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Unformatted text preview: doi 10.1098/rspb.2001.1709 Stabilimenta attract unwelcome predators
Wee Khee Seah and Daiqin Li*
Department of Biological Sciences, National University of Singapore, Singapore 119260
Conspicuous behaviour exposes animals to predation; prey-attraction thus often con£icts with
antipredator behaviour. The fact that a conspicuous ultraviolet-light re£ecting silken structure in the
orb-webs of certain spider species, known as a stabilimentum, makes the webs obvious to both prey and
predators has been used to argue that spiders bene¢t from building stabilimenta by attracting prey
and/or defending against visually hunting predators. Here, we provide experimental evidence that
stabilimenta can act as visual signals that attract web-invading spider-eating predators with acute
vision to the webs. We also show that the predators can learn to remember a particular type of
stabilimentum. Thus, stabilimentum-building spiders risk a high level of predation by attracting visually
Keywords: Argiope versicolor ; silk; stabilimenta; spider; Portia labiata ; predator attraction
1. INTRODUCTION Biologists have always been fascinated by the spider's web
in all its various forms (see Foelix 1996) and long been
puzzled by the function of the so-called stabilimenta in
the webs of certain orb-weaving spiders (Robinson &
Robinson 1970). The orb-web is well known as a device
used in foraging as well as in defending against predators
olbert 1975; Edmunds & Edmunds 1986; Eberhard
1990; Blackledge & Wenzel 1999). The stabilimentum,
however, is an extra silk structure that is added by certain
species of spider to the central portion of the orb-web
after web construction. Why these spiders, but not others,
include stabilimenta in their webs has been debated for
more than a century (Simon 1895). Although many
functional hypotheses have been proposed for silk stabilimenta, the most recent and plausible, but also contentious, hypotheses are the prey-attraction hypothesis
(Craig & Bernard 1990; Elgar et al. 1996; Tso 1996, 1998;
Watanabe 1999) and the predator-defence hypothesis
(Horton 1980; Eisner & Nowicki 1983; Schoener &
Spiller 1992; Kerr 1993; Blackledge 1998a ; Blackledge &
Wenzel 1999). Interestingly, both hypotheses have used
the fact that stabilimenta have a bright re£ectance across
wavelengths of light visible to some animals, including
ultraviolet (UV) light (Craig & Bernard 1990), to argue
that stabilimenta act as visual signals that make webs
more conspicuous either to prey or to predators. Thus,
spiders bene¢t from building conspicuous stabilimenta by
attracting prey to the webs, by advertising the presence of
sticky webs to predators or by distracting predators.
However, building stabilimenta may be costly because
stabilimenta provide visual cues that insect prey use to
avoid webs, thus reducing the web builder's foraging
e¤ciency (Blackledge 1998a ; Blackledge & Wenzel 2000,
2001). However, predators can also potentially perceive
these conspicuous visual signals. Therefore, one may
hypothesize that the stabilimenta could also operate to
the detriment of a spider by attracting unwelcome visual
predators to the orb-web, if UV-re£ecting stabilimenta
* Author for correspondence ([email protected]). Proc. R. Soc. Lond. B (2001) 268, 1553^1558
Received 20 December 2000 Accepted 12 April 2001 make webs more conspicuous. Surprisingly, no e¡ort has
been made to test this hypothesis. Here, we experimentally test this hypothesis using a species of orb-web
weaving spider, Argiope versicolor (Doleschall) (Araneae:
Araneidae), and a species of predator, Portia labiata
(Thorell), a web-invading and spider-eating jumping
spider (Araneae: Salticidae) from Singapore.
Like other Argiope species, A. versicolor build di¡erent
types of stabilimenta, including disc-like stabilimenta built
only by juveniles and cruciforms created by adults. A total
of ¢ve di¡erent types of stabilimenta have been described
in A. versicolor (¢gure 1; W. K. Seah and D. Li, unpublished
data). It is known that webs with stabilimenta are able to
attract prey (Craig & Bernard 1990; Tso 1996, 1998;
Watanabe 1999; but see Blackledge & Wenzel 1999). As
mentioned above, conspicuous signals may also attract
the attention of predators (Eisner & Nowicki 1983).
P. labiata is sympatric with A. versicolor and is known to
enter the webs of A. versicolor to prey on it, both in nature
and in the laboratory (D. Li, unpublished data). Like
other salticids, Portia spp. di¡er from most spiders in
having remarkably acute vision (Blest et al. 1990) that
permits them to discriminate visually between di¡erent
prey, predators and conspeci¢cs (Jackson & Blest 1982; Li
& Jackson 1996). Jumping spiders are also known to be
dichromatic, having the physiological capacity to distinguish colours using the UV-sensitive and greensensitive cells in their principal (anterior median) eyes
(Land 1969; DeVoe 1975; Nakamura & Yamashita 2000).
On the basis of visual cues alone, Portia spp. can distinguish a web from an empty cage, and can also distinguish between quiescent spiders, insects and egg sacs
(Jackson 1995; Jackson & Pollard 1996; Li & Jackson
1996). Therefore, we assume that P. labiata is able to
distinguish between decorated and undecorated webs and
that it can make a choice as to which web it wants to
In this study, we use P. labiata as a predator of
A. versicolor to test three hypotheses: ¢rst, that P. labiata
can use conspicuous UV-light re£ecting stabilimenta to
locate and catch A. versicolor ; second, that the presence of
A. versicolor has no e¡ect on the attraction of P. labiata by 1553 & 2001 The Royal Society 1554 W. K. Seah and D. Li Stabilimenta attract predators
experience of A. versicolor prior to testing. They were kept
individually in plastic cylindrical cages (diameter Â height:
100 mm Â 80 mm) and provided with a diet of fruit £ies
(Drosophila melanogaster Meigen) and house£ies (Musca domestica
L.) ad libitum before testing. No individual P. labiata was used in
more than one test of any one type. (b) Predator-attraction test Figure 1. Webs and forms of stabilimenta of Argiope versicolor.
The spiders either spin a new decoration each day, which
may vary in design, or do not decorate their webs at all.
(a) Disk-like (discoid) stabilimentum built by juveniles,
(b^e) cruciform stabilimenta with four, three, two and one
arms, respectively, and ( f ) a web without a stabilimentum.
Scale bar: ca. 1 cm. stabilimenta; and third, like stingless bees that can learn
to avoid webs (Craig 1994), P. labiata is able to remember
a particular type of decorated web.
2. METHODS (a) General Maintenance procedures, cage design, basic testing methods
and terminology were as reported in earlier spider studies (see
Jackson & Hallas 1986) and only essential details are given here.
All experiments were carried out in a controlled laboratory
(relative humidity, 80^90%; temperature, 24 Æ 1 8C; 12 L:12 D
photoperiod; laboratory light intensity, 332.8 Æ 4.3 lux).
We collected A. versicolor juveniles and females from Bukit
Timah Nature Reserve in Singapore in 1999. They were kept
individually in plastic-framed cages with removable glass sides
(200 mm Â 200 mm Â 50 mm). Wooden sticks were connected to
the sides of the cages to provide attachment sites for the webs.
A. versicolor fastens web lines primarily to the wooden sticks and
the plastic frame instead of to the glass, and so the sides could
be removed from the cages with minimal damage to the webs.
Except when otherwise stated, the webs were always exposed
during the tests to ensure that the predator, P. labiata, could see
the web without obstruction or re£ection from the cage.
A. versicolor juveniles were used to build discoid stabilimenta and
A. versicolor adult females were used to build cruciform stabilimenta with di¡erent numbers of arms (see ¢gure 1). Both vacant
webs (i.e. A. versicolor was not on its web) and webs with
A. versicolor were used. Vacant webs were obtained by removing
the host spiders 24 h before tests began.
We used laboratory cultures of P. labiata that originated from
specimens collected in the habitat of A. versicolor. All P. labiata
used were females (body length, 6^8 mm) that did not have any
Proc. R. Soc. Lond. B (2001) We designed a choice test to determine whether P. labiata could
be attracted to a web by the presence of a stabilimentum. A web
without a stabilimentum (¢gure 1f ) was compared with one with
a stabilimentum (¢gure 1a ^ e), making an e¡ort to pair similarly
sized webs. Each P. labiata was starved for 1 week before the test.
We conducted two series of tests: in one series, P. labiata were
tested using webs with A. versicolor left in them; in the other series,
the same P. labiata were tested using webs without A. versicolor.
esting began by putting P. labiata between two cages that bore
webs. The two webs were set 20 cm apart and P. labiata was placed
10 cm from each web. Each test ended when P. labiata chose one of
the two webs in the vacant-web tests, ate one A. versicolor in the
occupied-web tests or when 10 min had elapsed, whichever came
¢rst. However, observations were continued until the sequence
had terminated if P. labiata responded to a web, even after the
allocated time. All tests were conducted in a plastic box with a
glass bottom. After each test the glass panel was wiped with 70%
ethanol and with distilled water to ensure that all secretions and
pheromones deposited by the previous P. labiata were removed.
We ran all tests under the same light conditions (light intensity,
430.2 Æ 17.40 lux) and a sheet of black cardboard was provided as
a background for each web. We analysed the data using a w2 test of
goodness of ¢t (Zar 1996). (c) E¡ect of A. versicolor on attraction of predators
to webs A. versicolor may be attractive to predators, so the presence of
a spider may a¡ect the attraction of predators to the web. T test
the in£uence of A. versicolor, a naive P. labiata was given a choice
between webs with and without A. versicolor, making an e¡ort to
pair similarly sized webs with the same type of stabilimentum.
esting began by putting a P. labiata between two web-bearing
cages as in the predator-attraction tests. T
ests ended when
P. labiata chose one of the two webs or when 10 min had elapsed,
whichever came ¢rst. Comparisons were made using the w2 test
of goodness of ¢t (Zar 1996). (d) Learning experiments Portia spp. are known to exhibit trial-and-error learning
(Jackson 1992; Jackson & Wilcox 1993). T examine the ability
of P. labiata to detect and remember webs with stabilimenta, we
individually trained P. labiata to associate decorations with foraging success. P. labiata was ¢rst trained to associate the cruciform
stabilimentum with one arm in the web with food, A. versicolor.
Then, a web with a one-arm cruciform stabilimentum was used
in combination with a web with any other type of stabilimentum. The two webs were presented to P. labiata simultaneously, but A. versicolor was only present in the hub of the web
with the one-arm cruciform stabilimentum. In the other web, a
small piece of cotton ball was attached on the hub to replace the
absent prey. P. labiata was starved for 3 days before the experiments. Each P. labiata was allowed to take its time to ¢nd the
web containing the food. Those that went to the web with the
cotton ball were gently removed back to the starting point, and
the training continued until they got onto the web with the Stabilimenta attract predators W. K. Seah and D. Li 1555
Table 1. Results from predator-attraction tests in which one web with stabilimentum and one web without a
stabilimentum were presented to Portia labiata simultaneously in bright conditions (430.2 Æ 17.4 lux). web A web B presence/absence of
A. versicolor number choosing
web A first number choosing
web B first test of goodness of fit present 13 27 χ 2 =4.90; p <0.05 absent 12 28 χ 2 =6.40; p <0.01 present 13 27 χ 2 =4.90; p <0.05 absent 12 28 χ 2 =6.40; p <0.01 present 8 32 χ 2 =14.4; p <0.001 absent 10 30 χ 2 =10.0; p <0.005 present 11 29 χ 2 =8.10; p <0.005 absent 8 32 χ 2 =14.4; p <0.001 present 9 31 χ 2 =12.1; p <0.001 absent 7 33 χ 2 =15.7; p <0.001 one-arm cruciform stabilimentum. The training exercise was
performed eight times, twice for each pair of web types, with
each P. labiata. When P. labiata chose the right web, it was
allowed to feed on A. versicolor for 5 min as a reward. Subsequently, the trained P. labiata were kept without food for 3 days
after the date of learning. They were then used in the learning
experiments to determine whether they could associate a stabilimentum with food. We also used a choice test to examine
whether a predator remembered the familiar stabilimentum. In
these tests, two webs, one with a familiar stabilimentum (onearm cruciform) and the other with an unfamiliar stabilimentum, had A. versicolor present in the centre of the hub. Each
test began by putting a trained P. labiata in between the two
webs, as in the predator-attraction tests. T
ests ended when
P. labiata chose one of the two cages and entered the web or
when 10 min had elapsed, whichever came ¢rst.
3. RESULTS (a) Predator-attraction test When given a choice between webs with and without
stabilimenta, regardless of whether A. versicolor was in the
web or not, P. labiata preferred the webs with stabilimenta
to those without stabilimenta in all tests on all ¢ve combinations of webs (table 1). (b) E¡ect of the presence of A. versicolor on the
attractiveness of webs to predators When given a choice between webs with and without
A. versicolor, P. labiata chose the webs with A. versicolor as
often as they chose the webs without A. versicolor (table 2).
Proc. R. Soc. Lond. B (2001) (c) Learning experiments When P. labiata were trained to associate a one-arm
cruciform stabilimentum with the presence of A. versicolor,
P. labiata chose the web with the one-arm cruciform
(familiar) stabilimentum more often than they chose
other types of decorated webs (table 3). 4. DISCUSSION When a predator (receiver) encounters a spider
(sender) in a web with a conspicuous UV-re£ecting
stabilimentum (visual signal), there are three possible
responses: ¢rst, being trapped if the predator is unable
to determine the presence of the stabilimentum in the
web; second, avoiding the web passively or actively (e.g.
some arthropod predators and birds); and third, being
attracted to the web if it is able to see the stabilimentum.
Surprisingly, almost all studies investigating the adaptive
signi¢cance of stabilimenta as defensive structures have
tried to examine how a predator avoids a web with a
stabilimentum (e.g. Blackledge 1998a ; Blackledge &
Wenzel 1999, 2000). Horton (1980) stated that the web is
a negative stimulus to a bird because sticky silk adheres
to the bird's feathers and clumps them; Horton found
that the cross (cruciform) stabilimentum advertises the
presence of the noxious web, enabling birds to avoid
webs with stabilimenta. Eisner & Nowicki (1983)
measured the durability of 30 webs with stabilimenta
and 30 webs without stabilimenta in the ¢eld, and found
that 8% of the stabilimentum-free webs persisted
undamaged as compared with 60% of the webs with 1556 W. K. Seah and D. Li Stabilimenta attract predators Table 2. Results from tests of the e¡ect of the presence or absence of Argiope versicolor on the attraction of predators
to webs. A naive Portia labiata was given a choice between a web with A. versicolor left in it and a web without
A. versicolor; both webs had with the same type of stabilimentum.
web (B) first test of goodness of fit 14 χ 2 =0.133, not significant 17 13 χ 2 =0.533, not significant 18 12 χ 2 =1.2, not significant 17 13 χ 2 =0.533, not significant 18 web (B), without
A. versicolor number choosing
web (A) first 16 web (A), with
A. versicolor 12 χ 2 =1.2, not significant Table 3. Results from learning experiments in which a Portia labiata trained with a one-arm cruciform
stabilimentum plus food as a reward was given a choice between a web with a familiar stabilimentum (one-arm
cruciform) and a web with an unfamiliar stabilimentum; both webs contained Argiope versicolor.
(Data in the third and fourth columns only are used in these tests. Null hypothesis: if P. labiata chose a web, then the probability
of choosing web A, with a familiar stabilimentum, is equal to the probability of choosing web B, with an unfamiliar
stabilimentum.) number choosing
web B first test of goodness of fit 12 χ 2 =5.16; p <0.05 27 14 χ 2 =4.10; p <0.05 29 11 χ 2 =8.10; p <0.005 30 web B number choosing
web A first
26 web A 9 χ 2 =11.3; p <0.001 stabilimenta. Thus, `web durability was therefore due to
the presence of the visual markers' and `stabilimenta
serve as visual advertisements of webs' (Eisner &
Nowicki 1983, p.186). Blackledge & Wenzel (1999)
conducted experiments to determine the e¡ect of stabilimenta on web damage by birds, and their data support
the hypothesis that stabilimenta can function as a
defence against birds.
In this study, we have demonstrated experimentally
that predators are attracted to webs by the stabilimenta
and feed on the resident spiders: the web-invading spidereating jumping spider P. labiata consistently chose the
webs with stabilimenta more often than the webs without
Proc. R. Soc. Lond. B (2001) stabilimenta (table 1). We also found that the presence of
A. versicolor in the web had no e¡ect on P. labiata's attraction to the stabilimenta (table 2). Our results suggest that
the stabilimenta alone is a su¤cient visual signal to
attract predators to the web. Not surprisingly, stabilimenta acting as highly visual signals to prey and predators create trade-o¡s in signal design between foraging,
avoiding attack by predators and defence when actually
attacked. Like other visual signals that are designed for
communicating with conspeci¢cs that carry a greater risk
of predator attraction (Guilford 1990; Endler 1993; Ryan
& Rand 1993), conspicuous stabilimenta built by the orb
owners to attract visual prey or to warn some visual Stabilimenta attract predators W. K. Seah and D. Li 1557
predators of the presence of noxious sticky silk in the orbwebs may also increase the risk of predation by attracting
particular types of predators.
However, whether or not the silk is conspicuous to
insects is controversial and has important implications
for the function of stabilimenta (in terms of the preyattraction hypothesis versus the predator-defence
hypothesis) (Craig & Bernard 1990; Blackledge 1998a;
Blackledge & Wenzel 1999, 2000). The prey-attraction
hypothesis argues that UV re£ection makes stabilimenta
conspicuous to insects and attracts insects to webs (Craig
& Bernard 1990; Tso 1996, 1998; Watanabe 1999;
Herberstein et al. 2000). However, the predator-defence
hypothesis suggests that insects avoid webs, based upon
the presence of stabilimenta, and that re£ection of UV
light is crucial to make the stabilimenta cryptic to insects,
thereby reducing their ability to avoid webs (Blackledge
1998a ; Blackledge & Wenzel 2000, 2001). There are
several explanations for the incompatible di¡erences
between these two hypotheses. Many of the data relating
to the attraction of prey to stabilimenta are correlational
(Craig & Bernard 1990; Tso 1996, 1998; Watanabe 1999;
Herberstein et al. 2000), while the data supporting insect
avoidance of stabilimenta come from a manipulative
experiment (Blackledge & Wenzel 1999) that controlled
for a strong in£uence of foraging success on the presence
of stabilimenta in webs. Several studies have shown that
spiders capturing more prey are more likely to build
stabilimenta, which can produce a correlation between
the capture of high numbers of prey and the presence of
stabilimenta in webs (Blackledge 1998b ; Herberstein et al.
2000). Furthermore, choice experiments demonstrating
attraction of insects to stabilimenta have been performed
in laboratory settings (Craig & Bernard 1990; Watanabe
1999), while the choice experiment showing the cryptic
nature of the silk to bees was performed under natural
conditions (Blackledge & Wenzel 2000). The background
against which a signal is viewed is crucial to its conspicuousness (Blackledge 1998a). Animals can easily ¢nd a
signal conspicuous in the laboratory when that same
signal is inconspicuous outside. We have demonstrated
that P. labiata are capable of using stabilimenta to locate
A. versicolor prey in the laboratory, but it remains to be
determined how important this is for P. labiata in the
If a web-building spider consistently builds the same
form of stabilimentum in its web, it may increase the risk
of predation because predators may remember the form
of the stabilimentum. In the learning experiments, we
demonstrated that P. labiata can learn to associate the
form of the stabilimentum with food: when given a choice
between a web with a familiar form of stabilimentum and
a web with an unfamiliar form of stabilimentum, P. labiata
chose to enter the web with the familiar form of stabilimentum more often than the web with the unfamiliar
form of stabilimentum.
Building stabilimenta may attract more prey, thus
increasing foraging e¤ciency (Craig & Bernard 1990;
Elgar et al. 1996; Tso 1996, 1998; Watanabe 1999), or
reduce the risk of predation (Robinson & Robinson
1970; Eisner & Nowicki 1983; Blackledge 1998a,b ; Blackledge & Wenzel 1999). T increase their foraging success
by building stabilimenta to attract prey and, at the
Proc. R. Soc. Lond. B (2001) same time, reduce the risk of predation, spiders must
have strategies to balance the trade-o¡. Variation in
stabilimentum-building behaviour seems to be a strategy
that has evolved in these spiders to balance this trade-o¡.
In nature, when Argiope argentata spins a new web each
morning the spider produces a decoration whose pattern
or pattern orientation di¡ers from that spun the previous
day, and on some days spiders spin no decoration at all
(Craig 1994). By varying their web decorations randomly
and daily, A. argentata seem to have evolved a foraging
behaviour that inhibits bees both from learning to avoid
webs and from remembering their location (Craig 1994)
because learning is constrained by anatomical and
physiological limits to the animal's perceptual capabilities
and by the complexity of its neural-processing system
(Dukas 1993; Belisle & Cresswell 1997; Endler & Basolo
1998). Whether, by varying their web decorations
randomly and daily, Argiope spiders also prevent some
predators, such as Portia, from learning to be attracted to
their webs needs further study.
We are grateful to Koh T
eck Hui for helping us collect spiders,
Goh Poh Moi for rearing house£ies and Lim Lek Min for drawing the stabilimenta in tables 1, 2 and 3. Comments and
suggestions from T
odd A. Blackledge, Koh T
ech Hui and an
anonymous reviewer greatly improved the manuscript. This
work was supported by generous grants (R-154-000-060-112
and R-154-000-072-112) to D. Li from the National University
of Singapore Academic Research Fund.
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This note was uploaded on 01/27/2012 for the course ECOLOGY 300 taught by Professor Zumdahli during the Spring '11 term at St. Mary NE.
- Spring '11