numbers. Unfortunately, most of the specimens were immature and could not be identified to species level. However,
several of the species collected are known predators of spider
mites.
Vineyards
Spiders are found in and around vineyards and prey on
insect pests that are found associated with grapes (Vitis spp.).
Although there is no information available regarding the
spiders in vineyards in South Africa, Costello and Daane
(1995, 1997, 2005) and Roltsch et al. (1998) found that spiders
were by far the dominant predator group collected in grape
vineyards in central California, and in some years spiders
comprised over 95% of predators sampled. Eight species of
spiders represented more than 90% of the fauna from central
California, with two theridiids (Theridion dilutum and T.
melanurum) the most abundant, followed by the miturgid
Cheiracanthium inclusum (Miturgidae).
Three unpublished surveys have been undertaken in grape
vineyards in Southern Africa. The first was undertaken in 2008
on the border of Namibia and South Africa, where a total of
106 spiders representing eight species from six families were
collected. Two species, Cheiracanthium furculatum and an
unidentified Theridion sp. (Theridiidae), together constituted
86% of all the spiders collected.
The second survey was conducted in the Cape Floristic
Region of South Africa, where wine grape production and
biodiversity conservation are of major importance, and
innovative management of the landscape is necessary. A
study was done in June and October 2006 using pitfall traps to
look at the impacts of alternative farming methods, such as
organic and biodynamic farming, on biodiversity. A total of 17
spider families represented by 42 species were sampled
during this study (Gaigher, 2008).
In 2011, vineyards in the Western Cape Province were
surveyed and 940 spiders were sampled with paper traps or
by hand to investigate their possible role in the transfer of
vinestem virus (F. Halleen, personal communication, 2012). A
total of 16 families and 52 species were sampled and the most
abundant species was Cheiracanthium furculatum, followed
by Euryopis episinoides (Theridiidae) and Pelecopsis janus
(Linyphiidae).
Although spiders may be of benefit as predators in
vineyards, they may also pose problems as possible invasive
species when they land in containers of table grapes and are
inadvertently exported. When the grapes are harvested, some
spiders escape detection during the packing process by hiding
in silk retreats made in bunches of grapes. The grapes are
chilled prior to export, causing the spiders to become dormant
and immobile. Spiders are able to lower their metabolic rates,
enabling them to survive long periods of exposure to low
temperatures. When the exported containers are opened at
the retailers or by consumers in recipient countries, the
spiders try to escape after this period of inactivity. It is mainly
members of the family Miturgidae (Cheiracanthium spp.) that
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have been implicated in such scenarios (Dippenaar-Schoeman, 2007).
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Spider families and agrobiont species
Araneidae
A total of 36 araneid species have been sampled from
agroecosystems in South Africa (Appendix 1), but none of
them were sufficiently abundant in any crop to be regarded as
an agrobiont. On citrus, four species of araneids were
observed to prey on the citrus psylla (Van den Berg et al.,
1992), amongst which was an Araneus sp. Of the 18 species of
araneids collected on cotton, most were rare except for
Neoscona triangula, N. blondeli, N. subfusca and Nemoscolus
obscurus, which were regularly found. Araneids were also
the most abundant spiders collected from tomatoes.
Araneids spin their webs between plants and feed on a
variety of flying and jumping insects. Araneids were observed
preying on Helicoverpa armigera (Lepidoptera: Noctuidae)
moths and larvae, both in the laboratory and in the field
(Dippenaar-Schoeman et al., 1999). Members of the Araneidae
were also commonly found on cotton in other countries (Dean
et al., 1982; Nyffeler et al., 1989), while Lincoln et al. (1967)
found that species of Araneus were one of the most important
predators of bollworm moths. Other pest species preyed on
by araneid species (Kagan, 1943) include Adelphocoris rapidus
nymphs and Pseudatomoscelis seriatus adults (Hemiptera:
Miridae), Brevicoryne brassicae, Myzus cerasi, M. Iythri and
Rhopalosiphum padi aphids (Hemiptera: Aphididae), H. armigera
larvae and moths, and spur-throated locust nymphs.
Linyphiidae
Nineteen species have been sampled from crops in South
Africa (Appendix 1) and two species could be recognized as
agrobionts, namely the cosmopolitan species Ostearius
melanopygius, which was abundant on cotton, maize and
pistachio, and Eperigone fradeorum, which was abundant in
cotton fields. On cotton, the Linyphiidae were mostly
observed in small webs spun across the leaf lamina (Van
den Berg, 1989). Eperigone fradeorum was the most common
species, followed by Microlinyphia sterilis and O. melanopygius.
Erigone irrita and M. sterilis were found in association with the
red spider mite Tetranychus lombardinii on several vegetables
(Smith-Meyer, 1996). In South African maize, the most
abundant linyphiids were O. melanopygius, Meioneta habra
and Limoneta sirimoni (Dippenaar-Schoeman, unpublished
report, 1989). Ostearius melanopygius dominated the grounddwelling spider fauna sampled in pistachio orchards of
contrasting size and age (Haddad & Dippenaar-Schoeman,
2006). In vineyards it was Pelecopsis janus that was the most
abundant, followed by M. habra and L. sirimoni.
Although no studies have been conducted in South African
winter wheat, linyphiids often form the dominant group of
generalist predators in European wheat and barley (Schmidt
& Tscharntke, 2005; Öberg et al., 2008), where they are
particularly important in the control of aphids (Sunderland
et al., 1986a, b; Harwood et al., 2001, 2003). Some Iinyphiids
have been observed to attack second-instar bollworm
(Helicoverpa zea) larvae as they crawl up the main stems of
cotton plants (Whitcomb, 1967). In Texas, Nyffeler et al. (1988)
found that aphids were the dominant prey of the linyphiid
Frontinella pyramitela.
Lycosidae
Twenty-three species of lycosids have been found in crops
(Appendix 1) but only one species, Pardosa crassipalpis, was
sampled in abundance from strawberry and cotton fields and
could be considered an agrobiont in these crops. This species
was also collected from several other agroecosystems but in
lower numbers.
Lycosids are cursorial hunters and were observed on the
leaves and flowers of plants, or running on the ground and
hiding under dry leaves in cotton fields (Van den Berg, 1989).
The numerically dominant species, P. crassipalpis, was recorded from five cotton growing areas, and preyed on red
spider mites and various stages of cotton bollworm larvae
(Helicoverpa armigera) in the laboratory (Dippenaar- Schoeman
et al., 1999). They are the dominant family at the ground level
in Bt cotton fields, representing 62.5% of the spiders collected
(Mellet et al., 2006). Lycosidae was the dominant spider family
in strawberries, where P. crassipalpis was the dominant species
present, representing 82% of all the lycosids sampled and 63%
of all the spiders present (Dippenaar-Schoeman, 1979). The
biology of this species was described by Dippenaar-Schoeman
(1977). Lycosidae was also the most abundant family sampled
from sugar cane in South Africa (Leslie & Boreham, 1981).
During this study, the authors conducted cross-over electrophoresis on the stomach contents of arthropods sampled on
sugar cane, and determined that ants and spiders were the
most common predators that fed on the sugarcane borer,
Eldana saccharina (Lepidoptera: Pyralidae).
Coates (1974), Dippenaar-Schoeman (1976) and Botha (1986)
found P. crassipalpis to be active predators of red spider mites
in various crops in South Africa. In the USA, Whitcomb et al.
(1963) found that Iycosids swarm over cotton plants at night
in summer and that Pardosa spp. prey on pink bollworm
moths. They presumably destroy first and second-instar
larvae of the bollworm on the plant, as well as those that
fall to the ground. Bollworm moths are also vulnerable to
predation during their brief exposure on the soil surface
following emergence from their pupae (Lincoln et al., 1967;
Whitcomb, 1967). Laboratory studies on feeding strategies of
Pardosa hortensis showed that they can play a positive role in
controlling agricultural pests in a density-sensitive way (Samu
& Bı́ró, 1993). Lycosids also prey on Helicoverpa pupae (Lincoln
et al., 1967), cabbage looper moths (Whitcomb & Bell, 1964),
Heliothis virescens (Lepidoptera: Noctuidae) and Lygus Iineolaris
(Hemiptera: Lygaeidae) (Hayes & Lockley, 1990).
Miturgidae
Seven species of sac spiders were commonly found on crops
(Appendix 1). Cheiracanthium furculatum is the dominant sac
spider recorded from crops in South Africa and is an agrobiont
species in vineyards and pistachio nuts. It is also the dominant
sac spider recorded from citrus in South Africa, and Van den
Berg et al. (1992) observed that they prey mainly on the citrus
mite Panonychus ulmi (Acari: Tetranychidae), killing 29.3 mites/
spider/day. Their silk retreats, which are usually constructed
between rolled up leaves for resting, frequently trap citrus
psylla (Van den Berg et al., 1992). Two species, C. furculatum
and C. africanum, were occasionally collected during cotton
surveys. They are aggressive spiders and kill any prey that
they encounter, although not always feeding on them.
According to Brettell & Burgess (1973), C. furculatum was
frequently found in the narrow space between the bract and
boll of cotton in Zimbabwe. In the laboratory they preyed on
all larval stages of the cotton bollworm.
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A.S. Dippenaar-Schoeman et al.: Knowledge of Spider Diversity in Agroecosystems
The genus Cheiracanthium is one of the most common genera
found on cotton in the USA (Whitcomb & Bell, 1964; Young &
Lockley, 1985), Australia (Bishop & Blood, 1981), Israel
(Mansour, 1987), Zimbabwe (Brettell & Burgess, 1973) and
South Africa (Dippenaar-Schoeman et al., 1999). Carroll (1980)
found that they prey on citrus thrips, mites, insect eggs and
lepidopterous larvae, and that they are probably the most
promising natural enemies of citrus pest arthropods in
California. Cheiracanthium species have been reported to
prey on second and third instars of Helicoverpa spp. larvae,
cotton looper larvae, green vegetable bug and cotton seed bug
in cotton (Bishop & Blood, 1981). They also feed preferentially
on abundant pests during pest outbreaks, and show a direct
numerical relationship with changes in Helicoverpa spp. larval
abundance (Bishop, 1979). McDaniel & Sterling (1982) found
that an individual consumed on average 14.2 Heliothis virescens
eggs in the laboratory during a 24-hour period. Feeding
studies in Israel showed that C. mildei are important predators
of red spider mites (T. cinnabarinus). During feeding studies in
the laboratory, females of C. mildei on average preyed on 27.5
mites per day and C. mildei juveniles on 18.8 mites per day. In
contrast, the predacious mite Phytoseiulus persimilis (Acari:
Phytoseiidae), which is extensively used in biological control,
has been reported to prey on only 11.3 mites per day
(Mansour et al., 1995).
Oxyopidae
Nine oxyopid species have been sampled from crops in
South Africa (Appendix 1). Oxyopids are diurnal and nocturnal cursorial hunters that are found on cotton plants,
frequently near the flowering parts (Van den Berg, 1989).
Oxyopes bothai was occasionally found in all the cotton areas
sampled. Peucetia viridis dominated the ground cover fauna in
two of three pistachio orchards in the Northern Cape, and can
be considered an agrobiont, while O. bothai and O. hoggi were
low in abundance in the three orchards sampled (Haddad
et al., 2004a). All three species occurred in the pistachio
canopies but were low in abundance (Haddad et al., 2005).
Oxyopids were consistently among the most abundant
arthropod predators in cotton agroecosystems in the USA
(Young & Lockley, 1985; Nyffeler et al., 1987a,b; 1992a).
According to Nyffeler et al. (1987a), 0.12 million prey may be
killed per week per hectare by O. salticus spiders in unsprayed
cotton. In sprayed fields, spider density is often strongly
reduced, reducing their impact on pests. Whitcomb (1967)
found that Oxyopes spp. destroyed more second-instar larvae
of Helicoverpa zea than did any other arthropod predator in
cotton. They also preyed on mirids, cotton leafhoppers,
tarnished plant bugs (Whitcomb et al., 1963), Adelphocoris
rapidus (adults), Pseudatomoscelis seriatus adults, and aphids
(Kagan, 1943; Nyffeler et al., 1992b). In Australia, Bishop &
Blood (1981), found that O. mundulus fed preferentially on
Helicoverpa larvae during outbreaks and showed a numerical
response to changes in Helicoverpa larval abundance. In
feeding experiments in Israel, oxyopids consumed, on average, 16.8 red spider mites per day in the laboratory (Mansour
et al., 1995).
Salticidae
A total of 70 salticid species have been sampled from crops
(Appendix 1). Although salticids are a common part of the
predator complex in most agroecosystems, their species
composition and numbers vary between crops and between
regions. For example, salticids are sometimes the dominant
7
family present on citrus (Van den Berg et al., 1987, 1992), but
are less abundant on cotton (Dippenaar-Schoeman et al., 1999;
Mellet et al., 2006) and strawberries (Dippenaar-Schoeman,
1976).
In orchard crops particulartly, Salticidae has been shown to
be an important component of spider assemblages, where
several species are agrobionts. In a knock-down study of three
macadamia orchards in the Mpumalanga Lowveld in South
Africa, 73% of the sampled spiders were salticids, and four
species together represented more than 61% of all spiders
collected. Thyene coccineovittata was the most abundant species
and represented 30% of all the spiders collected, followed by
T. natali with 14%, Viciria alba with 9% and Tusitala guineensis
with 8%. These four species were present throughout the year
in all three orchards sampled (Dippenaar-Schoeman et al.,
2001a, b). On avocado in Mpumalanga, salticids comprised
31.0% of the spiders collected, and two of the four most
abundant species were salticids, viz. Thyene coccineovittata
(11.5%) and T. natali (11.0%). In citrus orchards in Mpumalanga, salticids accounted for 34.0% of all spiders collected
(Van den Berg et al., 1987, 1992). In pistachio orchards in the
Northern Cape, the fauna of ground covers in pistachio
orchards was dominated by Salticidae (31.8%), with
Heliophanus pistaciae (23.4%) and Phlegra karoo (5.8%) the most
abundant species (Haddad et al., 2004a). In pistachio canopies,
salticids accounted for 59.2% of the spiders collected, with H.
pistaciae by far the most abundant species, accounting for 53.4%
of the spiders collected (Haddad et al., 2005). On the ground
surface, salticids accounted for 20.0% of the spiders collected in
pitfall traps and 15.4% of the spiders collected by hand, with P.
karoo the most abundant salticid collected by both methods
(Haddad & Dippenaar-Schoeman, 2006). As H. pistaciae is
found in multiple habitat strata in high abundance, and occurs
throughout the year in the orchards, it can be considered a
pistachio agrobiont (Haddad & Louw, 2006).
Regarding field crops, various salticid genera were found in
cotton. They were common in all five areas sampled and were
found both in pit-traps and on plants, where they constructed
sacs in rolled-up leaves (Van den Berg, 1989). Most salticids
are diurnal and their prey is limited to species present on the
cotton plants. In the laboratory, a single salticid devoured 17
red spider mites within a 12-minute period and also preyed
on the first two larval stages of the cotton bollworm
Helicoverpa armigera (Dippenaar-Shoeman et al., 1999). Van
den Berg et al. (1987, 1992) observed several salticid species of
the genera Myrmarachne, Hyllus and Thyene preying on adult
citrus psylla in citrus orchards (Dippenaar-Schoeman, 2001).
In pistachio orchards, Heliophanus pistaciae was found to have a
limited role as a predator of the lygaeid bug Nysius natalensis
(Haddad et al., 2004b), although it does feed on a variety of
different arthropods and may play a role in their biological
control (Haddad, 2003).
Studies elsewhere in the world indicate that salticids prey on
a variety of insect and mite species, and that predation is not
limited to adult prey but includes the eggs and larval or
nymphal stages as well (Whitcomb, 1974; Nyffeler et al., 1994a).
Salticidae are highly polyphagous but can narrow their prey
spectrum when prey becomes available in high numbers
(Nyffeler et al., 1994a). Studies indicate that they prey on
several insect and mite species in citrus. Edwards (1981) and
Mansour et al. (1982) reported salticids preying on citrus
weevils, and Carroll (1980) recorded thrips, mites and midges
as prey of salticids. Feeding experiments conducted in Israel
showed salticids to consume on average 10.1 red spider mites
8
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per day (Mansour et al., 1995). In US cotton fields, salticids prey
on bollworms (first to third instar larvae), boll weevils, robber
flies (Whitcomb & Bell, 1964), Helicoverpa armigera larvae,
Alabama argillacea (Lepidoptera: Noctuidae) larvae and adults,
and nymphs of Adelphocoris rapidus (Kagan, 1943). They seize
second-instar larvae of H. zea from cotton foliage and under
bracts of squares and bolls of the plant (Whitcomb, 1967;
Young, 1989) and are able to destroy large numbers of first and
second-instar larvae in the field (Lincoln et al., 1967).
Theridiidae
Gumfoot-web spiders build webs on, between and around
leaves. Twenty-one species have been sampled from crops in
South Africa (Appendix 1). Theridion purcelli and Enoplognatha
molesta were the most abundant theridiids found in citrus
orchards in South Africa (Van den Berg et al., 1992). This
corresponds well with the findings of Mansour & Whitcomb,
(1986), who found that 34% of spiders in the undercover of
citrus groves in Israel belong to the genus Theridion. In cotton
fields, E. molesta was the most common species, followed by T.
purcelli (Dippenaar-Schoeman et al., 1999). In the laboratory
both species fed on red spider mites, the first three larval
stages and adult stages of Helicoverpa armigera, leafhoppers
and aphids. Van den Berg et al. (1992) observed that
Achaearanea sp., Enoplognatha sp. and two Theridion spp.
frequently spun their webs close to the leaf surface in citrus
trees. Citrus psylla nymphs and adults crawl into the webs,
where they are caught and eaten. In the laboratory at ARCPPRI, it was also observed that they feed on red spider mites
(Dippenaar-Schoeman et al., 1999).
Theridiids are important predators of several insect pest
species (Mansour & Whitcomb, 1986). In Israel, Mansour et al.
(1995) found that a Steatoda sp. was able to prey on 9.5
Tetranychus cinnabarinus mites per day in the laboratory. In
cotton, they have been found to prey on boll weevils and
bollworm larvae (Whitcomb & Bell, 1964). Prey records of
Theridion australe and Tidarren haemorrhoidale, often found on
cotton in Texas, showed aphids to be their main prey
(Nyffeler et al., 1988). In Australia, Achaearanea veruculata was
observed to prey on jassids, cotton aphids, looper larvae and
cotton seed bugs (Bishop & Blood, 1981).
Thomisidae
Fourty-four thomisids were sampled from crops
(Appendix 1). A total of 18 species were collected from citrus
in South Africa, with Misumenops rubrodecoratus the most
common. Van den Berg et al. (1992) observed that M.
rubrodecoratus prey on adult citrus psylla, and feeding experiments indicated that they also prey on aphids, red spider
mites and thrips (Dippenaar-Schoeman, unpublished data,
1998). On cotton, M. rubrodecoratus are mostly found on leaves
and stems, near bracts of cotton plants, as well as hiding
under dry leaves on the ground (Van den Berg, 1989); M.
rubrodecoratus was the most abundant species collected in five
cotton growing areas. In the laboratory they preyed on red
spider mites, the first two larval stages of Helicoverpa armigera,
as well as on aphids (Dippenaar-Schoeman et al., 1999). In two
cotton fields in Arizona, USA, Misumenops celer occurred in
high numbers and constituted a major portion (44.257.6%) of
the predator complex (Plagens, 1983). Oxytate argenteooculata
was the most abundant species on avocado trees and
represented 22.2% of all spiders collected (Dippenaar-Schoeman et al., 2005). They are less abundant in other crops.
Thomisids are active during the day (Leigh & Hunter, 1969)
and prey on a variety of small invertebrates, potentially
playing an important role in the natural control of pests
such as aphids, red spider mites and thrips (Bogya, 1999).
Several studies have highlighted the importance of thomisids
in biological control in cotton fields. Lincoln et al. (1967) found
Misumenops spp. on all parts of the plant, especially on or
under bracts of cotton bolls and squares, although very few
were taken from the ground. They capture second-instar
larvae of H. zea in cotton terminals, where they wait in
ambush (Whitcomb, 1967). They also prey on several other
insects, including Geocoris punctipes (Hemiptera: Geocoridae),
Lygus lineolaris (Hemioptera: Lygaeidae), Acalymna vittata
(Coleoptera: Chrysomelidae) (Lincoln et al., 1967), syrphid
flies and beetles (Whitcomb & Bell, 1964), Pseudatomoscelis
seriatus adults (Dean et al., 1987), and larvae of Alabama
argillacea and H. armigera (Kagan, 1943). In China, they have
been observed to feed on eggs and first instar larvae of H.
armigera (Wu et al., 1981).
CONCLUSION
Spiders are common and occur in fairly high numbers in
agroecosystems in South Africa where they form part of the
natural enemy complex as generalist predators. Their presence in crops should be encouraged and steps taken to
protect them from potentially lethal pesticides. Owing to the
different guilds they occupy, various families are affected
differently by pesticides (Pekár & Haddad, 2005). Judicious
use of pesticides in crop programmes may result in more
complex and abundant spider communities, thereby augmenting biological pest control. Although spiders may be
incapable of controlling major pest outbreaks by themselves,
their role in a complex predator community could be
important to regulate pest species at low densities early in
the season and between peaks of pest species activity. They
could therefore play an important role in keeping pests at
endemic levels and preventing outbreaks. It is important that
farmers and students be made aware of their role.
ACKNOWLEDGEMENTS
We thank the ARC-Plant Protection Research Institute for
providing the opportunity and the facilities for most of these
investigations. The technical assistance and helpfulness of the
staff at the different research stations is greatly appreciated.
We also want to thank all the people who assisted at various
times with fieldwork and for their encouragement. Vaughn
Swart (University of the Free State) is also thanked for
providing spider material from pecan nut orchards for
inclusion in this study.
REFERENCES
ANDERSON, J.F. 1974. Responses to starvation in the spiders Lycosa
lenta Hentz and Filistata hibernalis (Hentz). Ecology 55: 576585.
BEDFORD, E.C.G. 1978. Methods of controlling citrus pests. In Bedford,
E.C.G. (Ed.) Citrus PPests in the Republic of South Africa. Science Bulletin
391, Department of Agricultural Technical Services, Republic of South
Africa. pp. 1116.
BISHOP, A.L. 1979. The role of spiders as predators in a cotton
ecosystem. Working papers, Australian Applied Entomology Research
Conference, Queensland, 7678.
BISHOP, A.L. & BLOOD, P.R.B. 1981. Interactions between natural
populations of spiders and pests in cotton and their importance to
cotton production in Southeastern Queensland. General Applied
Entomology 13: 98104.
Downloaded by [ARC Central Office], [Professor A.S. Dippenaar-Schoeman] at 04:25 16 January 2013
A.S. Dippenaar-Schoeman et al.: Knowledge of Spider Diversity in Agroecosystems
BOGYA, S. 1999. Spiders (Araneae) as polyphagous natural enemies in
orchards. Unpublished PhD thesis, Landbouwuniversiteit, Wageningen. 189 pp.
BOTHA, J.H. 1986. An evaluation of the influence of some pesticides on
natural enemies of spider mite populations in cotton. Unpublished
PhD thesis, Rand Afrikaans University. 276 pp.
BRETTELL, J.H. & BURGESS, M.W. 1973. A preliminary assessment of the
effects of some insecticides on predators of cotton pests. Rhodesia
Agricultural Journal 70: 103104.
CARROLL, D.P. 1980. Biological notes on the spiders of some citrus
groves in central and Southern California. Entomological News 91: 147
154.
CATLING, H.D. 1970. The bionomics of the South African citrus psylla
Trioza erytreae (Del Guercio) (Homoptera: Psyllidae) 4. The influence
of predators. Journal of the Entomological Society of Southern Africa 34:
381391.
CATLING, H.D. & ANNECKE, D.P. 1968. Ecology of citrus psylla in the
Letaba district of Northern Transvaal. South African Citrus Journal 410:
817.
COATES, T.J.D. 1972. The influence of some natural enemies and
pesticides on various populations of Tetranychus cinnabarinus (Boisduval), T. lombardinii Baker and Pritchard and T. luderti Zacher (Acari:
Tetranychidae), with aspects of their biologies. Unpublished DSc
thesis, Rand Afrikaans University. 120 pp.
COATES, T.J.D. 1974. The influence of some natural enemies and
pesticides on various populations of Tetranyhus cinnabarinus (Boisduval),
T. lombardinii Baker & Pritchard and T. ludeni Zacher (Acari: Tetranychidae) with aspects of their biologies. Entomology Memoir, Department of Agricultural Technical Services, Republic of South Africa, vol. 42,
40 pp.
COSTELLO, M.J. & DAANE, K.M. 1995. Spider species composition and
seasonal abundance in San Joaquin Valley grape vineyards.
Environmental Entomology 24: 823831.
COSTELLO, M.J. & DAANE, K.M. 1997. Comparison of sampling
methods used to estimate spider (Araneae) species abundance and
composition in vine grape vineyards. Ecological Entomology 23: 3340.
COSTELLO, M.J. & DAANE, K.M. 2005. Day vs. night sampling for
spiders in grape vineyards. Journal of Arachnology 33: 2532.
DEPARTMENT OF AGRICULTURE, FORESTRY AND FISHERIES (DAFF) 2011a.
A profile of the South African tomato market value chain. DAFF, Directorate
Marketing, Arcadia.
DEPARTMENT OF AGRICULTURE, FORESTRY AND FISHERIES (DAFF) 2011b.
Trends in the Agricultural Sector 2011. DAFF, Pretoria.
DEAN, D.A., STERLING, W.L. & HORNER, N.V. 1982. Spiders in eastern
Texas cotton fields. Journal of Arachnology 10: 251260.
DEAN, D.A., STERLING, W.L., NYFFELER, M. & BREENE, R.G. 1987.
Foraging by selected spider predators on the cotton fleahopper and
other prey. Southwestern Entomologist 12: 263270.
DIPPENAAR-SCHOEMAN, A.S. 1976. An ecological study of the spider
population in strawberries with special reference to the role of Pardosa
crassipalpis Purcell (Araneae: Lycosidae) in the control of Tetranychus
cinnabarinus (Boisduval). Unpublished MSc thesis, Rand Afrikaans
University. 119 pp.
DIPPENAAR-SCHOEMAN, A.S. 1977. The biology of Pardosa crassipalpis
Purcell (Araneae: Lycosidae). Journal of the Entomological Society of
Southern Africa 40: 225236.
DIPPENAAR-SCHOEMAN, A.S. 1979. Spider communities in strawberry
beds: seasonal changes in numbers and species composition.
Phytophylactica 11: 14.
DIPPENAAR-SCHOEMAN, A.S. 1998. Spiders as predators of citrus pests.
In Bedford, E.C.G. & Van den Berg, M.A. (Eds). Citrus Pests in Southern
Africa. Nelspruit, Agricultural Research Council, pp. 3435.
DIPPENAAR-SCHOEMAN, A.S. 2001. Spiders as predators of pests of
tropical and non-citrus subtropical crops. In Van den Berg, M.A. & De
Villiers, E.A. (Eds). Pests of Tropical and Non-citrus Subtropical Crops in
the Republic of South Africa. Nelspruit, ARC-Institute for Tropical and
Subtropical Crops, pp. 1517.
DIPPENAAR-SCHOEMAN, A.S. 2007. Spiders in vineyards. SANSA
Newsletter 3: 12.
9
DIPPENAAR-SCHOEMAN, A.S., VAN DEN BERG, A.M. & VAN DEN BERG,
A. 1999. Spiders in South African cotton fields: species diversity
and abundance (Arachnida: Araneae). African Plant Protection 5:
93103.
DIPPENAAR-SCHOEMAN, A.S., VAN DEN BERG, M.A. & VAN DEN BERG,
A.M. 2001a. Spiders in macadamia orchards in the Mpumalanga
Lowveld of South Africa: species diversity and abundance (Arachnida: Araneae). African Plant Protection 7: 3646.
DIPPENAAR-SCHOEMAN, A.S., VAN DEN BERG, M.A. & VAN DEN BERG,
A.M. 2001b. Salticid spiders in macadamia orchards in the Mpumalanga Lowveld of South Africa (Arachnida: Araneae: Salticidae).
African Plant Protection 7: 4751.
DIPPENAAR-SCHOEMAN, A.S., VAN DEN BERG, A.M., VAN DEN BERG,
M.A. & FOORD, S.H. 2005. Spiders in avocado orchards in the
Mpumalanga Lowveld of South Africa: species diversity and abundance (Arachnida: Araneae). African Plant Protection 11: 816.
DIPPENAAR-SCHOEMAN, A.S., VAN DER MERWE, M. & VAN DEN BERG,
A.M. 2006. Habitat preferences and seasonal activity of the Microstigmatidae from the Ngome State Forest, South Africa (Arachnida:
Araneae). Koedoe 49: 8589.
DIPPENAAR-SCHOEMAN, A.S., HADDAD, C.R., FOORD, S.H., LYLE, R.,
LOTZ, L., HELBERG, L., MATHEBULA, S., VAN DEN BERG, A., VAN DEN
BERG, A.M., VAN NIEKERK, E. & JOCQUÉ, R. 2010. First Atlas of the
Spiders of South Africa. South African National Survey of Arachnida.
SANSA Technical Report version 1.
DU PLESSIS, J. 2003. Maize Production. Department of Agriculture,
Directorate Agricultural Information Services, Pretoria, 35 pp.
EDWARDS, G.B. 1981. Sound production by courting males of Phidippus
mystaceus (Araneae: Salticidae). Psyche 88: 199214.
FOORD, S., DIPPENAAR-SCHOEMAN, A.S. & HADDAD, C.R. 2011. The
faunistic diversity of spiders (Arachnida, Araneae) of the Savanna
Biome in South Africa. Transactions of the Royal Society of South Africa 66:
170201.
GAIGHER, R. 2008. The effect of different vineyard management
systems on the epigaeic arthropod assemblages in the Cape Floristic
Region, South Africa. Unpublished MSc thesis, University of Stellenbosch. 118 pp.
GREEN, J. 1996. Spiders in biological control an Australian perspective. Revue Suisse de Zoologie, hors série: 245253.
GREENSTONE, M.H. 1999. Spider predation: who and why we study it.
Journal of Arachnology 27: 333342.
GROENEWALD, A. 1999. Makadamiabedryf staan saam vir groter mark.
Landbouweekblad 1122: 14.
HADDAD, C.R. 2003. Spider ecology in pistachio orchards in South
Africa. Unpublished MSc thesis, University of the Free State. 173 pp.
HADDAD, C.R. & LOUW, S. vdM. 2006. Phenology, ethology and
fecundity of Heliophanus pistaciae Wesolowska (Araneae: Salticidae), an
agrobiont jumping spider in South African pistachio orchards. African
Plant Protection 12: 111.
HADDAD, C.R., LOUW, S.vdM. & DIPPENAAR-SCHOEMAN, A.S. 2004a.
Spiders (Araneae) in ground covers of pistachio orchards in South
Africa. African Plant Protection 10: 97107.
HADDAD, C.R., LOUW, S.VDM. & DIPPENAAR-SCHOEMAN, A.S. 2004b. An
assessment of the biological control potential of Heliophanus pistaciae
(Araneae: Salticidae) on Nysius natalensis (Hemiptera: Lygaeidae), a
pest of pistachio nuts. Biological Control 31: 8390.
HADDAD, C.R., DIPPENAAR-SCHOEMAN, A.S. & PEKÁR, S. 2005. Arboreal
spiders (Arachnida: Araneae) in pistachio orchards in South Africa.
African Plant Protection 11: 3241.
HADDAD, C.R. & DIPPENAAR-SCHOEMAN, A.S. 2006. Epigeic spiders
(Araneae) in pistachio orchards in South Africa. African Plant Protection
12: 1222.
HADDAD, C.R., LOUW, S.VDM. & PEKÁR, S. 2008. Commercial pistachio
orchards maintain lower density and diversity of spiders (Araneae): a
study from South Africa. African Plant Protection 14: 2436.
HARWOOD, J.D., SUNDERLAND, K.D. & SYMONDSON, W.O.C. 2001.
Living where the food is: web location by linyphiid spiders in
relation to prey availability in winter wheat. Journal of Applied Ecology
38: 8899.
Downloaded by [ARC Central Office], [Professor A.S. Dippenaar-Schoeman] at 04:25 16 January 2013
10
Transactions of the Royal Society of South Africa
HARWOOD, J.D., SUNDERLAND, K.D. & SYMONDSON, W.O.C. 2003.
Web-location by linyphiid spiders: prey-specific aggregation and
foraging strategies. Journal of Animal Ecology 72: 745756.
HAYES, J.L. & LOCKLEY, T.C. 1990. Prey and nocturnal activity of wolf
spiders (Araneae: Lycosidae) in cotton fields in the delta region of
Mississippi. Environmental Entomology 19: 15121518.
HAYNES, D.L. & SISOJEVIC, P. 1966. Predatory behaviour of Philodromus
rufus Walckenaer (Araneae: Thomisidae). Canadian Entomology 98:
113133.
HUNTLEY, B.J. 1984. Characteristics of South African biomes. In
Booysen, P. De, V. & Tainton, N.M. (Eds). Ecological Effects of Fire in
South African Ecosystems. Berlin, Springer, pp. 118.
KAGAN, M. 1943. The Araneida found on cotton in central Texas.
Annals of the Entomlogical Society of America 36: 257258.
KRÜGER, K. & DIPPENAAR-SCHOEMAN, A.S. 2000. Integrated pest management of insect and mite pests on tomatoes. Tomato Producers’ Organisation, ARC-PPRI, unpublished report.
LEIGH, T.F. & HUNTER, R.E. 1969. Predacious spiders in California
cotton. California Agriculture 23: 45.
LESLIE, G.W. & BOREHAM, P.F.L. 1981. Identification of arthropod
predators of Eldana saccharina Walker (Lepidoptera: Pyralidae) by
cross-over electrophoresis. Journal of the Entomological Society of Southern Africa 44: 381388.
LIMPOPO PROVINCE FREIGHT TRANSPORT DATA BANK
(LPFDB). 2012. Cotton production and processing. http://www.ldrt.
gov.za?wpcontent/Limpopo_Databank/industries,salt/index.html (accessed 24 July 2012).
LINCOLN, C., PHILLIPS, J.R., WHITCOMB, W.H., DOWELL, G.C., BOYER,
W.P., BELL, K.O. Jr., DEAN, G.L., MATTHEWS, E.J., GRAVES, J.B.,
NEWSOM, L.D., CLOWER, D.F., BRADLEY, J.R. Jr. & BAGENT, J.L. 1967.
The bollworm-tobacco budworm problem in Arkansas and Louisiana.
Arkansas Agricultural Experimental Station. Bulletin 720: 66.
LUCZAK, J. 1979. Spiders in agrocoenoses. Polish Ecological Studies 5:
151200.
MCDANIEL, S.G. & STERLING, W.L. 1982. Predation of Heliothis virescens
(F.) eggs on cotton in East Texas. Environmental Entomology 11: 6066.
MALONEY, D., DRUMMOND, F.A. & ALFORD, R. 2003. Spider predation
in agro- ecosystems, can spiders effectively control pest populations.
Maine Agricultural and Forest Experimental Station Technical Bulletin 190:
132.
MANSOUR, F.A. 1987. Spiders in sprayed and unsprayed cotton fields
in Israel, their interactions with cotton pests and their importance as
predators of the Egyptian cotton leaf worm, Spodoptera littoralis.
Phytoparasitica 15: 3141.
MANSOUR, F. & WHITCOMB, W.H. 1986. The spiders of a citrus grove in
Israel and their role as biocontrol agents of Ceroplastes floridensis
(Homoptera: Coccidae). Entomophaga 31: 269276.
MANSOUR, F.A., ROSEN, D. & SHULOV, A. 1981. Disturbing effect of a
spider on larval aggregations of Spodoptera littoralis. Entomologia
Experimentalis et Applicata 29: 234237.
MANSOUR, F.A., ROSS, J.W., EDWARDS, G.B., WHITCOMB, W.H. &
RICHMAN, D.B. 1982. Spiders of Florida citrus groves. Florida
Entomologist 65: 514522.
MANSOUR, F., WYSOKI, M. & WHITCOMB, W.H. 1985. Spiders inhabiting
avocado orchards and their role as natural enemies of Boarmia selenaria
Schiff. (Lepidoptera: Geometridae) larvae in Israel. Acta Oecologia 6:
315321.
MANSOUR, F., BERNSTEIN, E. & ABO-MOCH, F. 1995. The potential of
spiders of different taxa and a predacious mite to feed on the carmine
spider mite, a laboratory study. Phytoparasitica 23: 217220.
MELLET, M.A. 2005. Cotton cultivation practices with special reference
to the effect of Bt-cotton on Arthropoda populations. Unpublished
MSc thesis, University of Pretoria. 329 pp.
MELLET, M.A., SCHOEMAN, A.S. & DIPPENAAR-SCHOEMAN, A.S. 2006.
The effect of Bt-cotton cultivation on spider (Arachnida: Araneae)
populations in Marble Hall, South Africa. African Plant Protection 12:
4050.
MIDEGA, C.A.O., KHAN, Z.R., VAN DEN BERG, J., OGOL, C.K.P.O.,
DIPPENAAR-SCHOEMAN, A.S., PICKETT, J.A. & WADHAMS, L.J. 2008.
Response of ground-dwelling arthropods to a ‘push-pull’ habitat
management system: spiders as an indicator group. Journal of Applied
Entomology 132: 248254.
MOULDER, B.C. & REICHLE, D.E. 1972. Significance of spider predation
in the energy dynamics of forest-floor arthropod communities.
Ecological Monograph 42: 473498.
MYBURGH, A.C. (Ed.) 1988. Crop pests in southern Africa: Volume 3.
Potatoes and other vegetables. Bulletin 415. Plant Protection Research
Institute. Department of Agriculture and Water Supply, Pretoria.
94 pp.
NYFFELER, M. & BENZ, G. 1987. Spiders in natural pest control: a
review. Journal of Applied Entomology 103: 321339.
NYFFELER, M., DEAN, D.A. & STERLING, W.L. 1987a. Evaluation of the
importance of the striped lynx spider, Oxyopes salticus (Araneae:
Oxyopidae), as a predator in Texas cotton. Environmental Entomology
16: 11141123.
NYFFELER, M., DEAN, D.A. & STERLING, W.L. 1987b. Predation by the
green lynx spider, Peucetia viridans (Araneae: Oxyopidae), inhabiting
cotton and woolly croton plants in East Texas. Environmental
Entomology 16: 355359.
NYFFELER, M., DEAN, D.A. & STERLING, W.L. 1988. Prey records of the
web-building spiders Dictyna segregata (Dictynidae), Theridion australe
(Theridiidae), Tidarren haemorrhoidale (Theridiidae), and Frontinella
pyramitela (Linyphiidae) in a cotton agroecosystem. The Southwestern
Naturalist 33: 215218.
NYFFELER, M., DEAN, D.A. & STERLING, W.L. 1989. Prey selection and
predatory importance of orb-weaving spiders (Araneae: Araneidae,
Uloboridae) in Texas cotton. Environmental Entomology 18: 373380.
NYFFELER, M., BREENE, R.G., DEAN, D.A. & STERLING, W.L. 1990.
Spiders as predators of arthropod eggs. Journal of Applied Entomology
109: 490501.
NYFFELER, M., DEAN, D.A. & STERLING, W.L. 1992a. Diets, feeding
specialization, and predatory role of two lynx spiders, Oxyopes salticus
and Peucetia viridans (Araneae: Oxyopidae), in a Texas cotton
agroecosystem. Environmental Entomology 21: 14571465.
NYFFELER, M., STERLING, W.L. & DEAN, D.A. 1992b. Impact of the
striped lynx spider (Araneae: Oxyopidae) and other natural enemies
on the cotton fleahopper (Hemiptera: Miridae) in Texas cotton.
Environmental Entomology 23: 11781188.
NYFFELER, M., DEAN, D.A. & STERLING, W.L. 1994a. How spiders make
a living. Environmental Entomology 23: 13571367.
NYFFELER, M., STERLING, W.L. & DEAN, D.A. 1994b. Insectivorous
activities of spiders in United States field crops. Journal of Applied
Entomology 118: 113128.
ÖBERG, S., MAYR, S. & DAUBER, J. 2008. Landscape effects on
recolonisation patterns of spiders in arable fields. Agriculture, Ecosystems and Environment 123: 211218.
PEKÁR, S. & HADDAD, C.R. 2005. Can spiders (Araneae) avoid a surface
with pesticide residues? Pest Management Science 61: 11791185.
PLAGENS, M.J. 1983. Populations of Misumenops (Araneida: Thomisidae) in two Arizona cotton fields. Environmental Entomology 12: 572
575.
PLATNICK, N.I. 2012. The World Spider Catalogue, version 12. American
Museum of Natural History. Online at: http://research.amnh.org/iz/
spiders/catalog/ DOI:10.5531/db.iz.0001 (accessed October 2012).
POLIS, G.A. & MCCORMICK, S.J. 1986. Scorpions, spiders and solpugids:
predation and competition among distantly related taxa. Oecologia 71:
111116.
RIECHERT, S.E. 1999. The hows and whys of successful pest suppression by spiders: insights from case studies. Journal of Arachnology 27:
387396.
RIECHERT, S.E. & LAWRENCE, K. 1997. Test for predation effects of
single versus multiple species of generalist predators: spiders
and their insect prey. Entomologia Experimentalis et Applicata 84: 147
155.
RIECHERT, S.E. & LOCKLEY, T. 1984. Spiders as biological control
agents. Annual Review of Entomology 29: 299320.
ROLTSCH, W.R., HANNA, R., SHOREY, H., MAYSE, M. & ZALOM, F. 1998.
Spiders and vineyard habitat relationships in central California. In
Downloaded by [ARC Central Office], [Professor A.S. Dippenaar-Schoeman] at 04:25 16 January 2013
A.S. Dippenaar-Schoeman et al.: Knowledge of Spider Diversity in Agroecosystems
Pickett, C.H. & Bugg, R.L. (Eds). Enchancing Biological Control: Habitat
Management to Promote Natural Enemies of Agricultural Pests. Berkeley,
Berkeley, University of California Press, pp. 311338.
ROYAUTÉ, R. & BUDDLE, C.M. 2012. Colonization dynamics of agroecosystem spider assemblages after snow-melt in Quebec (Canada).
Journal of Arachnology 40: 4858.
SAMU, F. & Bı́RÓ, Z. 1993. Functional response, multiple feeding and
wasteful killing in a wolf spider (Araneae: Lycosidae). European Journal
of Entomology 90: 471476.
SAMU, F. & SZINETÁR, C. 2002. On the nature of agrobiont spiders.
Journal of Arachnology 30: 389402.
SAMU, F., SZINETÁR, C., SZITA, E., FETYKÓ, K. & NEIDERT, D. 2011.
Regional variations in agrobiont composition and agrobiont life
history of spiders (Araneae) within Hungary. Arachnologische
Mitteilungen 40: 105109.
SCHMIDT, M.H. & TSCHARNTKE, T. 2005. Landscape context of
sheetweb spider (Araneae: Linyphiidae) abundance in cereal fields.
Journal of Biogeography 32: 467473.
SHULOV, A. 1938. Observations on citrus spiders. Hardar 11: 206208.
SMITH-MEYER, M.K.P. 1996. Mite pests and their predators on cultivated
plants in Southern Africa. Vegetables and berries. Plant Protection
Research Institute Handbook no. 6. Pretoria, Agricultural Research
Council. 90 pp.
SUNDERLAND, K.D. 1999. Mechanisms underlying the effects of spiders
on pest populations. Journal of Arachnology 27: 308316.
SUNDERLAND, K.D., FRASER, A.M. & DIXON, A.F.G. 1986a. Distribution
of linyphiid spiders in relation to capture of prey in cereal fields.
Pedobiologia 29: 367375.
SUNDERLAND, K.D., FRASER, A.M. & DIXON, A.F.G. 1986b. Field and
laboratory studies on money spiders (Linyphiidae) as predators of
cereal aphids. Journal of Applied Ecology 23: 433447.
SWART, V.R. 2002. Insect-fungal ecology on three selected new crops
in South Africa. Unpublished MSc thesis, University of the Free State.
151 pp.
SYMONDSON, W.O.C., SUNDERLAND, K.D. & GREENSTONE, M.H. 2002.
Can generalist predators be effective biocontrol agents? Annual Review
of Entomology 47: 561594.
TURNBULL, A.L. 1965. Effects of prey abundance on the development
of the spider Agelenopsis potteri (Blackwall) (Araneae: Agelenidae).
Canadian Entomologist 97: 141147.
UNITED STATES DEPARTMENT OF AGRICULTURE (USDA). 2005. Avocados. Online: http://www.fas.usda.gov/htp/Hort_Circular/2005/03-05/
Avocados%203-7-05.pdf (accessed September 2012).
VAN DEN BERG, A.M. 1989. An investigation into the effects of two
commonly used pesticides on spider mite predator populations in
cotton with special reference to spiders. Upublished MSc thesis, Rand
Afrikaans University. 121 pp.
VAN DEN BERG, M.A., DEACON, V.E., FOURIE, C.J. & ANDERSON, S.H.
1987. Predators of the citrus psylla, Trioza erytreae (Hemiptera:
Triozidae), in the Lowveld and Rustenburg areas of Transvaal.
Phytophylactica 19: 285289.
VAN DEN BERG, A.M. & DIPPENAAR-SCHOEMAN, A.S. 1988. Spider
communities in a pine plantation at Sabie, eastern Transvaal: a
preliminary survey. Phytophylactica 20: 293296.
11
VAN DEN BERG, A.M., DIPPENAAR-SCHOEMAN, A.S. & SCHOONBEE, H.J.
1990. The effect of two pesticides on spiders in South African cotton
fields. Phytophylactica 22: 435441.
VAN DEN BERG, A.M. & DIPPENAAR-SCHOEMAN, A.S. 1991. Spiders,
predacious insects and mites on South African cotton. Phytophylactica
23: 8586.
VAN DEN BERG, M.A., DIPPENAAR-SCHOEMAN, A.S., DEACON, V.E. &
ANDERSON, S.H. 1992. Interaction between citrus psylla, Trioza erytreae
(Hem. Triozidae), and spiders in an unsprayed citrus orchard in the
Transvaal Lowveld. Entomophaga 37: 599608.
VAN DEN BERG, M.A., DE VILLIERS, E.A. & JOUBERT, P.H. 1999a.
Identification manual for avocado pests. Agricultural Research Council, Nelspruit. 53 pp.
VAN DEN BERG, M.A., STEYN, W.P. & GREENLAND, J. 1999b. Hemiptera
occurring on macadamia in the Mpumalanga Lowveld of South
Africa. African Plant Protection 5: 8992.
VAN DEN BERG M.A., DE VILLIERS E.A. & JOUBERT P.H. 2000. Macadamia
pests in South Africa. Nelspruit, ARC - Institute for Tropical and
Subtropical Crops. 68 pp.
VAN DER MERWE, C.P. 1923. The citrus psylla (Trioza merwei, Pettey).
Reprint no. 41. Department of Agriculture. Union of South Africa.
VAN DER MERWE, M. 1994. A comparative survey of cursorial spider
communities in indigenous Afromontane forests and in pine plantations. Unpublished MSc thesis, University of Pretoria. 133 pp.
VAN DER MERWE, M., DIPPENAAR-SCHOEMAN, A.S. & SCHOLTZ, C.H.
1996. Diversity of ground-living spiders at Ngome State Forest,
KwaZulu/Natal: a comparative survey in indigenous forest and pine
plantations. African Journal of Ecology 34: 342350.
WHITCOMB, W.H. 1967. Field studies on predators of the second-instar
bollworm Heliothis zea (Boddie) (Lepidoptera: Noctuidae). Journal of the
Georgia Entomological Society 2: 113118.
WHITCOMB, W.H. 1974. Natural populations of entomophagous
arthropods and their effect on the agoecosystem. In Maxwell, F.G.
& Harris, F.A. (Eds). Proceedings of the Summer Institute of Biological
Control of Plant, Insects and Diseases. Jackson, MI, University Press of
Mississippi, pp. 150164.
WHITCOMB, W.H. & BELL, K. 1964. Predaceous insects, spiders and
mites of the Arkansas cotton fields. Bulletin of the Arkansas Agricultural
Experimental Station 690: 183.
WHITCOMB, W.H., EXLINE, H. & HUNTER, R.C. 1963. Spiders of the
Arkansas cotton field. Annals of the Entomological Society of America 56:
653660.
WU, Y., LI, Y.P. & JIANG, D.Z. 1981. Integrated control of cotton pests
in Nanyang region. Acta Entomologica Sinica 24: 3441.
YOUNG, O.P. 1989. Field observations of predation by Phidippus audax
(Araneae: Salticidae) on arthropods associated with cotton. Journal of
Entomological Science 24: 266273.
YOUNG, O.P. & LOCKLEY, T.C. 1985. The striped lynx spider Oxyopes
salticus (Araneae: Oxyopidae), in agroecosystems. Entomophaga 30:
329346.
ZHANG, Z.-Q. 2011. Animal biodiversity: an introduction to higherlevel classification and taxonomic richness. Zootaxa 3148: 712.
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Transactions of the Royal Society of South Africa
APPENDIX 1. SPIDER SPECIES COLLECTED IN
AGROECOSYSTEMS IN SOUTH AFRICA.
Guild abbreviations: BGW Burrow ground dweller; FWB
Funnel-web; GW Ground wanderer; PW Plant wanderer;
RWB Retreat-web; GWB Gumfoot-web; OWB Orb-web;
SpWB Space-web; ShWB Sheet-web.
*agrobiont species.
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Species
AGELENIDAE
Agelena gaerdesi Roewer,
1995
Benoitia ocellata (Pocock,
1900)
AMAUROBIIDAE
Pseudauximus annulatus
Purcell, 1908
AMMOXENIDAE
Ammoxenus amphalodes
Dippenaar & Meyer, 1980
Ammoxenus coccineus Simon,
1893
ARANEIDAE
Aethriscus olivaceus Pocock,
1902
Arachnura scorpionoides
Vinson, 1863
Araneus apricus Karsch, 1884
Guild
Crops
FWB
pistachio
FWB
maize
RWB
citrus, pistachio, maize
GW
cotton
GW
cabbage, pistachio
OWB
avocado
OWB
macadamia
OWB
Araneus holzapfelae Lessert,
1936
Araneus nigroquadratus
Lawrence, 1937
Argiope australis (Walckenaar,
1805)
OWB
avocado, citrus,
macadamia, tomatoes
avocado, tomatoes
OWB
pine plantations, tomatoes
OWB
Argiope trifasciata (Forskål,
1775)
Caerostris sexcuspidata
(Fabricius, 1973)
Chorizopes sp.
Cyclosa insulana (Costa, 1834)
Cyclosa oculata (Walckenaer,
1802)
Cyphalonotus larvatus (Simon,
1881)
Cyrtophora citricola (Forskål,
1775)
OWB
OWB
avocado, peach, pine
plantations, pistachio,
pumpkin
cotton, kenaf, lucerne,
tomatoes
apple, citrus, pine
plantations, tomatoes
citrus
avocado, citrus, tomatoes
citrus, cotton, macadamia,
potatoes
avocado, citrus,
macadamia
citrus, lemon, pine
plantations, pistachio,
prickly pear, tomatoes
citrus
OWB
OWB
strawberry
citrus, potatoes
OWB
OWB
OWB
citrus, tomatoes
potatoes, pumpkin
cotton
OWB
avocado, pecans, tomatoes
OWB
sugar cane
OWB
citrus, cotton, pistachio
OWB
avocado, cotton, pecans,
pistachio, tomatoes,
vineyards
avocado, cotton, lucerne,
maize, pecans, pistachio,
tomatoes
Gasteracantha versicolor
(Walckenaer, 1842)
Gea infuscata Tullgren, 1910
Hypsosinga lithyphantoides
(Walckenaer, 1802)
Isoxya tabulata (Thorell, 1859)
Kilima decens (Blackwall, 1866)
Larinia natalensis (Grasshoff,
1971)
Lipocrea longissima (Simon,
1881)
Mahembea hewitti (Lessert,
1930)
Nemoscolus elongatus
Lawrence, 1947
Neoscona blondeli (Simon,
1885)
Neoscona moreli (Vinson,
1863)
OWB
OWB
OWB
OWB
OWB
OWB
OWB
APPENDIX 1 (Continued )
Species
Guild
Crops
Neoscona hirta (C.L. Koch,
1844)
Neoscona quincasea Roberts,
1983
Neoscona rapta (Thorell, 1899)
Neoscona rufipalpis (Lucas,
1858)
Neoscona subfusca (C.L. Koch,
1837)
OWB
vineyards
OWB
pine plantations, vineyards
OWB
OWB
pistachio
avocado, citrus,
macadamia
avocado, citrus, cotton,
grapefruit, macadamia,
pecans, pine plantations,
pistachio, tomatoes,
vineyards
avocado, macadamia,
onion, pecans, sorghum,
tomatoes
citrus
OWB
Neoscona triangula
(Keyserling, 1864)
OWB
Paraplectana walleri (Blackwall,
1865)
Pararaneus cyrtoscapus
(Pocock, 1898)
Pararaneus perforatus (Thorell,
1899)
Pararaneus spectator (Karsch,
1885)
Prasonica seriata Simon, 1895
Singa albodorsata Kauri, 1950
ARCHAEIDAE
Afrarchaea godfreyi (Hewitt,
1919)
Afrarchaea ngomensis Lotz,
1996
CAPONIIDAE
Caponia chelifera Lessert, 1936
CLUBIONIDAE
Clubiona abbajensis Strand,
1906
Clubiona africana Lessert, 1921
OWB
Clubiona annuligera Lessert,
1929
Clubiona pupillaris Lawrence,
1938
CORINNIDAE
Afroceto arca Lyle & Haddad,
2010
Afroceto martini (Simon, 1897)
Cambalida dippenaarae
Haddad, 2012
Cambalida fulvipes (Simon,
1896)
Castianeira sp.
OWB
OWB
avocado, citrus, pistachio,
tomatoes
maize
OWB
maize, strawberries
OWB
OWB
pistachio, tomatoes
tomatoes
GW
pine plantations
GW
pine plantations
GW
cotton, pine plantations
PW
PW
avocado, citrus, lemon,
macadamia
avocado, citrus,
macadamia, tomatoes,
vineyards
maize
PW
citrus, cotton
GW
citrus
GW
GW
macadamia, pistachio
citrus, cotton, grapefruit,
maize
maize, pistachio
PW
GW
GW
Copa flavoplumosa Simon,
1885
GW
Copuetta lacustris (Strand,
1916)
Corinnomma semiglabrum
(Simon, 1896)
Fuchiba aquilonia Haddad &
Lyle, 2008
Graptartia mutillica Haddad,
2004
Hortipes luytenae Bosselaers &
Ledoux, 1998
Hortipes merwei Bosselaers &
Jocqué, 2000
Hortipes schoemanae
Bosselaers & Jocqué, 2000
Orthobula radiata Simon, 1897
GW
cotton, pistachio, sunflower,
tomatoes
avocado, citrus, cotton,
macadamia, maize,
pistachio, strawberries
maize, pistachio, vineyards
GW
citrus
GW
citrus
GW
maize
GW
pine plantations
GW
pine plantations
GW
pine plantations
GW
cotton
A.S. Dippenaar-Schoeman et al.: Knowledge of Spider Diversity in Agroecosystems
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APPENDIX 1 (Continued )
13
APPENDIX 1 (Continued )
Species
Guild
Crops
Species
Guild
Crops
Poachelas striatus Haddad &
Lyle, 2008
Pronophaea natalica Simon,
1897
Pronophaea sp.
Thysanina capensis Lyle &
Haddad, 2006
Thysanina transversa Lyle &
Haddad, 2006
Trachelas pusillus Lessert,
1923
CTENIDAE
Anahita sp.
Ctenus pulchriventris (Simon,
1896)
CYATHOLIPIDAE
Isicabu zuluensis Griswold,
1987
Ulwembua denticulata
Griswold, 1987
CYRTAUCHENIIDAE
Ancylotrypa pusilla (Purcell,
1903)
Ancylotrypa sp.
Ancylotrypa vryheidensis
(Hewitt, 1915)
Homostola zebrina Purcell,
1902
DEINOPIDAE
Deinopis cornigera Gerstäcker,
1873
Menneus camelus Pocock,
1902
DICTYNIDAE
Archaeodictyna sp.
Dictyna sp.
GW
maize
GW
macadamia, pear, pistachio
GW
pine plantations
GW
pistachio
GW
GW
cotton
pine plantations
GW
cabbage
GW
macadamia
PW
cotton, maize, pistachio,
tomatoes
GW
GW
GW
GW
pistachio
maize
cotton, pistachio
mango, pine plantations,
pistachio
citrus, cotton, pistachio
GW
GW
maize
citrus, maize, pine
plantations, strawberries
Drassodes lophognathus
Purcell, 1907
Drassodes sesquidentatus
Purcell, 1908
Drassodes stationis Tucker,
1923
Echemus erutus Tucker, 1923
Ibala arca (Tucker, 1923)
Ibala bilinearis (Tucker, 1923)
Latonigena africanus Tucker,
1923
Megamyrmaekion
transvaalense Tucker, 1923
Micaria sp.
OWB
pine plantations
OWB
avocado, citrus
GW
pistachio
GW
GW
cotton, maize
pine plantations
GW
pine plantations
OWB
avocado
OWB
avocado
DYSDERIDAE
Dysdera crocata C.L. Koch,
1838
ERESIDAE
Stegodyphus dumicola Pocock,
1898
GALLIENIELLIDAE
Austrachelas bergi Haddad,
Lyle, Bosselaers & Ramirez,
2009
Austrachelas natalensis
Lawrence, 1942
GNAPHOSIDAE
Aneplasa nigra Tucker, 1923
Asemesthes ceresicola Tucker,
1923
Asemesthes decoratus Purcell,
1908
Asemesthes lineatus Purcell,
1908
Asemesthes purcelli Tucker,
1923
Camillina aldabrae (Strand,
1907)
Camillina cordifera (Tullgren,
1910)
Camillina corrugata (Purcell,
1907)
Camillina setosa Trucker,
1923
Drassodes ereptor Purcell,
1907
RWB
RWB
pistachio
cabbage, citrus, cotton,
tomatoes
GW
pear
RWB
citrus
GW
avocado
GW
tomatoes
GW
GW
pistachio
cotton, tomatoes
GW
cotton
GW
pistachio
GW
pistachio
GW
maize
GW
GW
citrus, cotton, maize,
pistachio, sunflower
pistachio
GW
sugar cane
GW
pistachio
Nomisia varia (Tucker, 1923)
Odontodrassus aphanes
(Thorell, 1897)
Poecilochroa capensis Strand,
1909
Pterotricha auris (Tucker, 1923)
Pterotricha varius (Tucker,
1923)
Scotophaeus relegatus Purcell,
1907
Setaphis browni (Tucker, 1923)
Setaphis subtilis (Simon, 1897)
Trachyzelotes jaxartensis
(Kroneberg, 1875)
Trephopoda hanoveria Tucker,
1923
Upognampa biamenta Tucker,
1923
Upognampa parvipalpa Tucker,
1923
Urozelotes rusticus (L. Koch,
1872)
Xerophaeus capensis Purcell,
1907
Xerophaeus pallidus Tucker,
1923
Xerophaeus vickermani Tucker,
1923
Zelotes bastardi (Simon, 1896)
Zelotes corrugatus (Purcell,
1907)
Zelotes frenchi Tucker, 1923
Zelotes fuligineus (Purcell,
1907)
Zelotes humilis (Purcell, 1907)
Zelotes natalensis Tucker, 1923
Zelotes oneilli (Purcell, 1907)
Zelotes pallidipes Tucker, 1923
Zelotes sclateri Tucker, 1923
Zelotes scrutatus (O.P.Cambridge, 1872)
Zelotes uquathus Fitzpatrick,
2007
HAHNIIDAE
Hahnia lobata Bosmans, 1981
Hahnia schubotzi Strand, 1913
Hahnia tabulicola Simon, 1898
HERSILIIDAE
Hersilia sericea Pocock, 1898
Tyrotama bicava (Smithers,
1945)
GW
GW
GW
GW
apple, maize, pistachio,
potatoes
pistachio
cotton
GW
pistachio
GW
GW
maize, pistachio, vineyard
pistachio
GW
cotton, vineyard
GW
GW
GW
cotton, pistachio
cabbage, citrus, cotton,
maize, pistachio, tomatoes
cotton, grapefruit, minneola
GW
pistachio
GW
vineyards
GW
cotton
GW
cabbage
GW
vineyards
GW
pine plantations
GW
pistachio
GW
GW
cabbage, cotton
pistachio
GW
GW
cotton
citrus, pistachio
GW
GW
GW
pine plantations
avocado, citrus, cotton,
sunflower
pistachio
cotton
tomatoes
citrus, cotton, maize,
pistachio, sunflower
citrus
SWB
SWB
SWB
pine plantation
apple
cotton, maize
PW
avocado, citrus, grapefruit,
macadamia
cotton
GW
GW
GW
GW
PW
14
Transactions of the Royal Society of South Africa
APPENDIX 1 (Continued )
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Species
IDIOPIDAE
Ctenolophus fenoulheti Hewitt,
1913
Gorgyrella schreineri Purcell,
1903
LINYPHIIDAE
Ceratinopsis dippenaari
Jocqué, 1984
Ceratinopsis idanrensis Locket
& Russell-Smith, 1980
Erigone irrita Jocqué, 1984
Erigone prominens Bosenberg
& Strand, 1906
Limoneta sirimoni (Bosmans,
1979)
Meioneta habra Locket, 1968
Meioneta natalensis Jocqué,
1984
Meioneta prosectes Locket,
1968
Meioneta prosectoides Locket
& Russell-Smith, 1980
Mermessus fradeorum
(Berland, 1932)
Metaleptyphantes familiaris
Jocqué, 1984
Metaleptyphantes perexiguus
(Simon & Fage, 1922)
Microlinyphia sterilis (Pavesi,
1883)
Ostearius melanopygius (O.P.Cambridge, 1879)*
APPENDIX 1 (Continued )
Guild
Crops
GW
cotton
GW
pistachio
SWB
citrus
SWB
cotton
SWB
SWB
SWB
citrus, cotton
cotton, maize, strawberries,
tomatoes
lucerne, maize,
strawberries
cotton, lucerne, maize,
minneola, pecans,
pistachio, strawberries,
sunflowers, tomatoes
cotton, pine plantations
SWB
potatoes
SWB
cotton
SWB
cotton, tomatoes kenaf,
pistachio
apple, cotton, lucerne,
pistachio
citrus, cotton
SWB
SWB
SWB
SWB
SWB
SWB
avocado, cotton, kenaf,
lucerne, pecans, pistachio,
strawberries
apple, cabbage, citrus,
cotton, maize, pistachio,
strawberries, tomatoes,
vineyards
cotton, kenaf, lucerne,
maize, pistachio, sorghum,
vineyards
citrus
SWB
cotton, kenaf, potatoes
SWB
citrus, cotton
GW
pistachio
SWB
Pelecopsis janus Jocqué, 1984 SWB
Tybaertiella convexa (Holm,
1962)
Tybaertiella krugeri (Simon,
1894)
Typhistes gloriosus Jocqué,
1984
LIOCRANIDAE
Rhaeboctesis trinotatus Tucker,
1920
LYCOSIDAE
Allocosa lawrencei (Roewer,
1951)
Allocosa tuberculipalpa
(Caporiacco, 1940)
Amblyothele ecologica RussellSmith, Jocqué &
Alderweireldt, 2009
Arctosa promontorii (Pocock,
1900)
Evippa sp.
Evippomma squamulatum
(Simon, 1898)
Foveosa foveolata (Purcell,
1903)
Hippasa affinis Lessert, 1933
Hogna bimaculata (Purcell,
1903)
Hogna spenceri (Pocock,
1898)
GW
vineyards
GW
citrus, grapefruit
GW
tomatoes
GW
vineyards
GW
GW
pistachio
pistachio
GW
citrus, pistachio
GW
GW
maize
cotton
GW
cotton
Species
Guild
Crops
Hogna transvaalica (Simon,
1898)
Lycosa sp.
Ocyale guttata (Karsch, 1878)
Pardosa crassipalpis Purcell,
1903*
GW
cotton, maize, strawberries
GW
GW
GW
GW
GW
GW
strawberry
tomatoes
apple, cabbage, citrus,
cotton, lucerne, maize,
pear, pecans, pine
plantations, pistachio,
potatoes, sorghum,
strawberries, sugar cane,
sunflower, tomatoes
litchi, strawberries
cotton
pistachio, vineyards
GW
vineyards
GW
cotton
GW
cotton, maize, tomatoes
GW
GW
pistachio
cotton
GW
maize
GW
pine plantations
GW
pine plantations
PW
avocado, citrus, cotton,
pine plantations
PW
cotton
PW
PW
avocado, citrus, cotton,
lucerne, macadamia,
maize, mango, pecans,
pistachio, potatoes,
strawberries, tomatoes,
vineyards
maize
PW
cotton, pistachio
PW
PW
avocado, citrus, minneola,
tomatoes
macadamia
PW
pine plantations
OWB
citrus, prickly pear
GWB
macadamia, maize,
sunflower, tomatoes
GW
citrus
GW
pine plantations
GW
cotton, pine plantations,
sunflower
GW
pistachio
GW
pine plantations
Pardosa oncka Lawrence, 1927
Pardosa umtalica Purcell, 1903
Pterartoria arbuscula (Purcell,
1903)
Pterartoriola sagae (Purcell,
1903)
Schizocosa darlingi (Pocock,
1898)
Trabea natalensis RussellSmith, 1982
Trabea purcelli Roewer, 1951
Trochosa albipilosa (Roewer,
1960)
Zenonina mystacina Simon,
1898
MICROSTIGMATIDAE
Microstigmata longipes
(Lawrence, 1938)
Microstigmata zuluensis
(Lawrence, 1938)
MIMETIDAE
Mimetus natalensis Lawrence,
1938
MITURGIDAE
Cheiracanthium dippenaarae
Lotz, 2007
Cheiracanthium furculatum
Karsch, 1879*
Cheiracanthium molle L. Koch,
1875
Cheiracanthium vansoni
Lawrence, 1936
Cheiramiona krugerensis Lotz,
2002
Cheiramiona paradisus Lotz,
2002
Cheiramiona silvicola
(Lawrence, 1938)
NEPHILIDAE
Nephila fenestrata Thorell,
1859
NESTICIDAE
Nesticella benoiti (Hubert,
1970)
OONOPIDAE
Dysderina speculifera Simon,
1907
Gamasomorpha australis
Hewitt, 1915
Opopaea speciosa (Lawrence,
1952)
ORSOLOBIDAE
Afrilobus australis Griswold &
Platnick, 1987
Azanialobus lawrencei
Griswold & Platnick, 1987
A.S. Dippenaar-Schoeman et al.: Knowledge of Spider Diversity in Agroecosystems
APPENDIX 1 (Continued )
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Species
OXYOPIDAE
Hamataliwa kulczynskii
(Lessert, 1915)
Oxyopes bothai Lessert, 1915
Oxyopes flavipalpis (Lucas,
1858)
Oxyopes hoggi Lessert, 1915
Oxyopes jacksoni Lessert,
1915
Oxyopes longispinosus
Lawrence, 1938
Oxyopes pallidecoloratus
Strand, 1906
Oxyopes schenkeli Lessert,
1927
Peucetia viridis (Blackwall,
1858)
PALPIMANIDAE
Diaphorocellus biplagiatus
Simon, 1893
Palpimanus transvaalicus
Simon, 1893
PHILODROMIDAE
Gephyrota sp.
Hirriusa arenacea (Lawrence,
1927)
Philodromus brachycephalus
Lawrence, 1952
Philodromus browningi
Lawrence, 1952
Philodromus guineensis Millot,
1942
Philodromus maestrii
Caporiacco, 1941
Philodromus thanatellus
Strand, 1909
Thanatus dorsilineatus
Jezequel, 1964
Thanatus vulgaris Simon, 1870
Tibellus minor Lessert, 1919
PHOLCIDAE
Quamtana bonamanzi Huber,
2003
Quamtana ciliata (Lawrence,
1938)
Quamtana embuleni Huber,
2003
Quamtana merwei Huber, 2003
Smeringopus natalensis
Lawrence, 1947
Smeringopus sambesicus
Kraus, 1957
PHYXELIDIDAE
Pongolania chrysionaria
Griswold, 1990
Vidole capensis (Pocock, 1900)
Vidole sothoana Griswold, 1990
Xevioso aululata Griswold,
1990
Xevioso colobata Griswold,
1990
Xevioso kulufa Griswold, 1990
Xevioso tuberculata (Lawrence,
1939)
PISAURIDAE
Charminus aethiopicus
(Caporiacco, 1939)
15
APPENDIX 1 (Continued )
Guild
Crops
PW
avocado, macadamia
PW
PW
cotton, pistachio
sugar cane
PW
PW
pistachio
citrus, macadamia, maize,
sunflower
macadamia
PW
PW
PW
PW
cabbage, citrus, maize,
tomatoes
avocado, citrus,
macadamia
pistachio
GW
pistachio
GW
citrus, cotton
PW
GW
macadamia, maize,
pistachio, sunflower,
tomatoes
pistachio
GW
avocado, macadamia
PW
pecans, pistachio
GW
avocado, citrus
GW
mango
GW
vineyards
GW/
PW
GW
maize
PW
cotton, lucerne, maize,
potatoes, strawberries
cotton, maize, vineyards
SpWB citrus
SpWB citrus, pine plantations
SpWB citrus
SpWB pine plantations
SpWB cotton, pistachio
SpWB citrus
RWB
citrus, maize
RWB
RWB
RWB
citrus
cotton, maize
pine plantations
RWB
citrus
RWB
RWB
macadamia
citrus
PW
macadamia, maize
Species
Guild
Crops
Chiasmopes lineatus (Pocock,
1898)
Maypacius curiosus Blandin,
1975
Perenethis simoni (Lessert,
1916)
Perenethis symmetrica
(Lawrence, 1927)
Rothus purpurissatus Simon,
1898
Rothus vittatus Simon, 1898
PRODIDOMIDAE
Prodidomus purpurascens
Purcell, 1904
Theuma fusca Purcell, 1907
SALTICIDAE
Aelurillus sp.
Afraflacilla sp.
Asemonea maculata Wanless,
1980
Baryphas ahenus Simon, 1902
SWB
maize
SWB
maize
SWB
citrus
SWB
maize
PWB
lucerne, pecans, vineyards
PWB
pistachio
GW
cotton
GW
pistachio
GW
PW
PW
cotton
macadamia
avocado, lemon
PW
GW
citrus, maize, pine
plantations, strawberries,
tomatoes
cotton
PW
GW
avocado, cotton
vineyards
GW
PW
maize
pine plantations
PW
pine plantations
GW
GW
pistachio
tomatoes
PW
cotton, kenaf, maize
GW
apple, citrus, kenaf
PW
PW
avocado, citrus,
macadamia
lucerne, pine plantations,
pistachio
cotton, pistachio
PW
potatoes
PW
cotton, kenaf, sorghum
PW
citrus, cotton, kenaf
PW
lucerne, vineyards
PW
cotton, pecans, potatoes,
tomatoes
cotton
Bianor albobimaculatus (Lucas,
1846)
Brancus bevisi (Lessert, 1925)
Cyrba dotata Peckham &
Peckham, 1903
Cyrba nigrimana Simon, 1900
Dendryphantes hararensis
Wesolowska & Cumming,
2008
Dendryphantes purcelli
Peckham & Peckham, 1903
Euophrys sp.
Evarcha culicivora Wesolowska
& Jackson, 2003
Evarcha dotata (Peckham &
Peckham, 1903)
Evarcha flagellaris Haddad &
Wesolowska, 2011
Goleba puella (Simon, 1885)
Heliophanus charlesi
Wesolowska, 2003
Heliophanus debilis Simon,
1901
Heliophanus fascinatus
Wesolowska, 1986
Heliophanus hastatus
Wesolowska, 1986
Heliophanus insperatus
Wesolowska, 1986
Heliophanus modicus Peckham
& Peckham, 1903
Heliophanus nanus
Wesolowska, 2003
Heliophanus orchesta Simon,
1886
Heliophanus pistaciae
Wesolowska, 2003*
PW
PW
PW
Heliophanus proszynski
PW
Wesolowska, 2003
Heliophanus trepidus Simon,
PW
1910
Hyllus argyrotoxus Simon, 1902 PW
Hyllus brevitarsis Simon, 1902
PW
cotton, kenaf, pecans,
pistachio, potatoes,
pumpkin, sugar cane
cotton, tomatoes, wheat
cotton, pistachio
avocado, citrus, cotton,
macadamia
avocado, lemon,
macadamia
16
Transactions of the Royal Society of South Africa
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APPENDIX 1 (Continued )
APPENDIX 1 (Continued )
Species
Guild
Crops
Species
Guild
Crops
Icius insolidus Wesolowska,
1999
Langona manicata Simon, 1901
Menemerus bifurcus
Wesolowska, 1999
Menemerus pilosus
Wesolowska, 1999
Myrmarachne ichneumon
Simon, 1886
Myrmarachne marshalli
Peckham & Peckham, 1903
Myrmarachne solitaria
Peckham & Peckham, 1903
Myrmarachne uvira Wanless,
1978
Natta horizontalis Karsch, 1879
Nigorella hirsuta Wesolowska,
2009
Pachyballus castaneus Simon,
1900
Pachyballus sp.
Pachyballus transversus
Simon, 1900
Parajotus obscurofemoratus
Peckham & Peckham, 1903
Pellenes bulawayoensis
Wesolowska, 1999
Pellenes geniculatus (Simon,
1868)
Phintella aequipes (Peckham &
Peckham, 1903)
Phlegra certa Wesolowska &
Haddad, 2009
Phlegra karoo Wesolowska,
2006
Phlegra nuda Próchiewicz &
Heciak, 1994
Phlegra simplex Wesolowska &
Russell-Smith, 2000
Portia schultzi Karsch, 1878
Pseudicius africanus Peckham
& Peckham, 1903
Rhene machadoi Berland &
Millot, 1941
Stenaelurillus cristatus
Wesolowska & RussellSmith, 2000
Stenaelurillus guttiger (Simon,
1901)
Thyene aperta (Peckham &
Peckham, 1903)
Thyene bucculenta
(Gerstäcker, 1873
Thyene coccineovittata (Simon,
1886)
Thyene inflata (Gerstäcker,
1873)
Thyene natalii Peckham &
Peckham, 1903
Thyene ogdeni Peckham &
Peckham, 1903
Thyene semiargentea (Simon,
1884)
Thyene thyenoides (Lessert,
1925)
Thyenula aurantiaca (Simon,
1902)
Thyenula fidelis Wesolowska &
Haddad, 2009
Tusitala barbata Peckham &
Peckham, 1902
GW
pistachio
PW
avocado, macadamia
GW
PW
cotton, maize
cotton
PW
PW
avocado, citrus,
macadamia
macadamia
PW
pistachio
PW
avocado, macadamia
PW
citrus
PW
macadamia
PW
citrus, minneola, pecan
PW
lucerne
PW
cotton
GW
GW
pine plantations
maize
GW
GW
GW
avocado, citrus, mango
GW
pine plantations
PW
cotton, pistachio
cotton, maize, pecans,
strawberries
citrus
PW
PW
lucerne, onion
minneola
TWB
avocado, citrus, pistachio
PW
PW
macadamia
PW
avocado, citrus,
macadamia
pine plantations
GW
cotton, kenaf, pecans,
potatoes
cotton, lucerne, maize
Tusitala guineensis Berland &
Millot, 1941
Tusitala hirsuta Peckham &
Peckham, 1902
Veissella durbani (Peckham &
Peckham, 1903)
Viciria alba Peckham &
Peckham, 1903
Zulunigma incognita
(Wesolowska & Haddad,
2009)
SCYTODIDAE
Scytodes caffra Purcell, 1904
Scytodes elizabethae Purcell,
1904
Scytodes flagellata Purcell,
1904
Scytodes maritima Lawrence,
1938
SEGESTRIIDAE
Ariadna spp.
SELENOPIDAE
Anyphops fitzsimonsi
(Lawrence, 1940)
Anyphops lawrencei (Roewer,
1951)
Anyphops minor (Lawrence,
1940)
Anyphops rubicundus
(Lawrence, 1940)
Selenops radiatus Latreille,
1819
SICARIIDAE
Loxosceles spinulosa Purcell,
1904
SPARASSIDAE
Olios auricomis (Simon, 1880)
Olios sjostedti Lessert, 1921
Olios tuckeri Lawrence, 1927
Palystes superciliosus L. Koch,
1875
Panaretella minor Lawrence,
1952
Palystes ansiedippenaarae
Croeser, 1996
Palystes superciliosus L. Koch,
1875
TETRAGNATHIDAE
Leucauge decorata (Blackwall,
1864)
Leucauge festiva (Blackwall,
1866)
Leucauge medjensis Lessert,
1930
Leucauge thomeensis Kraus,
1960
Pachygnatha leleupi Lawrence,
1952
Tetragnatha jaculator Tullgren,
1910
Tetragnatha subsquamata
Okuma, 1985
THERAPHOSIDAE
Brachionopus tristis Purcell,
1903
THERIDIIDAE
Achaearanea sp.
Anelosimus nelsoni Agnarsson,
2006
Argyrodes convivans
Lawrence, 1937
PW
sugar cane
PW
avocado, macadamia
PW
potatoes
GW
cotton
PW
PW
PW
PW
PW
avocado, macadamia
macadamia
macadamia
avocado, cabbage,
macadamia
citrus
PW
pine plantations
PW
pine plantations
OWB
citrus, maize
OWB
avocado, macadamia,
pumpkin, tomatoes
citrus
GW
PW
GW
avocado, grapefruit,
macadamia
cotton
GW
pecans, pistachio
GW
cotton
GW
cotton, maize, tomatoes
PW
PW
macadamia
pine plantations
PW
lucerne
GW
maize
GW
cotton
PW
pistachio
PW
cotton, lucerne
PW
PW
avocado, citrus, grapefruit,
macadamia
avocado, citrus, cotton,
grapefruit, pistachio
avocado, citrus,
macadamia, tomatoes
citrus
PW
cotton
PW
pecans
PW
avocado, macadamia
GW
pine plantation
PW
citrus, pistachio
PW
PW
OWB
OWB
OWB
avocado, citrus,
macadamia
maize, potatoes
OWB
cotton, maize
OWB
avocado, macadamia,
maize
GW
citrus
GWB
GWB
citrus, vineyards
citrus
GWB
citrus, cotton
A.S. Dippenaar-Schoeman et al.: Knowledge of Spider Diversity in Agroecosystems
APPENDIX 1 (Continued )
APPENDIX 1 (Continued )
Species
Guild
Crops
Species
Guild
Crops
Coleosoma blandum O.P.Cambridge, 1882
Dipoena sp.
Dipoenura sp.
Enoplognatha molesta O.P.Cambridge, 1904*
GWB
grapefruit, minneola
PW
citrus
GWB
GWB
GWB
citrus, cotton
citrus, tomatoes
citrus, cotton, maize,
potatoes, sorghum,
strawberries, sugarcane,
tomatoes
vineyards
Phrynarachne melloleitaoi
Lessert, 1933
Runcinia aethiops (Simon,
1901)
Runcinia affinis Simon, 1897
PW
pine plantations,
strawberries
cotton, lucerne,
strawberries
pistachio, potatoes, sugar
cane, strawberries
kenaf
Euryopis episinoides
(Walckenaer, 1847)
Euryopis sp.
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17
Latrodectus cinctus Blackwall,
1865
Latrodectus geometricus C.L.
Koch, 1841
Latrodectus indistinctus O.P.Cambridge, 1904
Latrodectus renivulvatus Dahl,
1902
Phoroncidia sp.
Steatoda capensis Hann, 1990
Steatoda erigoniformis (O.P.Cambridge, 1872)
Steatoda grossa (C.L. Koch,
1838)
Theridion pictum (Walckenaer,
1802)
Theridion purcelli O.P.Cambridge, 1904
Theridula sp.
Tidarren cuneolatum (Tullgren,
1910)
THOMISIDAE
Ansiea tuckeri (Lessert, 1919)
Borboropactus silvicola
(Lawrence, 1938)
Camaricus nigrotesselatus
Simon, 1895
Diaea puncta Karsch, 1884
Firmicus bragantinus (Brito
Capello, 1866)
Heriaeus transvaalicus Simon,
1895
Misumenops rubrodecoratus
Millot, 1941*
Monaeses austrinus Simon,
1910
Monaeses fuscus DippenaarSchoeman, 1984
Monaeses quadrituberculatus
Lawrence, 1927
Oxytate argenteooculata
(Simon, 1886)
Oxytate concolor (Caporiacco,
1947)
Ozyptila caenosa Jézéquel,
1966
Parasmodix quadrituberculata
Jézéquel, 1966
Pherecydes tuberculatus O.P.Cambridge, 1883
GWB
GWB
GWB
GWB
GWB
GWB
cotton, minneola, maize,
pear
citrus, tomatoes
avocado, citrus, cotton,
maize, pistachio, prickly
pears, vineyards
pistachio
GWB
cotton, maize, strawberries,
sugarcane, vineyards
citrus, tomatoes
maize, pear, tomatoes, pine
plantations, vineyards
cotton, maize, tomatoes
GWB
cotton
GWB
avocado, macadamia
GWB
citrus, cotton, macadamia,
maize, strawberries,
sunflower, tomatoes
citrus, cotton
avocado, citrus,
macadamia, pistachio
GWB
GWB
GWB
GWB
PW
PW
avocado, macadamia, pine
plantations
maize
PW
tomatoes
PW
avocado, cotton, pistachio,
strawberries
avocado, citrus, grapefruit
PW
PW
PW
avocado, cotton,
strawberries
avocado, citrus, cotton,
kenaf, lucerne, macadamia,
maize, pecans, pine
plantations, pistachio,
pumpkin, sugar cane,
sunflower, strawberries,
tomatoes
cotton, pistachio
PW
cotton, potatoes
PW
lucerne, pistachio
PW
PW
avocado, citrus,
macadamia, tomatoes
avocado, citrus,
macadamia
maize
PW
maize
PW
lucerne, pine plantations
PW
PW
PW
Runcinia depressa Simon,
PW
1906
Runcinia erythrina Jézéquel,
PW
1964
Runcinia flavida (Simon, 1881) PW
Runcinia grammica (L. Koch,
1937)
Simorcus cotti Lessert, 1936
Sylligma theresa Lewis &
Dippenaar-Schoeman, 2011
Synema decens (Karsch, 1878)
Synema diana (Audouin, 1826)
Synema imitator (Pavesi, 1883)
Synema langheldi Dahl, 1907
PW
pine plantations,
strawberries
tomatoes
PW
PW
citrus
maize
PW
PW
PW
PW
tomatoes
tomatoes
tomatoes
citrus, macadamia,
minneola
citrus
Thomisops senegalensis Millot, PW
1941
Thomisus australis Comellini,
PW
1957
Thomisus blandus Karsch,
PW
1880
Thomisus citrinellus Simon,
1875
Thomisus congoensis
Comellini, 1957
Thomisus dalmasi Lessert,
1919
Thomisus daradioides Simon,
1890
Thomisus granulatus Karsch,
1880
Thomisus kalaharinus
Lawrence, 1936
Thomisus machadoi Comellini,
1959
Thomisus scrupeus (Simon,
1886)
Thomisus stenningi Pocock,
1900
Thomisus unidentatus
Dippenaar-Schoeman & Van
Harten, 2007
Tmarus cameliformis Millot,
1941
Tmarus comellinii Garcia-Neto,
1989
Tmarus foliatus Lessert, 1928
Tmarus natalensis Lessert,
1925
Xysticus natalensis Lawrence,
1938
Xysticus urbensis Lawrence,
1952
TROCHANTERIIDAE
Platyoides walteri (Karsch,
1886)
ULOBORIDAE
Uloborus plumipes Lucas, 1846
PW
peach, pine plantations,
tomatoes
citrus, cotton, maize,
papaya, pumpkin,
strawberries
pecans
PW
cotton, minneola
PW
lucerne, strawberries
PW
citrus, macadamia
PW
citrus
PW
pistachio
PW
pistachio
PW
avocado, citrus, cotton,
minneola, sunflower
cotton, potatoes, lucerne,
pecans, pine plantations,
pistachio, strawberries,
wheat
citrus
PW
PW
PW
PW
avocado, citrus, cotton,
macadamia
avocado, mango
PW
PW
sugar cane
macadamia
GW
GW
avocado, cotton,
strawberries, sugar cane
maize
PW
avocado, pine plantations
OWB
avocado, citrus, cotton,
pistachio, prickly pear,
strawberries, tomatoes
18
Transactions of the Royal Society of South Africa
APPENDIX 1 (Continued )
Downloaded by [ARC Central Office], [Professor A.S. Dippenaar-Schoeman] at 04:25 16 January 2013
Species
ZODARIIDAE
Cydrela schoemanae Jocqué,
1991
Diores recurvatus Jocqué, 1990
Diores strandi Caporiacco,
1949
Diores triangulifer Simon, 1910
Heradida sp.
Microdiores sp.
Ranops caprivi Jocqué, 1991
ZOROPSIDAE
Phanotea cavata Giswold,
1994
Guild
Crops
GW
maize, pine plantations
GW
GW
cotton
maize
GW
GW
GW
GW
pistachio
cotton
citrus, maize
cotton
RWB
citrus