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Title:Construction of the orb web in constant and changing abiotic conditions
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Construction of the orb web in constant and
changing abiotic conditions
Joshua Singh Phangurha
A dissertation submitted to the University of Bristol in the accordance with the
requirements for award of the degree of MSc by Research in the Faculty of Science.
School of Biological Sciences
Date of submission: 31/08/2018
Word count: 15,060
Abstract
Orb weaving spiders are thought to alter their web-building in different abiotic conditions to
maximise prey capture success, while being as silk-efficient as possible. However, there is
limited evidence to support this, along with a lack of focus on web variation over time under
constant conditions. It is important to better understand web-building in constant and
changing abitoic environments to determine if spiders perform adaptive manipulations of
their webs. Here, both of these concepts are investigated and the hypothesis that web
construction under constant abiotic conditions will not change over time, due to a lack of
environmental cues for spiders to respond to, is tested. It is also predicted that Zygiella x-
notata will reduce the prey capture aspects in high light intensities, when prey would be more
active, and silk can be preserved. The webs of both species were monitored in a constant
abiotic environment and Z. x-notata web building behaviour was compared in varying
environmental light intensities. These experiments explored patterns of variation in the web
characteristics of Argiope bruennichi and Zygiella x-notata over time in a constant abiotic
environment and identified significant differences in the web building of Z. x-notata in bright
and dark conditions. Results show an overall lack of variation in the webs of A. bruennichi
over time apart from radii number in webs of adult females and upper mesh spacing in
juvenile webs, and no significant variation in the webs of Z. x-notata over time. No
significant differences in the web geometry of Z. x-notata webs occurred in the light intensity
treatment, except for lower mesh height becoming reduced in darker conditions. These
findings suggest that varying abiotic factors, overall, do not influence the adaptive web
building decisions of an orb weaver and could only marginally impact specific web aspects.
Dedication and acknowledgments
I would like to thank Beth Mortimer and Daniel Robert for supervising this project in the Life
Sciences building, Paul Chappell and his colleagues at the chemistry department for building
the frame set up and the British Ecological Society and Bristol Alumni foundation for funding
my travel to Mexico to share this research at the American Arachnological Society’s annual
meeting.
All photographs used in this thesis were taken by the author. Image 1.1 (2.2MB), 1.2 (8.69MB),
1.4 (467KB), 2.1 (5.89MB), 2.2 (6.35MB), 2.3 (6.58MB), 2.4 (7.26MB), 4.1 (337KB), 4.2
(1.11MB), 4.3 (306KB) and 4.4 (435KB) were all edited in Picasa®. Image 1.3 (9MB) was
downloaded from Leica® microscopy software.
Author’s declaration
I declare that the work in this dissertation was carried out in accordance with the
requirements of the University's Regulations and Code of Practice for Research Degree
Programmes and that it has not been submitted for any other academic award. Except where
indicated by specific reference in the text, the work is the candidate's own work. Work done
in collaboration with, or with the assistance of, others, is indicated as such. Any views
expressed in the dissertation are those of the author.
SIGNED: ............................................................. DATE:..........................
Table of contents
Chapter 1 – Introduction and background ………………………………………………................ 1
1.1 – Background on foraging method variation in the natural world ………………………………… 1
1.2 – Biotic influence on web construction ………………………………………………………………………… 2
1.3 – Spider condition ……………………………………………………………………………………………………….. 3
1.4 – Web building influenced by predators …………………………………………………………………………………… 3
1.5 – Wind ……………………………………………………………………………………………………………………………………... 4
1.6 – Light ………………………………………………………………………………………………………………………………………. 5
1.7 – Temperature …………………………………………………………………………………………………………………………. 6
1.8 – Humidity ……………………………………………………………………………………………………………………………….. 7
1.9 – Intraspecific web variation ……………………………………………………………………………………………………. 8
1.10 – Other factors affecting web building ………………………………………………………………………………….. 10
1.11 – Stabilimenta – a prey attractant? ……………………………………………………………………………………….. 11
1.12 – Web damage prevention function of stabilimenta ……………………………………………………………… 12
1.13 – Stabilimenta for predator avoidance ………………………………………………………………………………….. 13
1.14 – Unexplained stabilimenta variation ……………………………………………………………………………………. 14
1.15 – Aim of study ………………………………………………………………………………………………………………………. 14
1.16 – Study species’ ……………………………………………………………………………………………………………………. 15
Chapter 2 – Methodology …………………………………………………………………………………………………. 18
2.1 – Argiope bruennichi and Zygiella x-notata collection ……………………………………………………………... 18
2.2 – Maintenance ………………………………………………………………………………………………………………………… 18
2.3 – Morphometrics ……………………………………………………………………………………………………………………. 19
2.4 – Environmental conditions …………………………………………………………………………………………………….. 21
2.5 – Light intensity experiment (Zygiella x-notata) ………………………………………………………………………. 22
2.6 – Web measurements …………………………………………………………………………………………………………….. 22
2.7 – Statistical methods ………………………………………………………………………………………………………………. 26
Chapter 3 – Results ……………………………………………………………………………………………………………… 26
3.1 – Web building variation under constant abiotic conditions …………………………………………………… 27
3.2 – Juvenile Argiope bruennichi …………………………………………………………………………………………………. 27
3.3 – Adult Argiope bruennichi …………………………………………………………………………………………………….. 34
3.4 – Stabilimenta building ………………………………………………………………………………………………………….. 40
3.5 – Zygiella x-notata …………………………………………………………………………………………………………………. 40
3.6 – Light intensity experiment ………………………………………………………………………………………………….. 47
Chapter 4 – Discussion ……………………………………………………………………………………………………… 49
4.1 – Juvenile upper mesh variation and overall mesh consistency …………………………………………….. 50
4.2 – Spatial constraints ………………………………………………………………………………………………………………. 52
4.3 – Web building experience ……………………………………………………………………………………………………. 54
4.4 – Size limitation hypothesis …………………………………………………………………………………………………… 54
4.5 – Adult Argiope bruennichi radii variation …………………………………………………………………………….. 55
4.6 – Biotic factors more influential on web construction? …………………………………………………………. 56
4.6.1 – Effect of food intake ……………………………………………………………………………………………….. 56
4.6.2 – Prey type …………………………………………………………………………………………………………………. 57
4.6.3 – Prey size …………………………………………………………………………………………………………………. 58
4.6.4 – Hydration ……………………………………………………………………………………………………………….. 58
4.6.5 – Effect of aging ………………………………………………………………………………………………………… 59
4.6.6 – Pollen consumption ………………………………………………………………………………………………… 59
4.6.7 – Effect of reproduction on web construction …………………………………………………………… 59
4.7 – Light intensity experiment (Zygiella x-notata) …………………………………………………………………… 62
4.8 – Hierarchy of variation ……………………………………………………………………………………………………….. 64
4.9 – Lack of stabilimenta ………………………………………………………………………………………………………….. 65
4.10 – Preliminary observations of stabilimenta ……………………………………………………………………….. 66
4.11 – Conclusion ……………………………………………………………………………………………………………………… 67
Appendix ……………………………………………………………………………………………………………………….. 69
References …………………………………………………………………………………………………………………….. 73
Figures, tables and images:
Chapter 1 – Introduction and background
Image 1.1 & 1.2 …………………………………………………………………………………………………………………………. 16
Image 1.3 & 1.4 …………………………………………………………………………………………………………………………. 17
Chapter 2 - Methodology
Figure 2.1 & 2.2 …………………………………………………………………………………………………………………………. 20
Figure 2.3 ………………………………………………………………………………………………………………………………….. 21
Image 2.1 ………………………………………………………………………………………………………………………………….. 23
Image 2.2 & 2.3 ………………………………………………………………………………………………………………………… 24
Image 2.4 …………………………………………………………………………………………………………………………………. 25
Chapter 3 - Results
Table 3.1 & Figure 3.1 ………………………………………………………………………………………………………. 28
Figure 3.2 …………………………………………………………………………………………………………………………. 29
Table 3.3 & 3.3 …………………………………………………………………………………………………………………. 30
Figure 3.3 & 3.4 ………………………………………………………………………………………………………………… 31
Figure 3.5 & 3.6 ………………………………………………………………………………………………………………… 32
Figure 3.7 & 3.8 ………………………………………………………………………………………………………………….33
Table 3.4 .…..…………….………………………………………………………………………………………………………. 34
Figure 3.9 & 3.10 ……….…………………………………………………………………………………………………….. 35
Table 3.5 & 3.6 …………..…………………………………………………………………………………………………….. 36
Figure 3.11 & 3.12 …………………………………………………………………………………………………………….. 37
Figure 3.13 & 3.14 …………………………………………………………………………………………………………….. 38
Figure 3.15 & 3.16 …………………………………………………………………………………………………………….. 39
Table 3.7 ……………………………………………………………………………………………………………………………. 40
Table 3.8 ……….…………………………………………………………………………………………………………………… 41
Table 3.9 & Figure 3.17 ……………………………………………………………………………………………………… 42
Figure 3.18 & 3.19 ……………………………………………………………………………………………………………… 43
Figure 3.20 & 3.21 ……………………………………………………………………………………………………………... 44
Figure 3.22 & 3.23 ………………………………………………………………………………………………………......... 45
Figure 3.24 & 3.25 ………………………………………………………………………………………………………………. 46
Figure 3.26 ………………………………………………………………………………………………………………………….. 47
Table 3.10 …………………………………………………………………………………………………………………………… 48
Figure 3.27 …………………………………………………………………………………………………………………………. 49
Chapter 4 – Discussion
Image 4.1 ……………………………………………………………………………………………………………………………. 60
Image 4.2 & 4.3 …………………………………………………………………………………………………………………… 61
Image 4.4 ……………………………………………………………………………………………………………………………. 62
1
Chapter 1 - Introduction and background 1.1 - Background on foraging method variation in the natural world
Much intrigue has been evoked by the way in which organisms alter foraging/prey capture
techniques in response to environmental change. Changes in environmental abiotic factors such
as light intensity, temperature and humidity can influence foraging effort in many species.
There is limited knowledge on how animals alter their foraging technique/activity in response
to varying light intensity levels. For Coccinellid beetle species, consumption rates and foraging
behaviour vary in response to different visual cues, even though the same prey is hunted1.
Temperature can also have an effect, as shown by South American Gastropods that show
increased drilling predation on bivalve prey in response to higher temperatures existing towards
the equator2. In theory, ectotherms in particular are more likely to show this response owing to
increased metabolic rate at higher temperatures. Temperature can therefore increase/decrease
the activity and conspicuousness of mainly ectothermic prey, which would ultimately influence
predator activity. This has been shown by Eurasian Kestrel predation on Common Lizards in
Norway as the probability of lizards, as opposed other endothermic prey, being delivered by
kestrels to their nests tends to increase as temperatures rise towards midday when lizard activity
is greater3. Recent literature has also shows that the activity of an important North American
predator, the Western Rat Snake, is increased by another abiotic factor, namely humidity4. The
increased activity of snakes under higher humidity leads to higher predation rates of mammals
and birds. Humidity has also been shown to affect foraging activity in ants in similar fashion5.
As activity levels of a predator change in response to these environmental factors, the exposure
to danger, such as risk of predation to the predator itself, will fluctuate. The exposure to such
dangers can cause evolutionary, physiological and/or ecological trade-offs to maximise
foraging success, while being as resource efficient and undetectable as possible6.
Spiders, the focal animals in this study, are an ectothermic group of organisms where activity
and predation rate may be influenced by abiotic factors. Aerial spider webs come in many forms
and primarily function as a trap for mostly insect prey and a vibrational device to communicate
with conspecifics7. This makes spiders ideal study organisms in regard to the impact abiotic
factors have on a predator as their webs are direct indicators of their foraging effort. The orb
web is the most familiar of these traps as a two-dimensional, circular meshed structure. Sticky
2
spirals are fastened onto radial threads that converge to the central hub of the web and an array
of silk types are used8, 9. The evolution of the orb web is not clear and there is controversy over
a monophyletic or polyphyletic origin10, 11. It is thought that Araneomorph spiders (Arthropoda:
Arachnida) alter their orb webs in a variety of forms to respond to pressures in the environment,
such as climate, vegetation structure, prey preference, prey abundance, predator presence and
parasitic infection12. This suggests that this web type is highly advanced in its functional
plasticity.
1.2 Biotic influence on orb web construction
There has been interest in how spiders change the structure of their webs in response to varying
abiotic and biotic factors to maximise foraging and reproductive success, while being as
silkefficient as possible13. The biotic influence on web building has been shown in Araneus
diadematus, which has been observed to increase the symmetry and capture area of its web in
agricultural areas when compared to spiders of the same species in a more prey abundant
habitat14. Thus is provided a more efficient structure for trapping prey in agricultural land where
insect abundance is much lower due to the use of pesticides. This suggests that A. diadematus
is willing to invest more energy into producing more silk to increase the chance of catching
insects in order to feed sufficiently. A. diadematus can even modify the fine details of its web
that are associated with prey capture, as this species was also observed increasing mesh spacing
when exposed to larger prey items (small Drosophila to larger mosquito prey)15. Larger mesh
height would presumably facilitate the trapping of larger prey rather than smaller and a larger
web can be constructed with the same/less silk investment required to build a smaller, more
densely woven web16, 17. However, prey taxa must be taken into consideration when evaluating
the effectiveness of larger mesh spacing on capturing larger prey items as deer flies (Chrysops
sp.) are retained for less time in the orb webs of Argiope with large mesh spacing, whereas
hanging flies (Hylobitacus sp.), which differ greatly in their morphology, show no difference
in their retention times in webs with smaller or larger mesh18. This effect of prey availability
on web construction may provide an explanation as to why Larinoides cornutus was observed
in vitro building smaller, rounder orb webs with shorter spiral lengths and fewer, thinner radial
threads than spiders in the field19. Reduced silk investment in a controlled environment may be
linked to prey abundance and spider condition, as prey tends to be more abundant in the field
3
and impacts the web more frequently19. In the wild, spiders would therefore need to build more
and thicker radii to sufficiently dissipate the kinetic energy of prey impact, also making the
web more robust to damage by abiotic factors, such as wind. Reducing the frequency of prey
available to orb weavers has also been shown to result in an increase in the overall web area,
perhaps as an adaptation to increase the chance of catching prey when it is less abundant20.
1.3 - Spider condition
The physical condition of the spider can also be influential on web building. Spiders close to
the end of their life expectancy naturally have reduced health, which causes a decline in
mobility and coordination when building webs, resulting in an ‘untidy’ arrangement of spiral
turns between radii21. These older spiders can also exhibit reduced silk investment, particularly
in flagelliform spiral silk that is associated with prey capture, which would hinder foraging
success. If orb weaving spiders are missing two or more legs, it does not appear to impact their
web building and foraging success significantly. However, if they are missing three or more
legs, this can result in an increased asymmetry in their orb webs22, which may negatively affect
predatory success. Spider mass has previously been shown to influence web building
behaviour, an example being orb weavers which tend to reduce silk investment in web building
as they mature (and increase in mass) thus to preserve energy for reproductive processes23, 24.
However, later studies concluded that although heavier spiders reduce their silk investment in
the orb web, more energy is required to build it25. This is potentially due to the increased impact
of gravity on mobility in larger spiders over a larger area, even if silk investment is reduced.
1.4 - Web building influenced by predators
One of the main influences on foraging behaviour in the natural world is predation. Foraging
effort can be reduced in the presence of a predator to reduce detectability. This appears to apply
to spiders and their web building behaviour. For example, Argiope versicolor is known to
reduce the total area of its orb web and to increase mesh spacing in the presence of a visually
acute predator26, which may reduce visibility of the web. A smaller web would enable the spider
to reach captured prey with less movement, which would reduce the chance of attracting the
4
attention of a predator27. Smaller webs perhaps save energy for other behaviours, such as
moving between both sides of the orb web and web shaking to blur the spider’s appearance28.
However, reduced web area would result in a smaller capture area, negatively impacting prey
capture rate. So it seems that, at least in some orb weaving species, there is a trade-off between
foraging success and conspicuousness to predators. Also, barrier webs, three-dimensional
tangled silken structures positioned in front of both sides of the orb web, are frequently built
by Nephila species. These structures are thought to protect the spider from predatory attacks,
with smaller, more vulnerable spiders building denser barrier webs compared to larger sexually
mature adults29.
Infection by parasitoid wasps, which are frequent predators of arachnids, can alter an orb
weaver’s web building for self-benefit. Spiders infected with parasitoid wasp larvae have
previously been observed reducing the number of sticky spirals spun during web building30.
This may benefit the parasite during pupation by reducing the chance of prey being captured
and damaging the web, which suspends the wasp cocoon out of the reach of predators.
Preventing the spider from investing in flagelliform spiral silk, which accounts for the majority
of total web mass, may maintain spider weight to improve nourishment of the parasite31.
1.5 - Wind
The abiotic influence of the environment on web building occurs less frequently in the
literature, but the few studies that are present have produced findings of interest. One abiotic
factor that has been of much focus, is the effect of wind on web architecture. It has been shown
that orb weavers can adjust the properties of their silk, as well at the geometry of the web in
response to wind exposure. This has been shown in two Cyclosa species from Taiwan, where
the seashore-dwelling C. mulmeinensis appears to build fewer radial threads, larger mesh
spacing and smaller catchment area than the interior forest-dwelling C. ginnaga32. This study
suggests that the building of smaller, less dense webs by C. mulmeinensis is an adaptation to
reduce drag on the web in a windy coastal environment, thus preventing web damage. Larger
mesh spacing is also considered to be more efficient in the catching of larger insects, which are
able to fly in windier conditions, as less silk is used. The major ampullate (MA) silk in the webs
5
of C. mulmeinensis, a silk type that has been of much engineering interest due to its
extraordinary tensile strength33, was shown to be significantly stronger than the MA silk of C.
ginnaga. This is thought to be another adaptation to windy conditions, by perhaps investing
more energy into making stronger radii than producing more radii. C. ginnaga built larger,
more densely woven webs in its forest habitat where wind speed is lower and more constant,
suggesting that this species may be able to invest more energy into building a more formidable
trap in this habitat. Araneus diadematus behaves in a similar manner, showing reduced radii
and capture area, but increased mesh spacing and reduced mesh number when exposed to windy
conditions34. This study additionally tested the impact of wind on the eccentricity of webs and
found that when weights were applied to wind exposed webs that were orientated horizontally,
the web deformed significantly less from the horizontal plane, compared to control webs that
were not exposed to windy conditions. Building a web like this would hinder prey capture
capabilities, but would limit damage in a windy environment. This suggests that it may be more
energy costly to rebuild a damaged web than to consume fewer prey items. Furthermore, more
recent research has indicated that orb weaving spiders increase the volume of glue droplets on
the capture spirals of their webs, while positioning these droplets closer together, to prevent
desiccation in windy conditions35. The glue droplets constitute an important web component
that is essential for prey capture and if these glue droplets dried to too great a degree, the web
would lose its stickiness.
1.6 - Light
There have been previous studies focusing on the impact of light intensity on web building
behaviour. Neoscona crucifera is known to significantly reduce the average radius of its web
after choosing to position the web near the brightest of four artificial lights in the field. Prey
was most abundant at the brightest light, suggesting that the spiders chose to build webs in the
most resourceful area where they could reduce silk investment owing to the increased chance
of catching prey36. Similar results occurred from a study on Larinoides cornutus, which also
showed supporting evidence that adult females are genetically predetermined to select brightly
lit web building sites as both wild and lab reared individuals exhibited the same behaviour in
this respect37. Later studies found a similar relationship between light intensity and web
building, but unlike N. crucifera, Argiope keyserlyngi showed a significant increase in overall
6
web area when exposed to higher environmental light intensity in the lab38. Silk also reflects
light intensely39, rendering the orb web more visible to a wide range of predators in bright
conditions40. Orb weavers spend much of their time motionless in a retreat or at the hub of the
web41, however they are most vulnerable to predation when highly mobile during web
building42. It is assumed that the duration of mobility when building a smaller web in brighter
conditions would be shorter, reducing the chance of being detected.
Furthermore, spiders have been documented as selecting brighter, open areas over sheltered
areas in which to build their webs in the field and, in a sheltered habitat, will even position their
webs to face an open space43. This makes ecological sense, as flying insects tend to be more
frequent in bright, open areas, such as the edge of woodland, where prey interception by the
web is more likely44. However, other studies have indicated that frequently building webs in
this type of environment is a genetically derived character45.
1.7 - Temperature
Although the geometric influence of temperature on webs has been poorly studied, the few
studies that have been carried out have generated considerable results on this topic.
Temperature has been shown to impact the geometry of orb webs, as spiders have been
observed increasing web area and mesh spacing when exposed to cooler conditions from a
warmer control condition34. This enables a larger web to be built, while investing less energy
in producing flagelliform spiral silk. However, as spiders are ectothermic, cooler temperature
may just simply hinder levels of activity, causing the spider to produce less silk.
Other studies have shown that temperature can affect web building site selection. When given
the choice of multiple environmental temperatures, the sheet web-building Theridiid
Archaearanea tepidoriorum selected areas to build webs where the ambient temperature was
approximately 20°C46. This study subsequently found that the spiders were able to build the
heaviest webs when exposed to this temperature and that web mass increased with strand
density. Increased strand density is known to positively correlate with prey capture success47,
7
suggesting that spiders can select environments to build their webs where foraging is most
lucrative.
Furthermore, the physical properties of silk can be affected by changes in temperature as the
MA silk of Nephila edulis is known from lab experiments to significantly increase in tensile
strength at very low temperatures (-60 to 0°C) and then drastically decrease at very high
temperatures (~70°C)48. However, it is highly unlikely that spider silk in nature would be
exposed to these extreme temperatures.
1.8 - Humidity
Although humidity has not been previously shown to clearly affect web geometry, evidently, it
can impact the physical properties of flagelliform spiral silk, which has a primarily prey capture
function. For example, the viscous glue droplets within flagelliform silk are known to absorb
more water in humid environments, causing the glue droplets to expand, increasing surface area
contact with prey and amount of air drag, which acts to dissipate prey impact emergy49, 50. This
then led to studies focusing on the effectiveness of stickier webs in humid environments. In one
such study, house flies placed in the webs of Araneus marmoreus constructed in 55% relative
humidity (RH) were retained eleven seconds longer than in webs built in 37% RH51.
MA silk has been observed supercontracting52. This is where, in humid conditions, water causes
MA thread length to shrink up to 50% and increase in volume. This silk type can supercontract
at approximately 70% RH at 20°C, causing the web to increase in tension53. This tautening of
the web via supercontraction has been considered an adaptive function to limit unwanted web
deformation under the weight of liquid water or following prey damage54. Wet MA silk in the
webs of Argiope spiders under high humidity has been shown to reduce in stiffness and extend
40% more than dry webs when prey impacts the web55. This may make sense in regard to prey
capture as significantly more web deformation, in response to prey impact, could reduce
deceleration, which would ultimately reduce web damage as the web would need to cope with
lower peak forces. More recent research suggests that supercontraction of MA silk may have a
8
sensory function, as well as a mechanical function. Constrained supercontraction of MA silk
generates a stress of approximately 50MPa on these threads, which increases the speed and
amplitude of transverse waves that are important for detecting prey and conspecifics,
compensating for the ‘sagging’ deformation of the web under the weight of additional water56.
In theory, this would also limit the strain that major ampullate threads must withstand from
kinetic energy produced by prey impacting the web57. Before web building even takes place,
>70% RH is the key environmental trigger for web building in Pasilobus spiders, which is
perhaps an adaptation for build in conditions when the stickiness and capture performance of
the web is maximised58.
1.9 - Intraspecific web variation
When studying the impact of the environment on web building behaviour, it is important to
consider the fact that web variation can occur among individuals of the same species. This is
particularly evident at different ontogenetic stages. For example, juvenile Clitaetra irenae are
known to build rounder, more typical orb webs, but as they mature their webs become
increasingly elongated vertically to form a ‘ladder’ web59. Furthermore, ontogenetic web
changes have been shown in Uloborus spiderlings, which immediately leave the egg sac after
hatching to build an orb web. These early webs differ from adult webs in that they contain more
radii and spiderlings only build the auxiliary spirals, which are used to guide capture spiral
construction60, with a lack of genuine capture spirals61. Uloborus spins cribellate silk, which
requires a fully formed cribellum and calamistrum. Spiderlings only obtain these cribellate silk
spinning organs after their second moult, which may explain the lack of spirals62. Hub
symmetry can also be influenced by ontogeny and this is most apparent in larger, mature
spiders, which tend to orientate the hub near the top of the web, exposing a larger catchment
area in the lower web portion where prey is more likely to be captured. It has been hypothesised
that this is an adaptation to enable heavier spiders to move with minimum energy expenditure
in a downwards direction with gravity, rather than against it63. Moreover, ontogeny can
influence the structural properties of webs. Neoscona arabesca was observed investing more
in the size of glue droplets on capture spirals as the spiders grew over time, which improves
the performance of catching and retaining larger prey64. This could be due to a requirement of
9
much larger prey during the later stages of a spider’s ontogeny to ensure successful
reproduction65.
In many orb weavers, the central nervous system (CNS) of newly hatched spiderlings only
contains neuroblasts and is still not fully developed64. This may explain why the first few webs
that are built by young spiders of some species are asymmetric and become increasingly
symmetric over several subsequently built webs while the CNS develops. The variability in the
webs of 1st instar Nephilengys cruentata is also much higher than in the webs built later in their
ontogeny66. Further details of the plasticity of web architecture by juvenile spiders has been
shown by Nephila madagascarienesis, at instars one and two, which construct multiple types
of radii, which are defined by their division and different attachment points to the periphery of
the web67. The use of these radii types was variable between individual spiders.
In some species, such as Wixia abdominalis, intraspecific web variation occurs among adults
as the construction of a missing sector, with a signal thread running through this sector, is
optional and can vary from spider to spider. However, this species tends to build more
homogenous mesh spacing in the lower portion of missing sector webs, which is thought to
improve prey retention68. This extended retention time enables the spider to reach prey before
it escapes while travelling further distances across the web from a retreat at the periphery. This
study further showed intraspecific variation in the presence of the sector, as some adults built
complete webs with no sector where the spider occupied the hub. These webs had more
heterogeneous mesh spacing, indicating that retention time is not prioritised by the spider as
the distance to prey is much shorter. Flexibility in the use of a missing sector and sector
orientation is also known to occur in other genera, such as Zygiella69. Furthermore, Araneus
diadematus has been observed building ‘pilot webs’ when introduced to new web building
sites. These initial pilot webs are usually smaller, less planar and contain fewer sticky spirals
that subsequent webs, which become larger, are more planar and possess stickier spirals70. From
these findings, it was suggested that A. diadematus build low investment pilot webs to assess
capture success and web damage at a new site, before investing more energy in ‘proper’
subsequent webs and that the anchor points from the pilot web are used as reference points
when constructing a new web. Many experiments that investigate web building behaviour
10
involve purposely destroying webs to ensure that a sample spider builds a new one. However,
orb weavers of the same species that are left undisturbed to ingest their entire webs when
rebuilding tend to build subsequent webs with a larger area, indicating that the amount of web
remnants that can be ingested when rebuilding is energetically important for spiders and can
cause intraspecific and intra-individual web geometry variation71. As with all species, there are
morphological differences from one individual to another. In spiders, some individuals will
naturally be larger and heavier with greater leg spans than other individuals as fully grown
adults of the same species. This automatically causes variations in web area for example, as the
area of a web tends to increase with spider size72.
1.10 - Other factors affecting web building
Web building experience, rather than developmental ontogeny of the CNS, can also be
influential on the symmetry of orb webs. One study showed that rearing spiders in small boxes,
where webs could not be built, before allowing them to build webs in larger spaces impacts
their ability to build an efficient prey capture web design, as spiders reared in larger spaces,
where they could build webs for a longer duration, built more asymmetric webs. The study also
found that when prey was placed in the lower section of the web over a 6-day period, the
experienced spiders augmented the area of the lower web portion, presumably in anticipation
of prey being caught here73. In contrast, Zygiella x-notata that were reared in small boxes
(where they could not build a web) for an extended period built more asymmetric webs when
entered into a larger web building space. These asymmetric webs had longer lower web sections
than the webs of the control spiders that were immediately entered into a web-building frame
after collection74. These findings may indicate that web symmetry could have an element of
species-dependant variability.
Web geometry can also fluctuate with web repair. Web repair, according to recent literature,
seems to be triggered by damage of anchor threads. Breakage of these structurally supportive
threads causes a great loss of capture area, which seems to trigger the quickest web repair
response by Araneus diadematus75. The repair response to anchor thread damage was variable
in this study, as some spiders built a new anchor thread, while others attached the broken thread
to existing anchor threads to increase web tension.
11
1.11 - Stabilimenta – a prey attractant?
In addition to the orb web, some species construct stabilimenta/web decorations in the central
vicinity of the web. The function of stabilimenta is thoroughly debated with several proposed
theories. It was originally thought that stabilimenta serves as additional support for the web,
hence the name of the structure76, 77. However, since then a multitude of theories have been
proposed. These include advertisement to larger animals to prevent web destruction78,
camouflage79, the provision of a ‘sunshield’ for thermoregulatory purposes80 and UV
reflectance to attract insects towards the web81, 82.
The UV reflectance theory has received much attention. Argiope spiders have been shown to
reduce the area of stabilimenta under high light intensity and increase stabilimenta area in low
light intensities83. This may support the UV reflectance theory, as the spider can afford to invest
less in stabilimenta owing to high UV reflectance of the structure and visibility to insects in
bright conditions. Insect activity also tends to be higher in bright conditions as temperature
usually positively correlates. This would increase the chance of prey capture and reduce the
necessity of an extra prey attractant. A later field study on this topic showed supporting results
for the theory. Prey capture in the webs of Argiope bruennichi with stabilimenta and webs
without stabilimenta were compared, and it was found that decorated webs captured 331 prey
items, while undecorated webs captured 119 (total of 450 prey items between both webs) during
five hour trials per day over 10 days. This study also found that webs with stabilimenta captured
more than twice the amount of large prey (>5mm) than webs without stabilimenta, which
indicates that stabilimenta may be more effective in catching larger prey84. Early and recent
studies have shown coherent results to support this prey luring function of stabilimenta via UV
reflection85, 86, but recent studies have also found no difference in prey capture between
decorated and undecorated webs87.
The morphology of web decorations differs between and within Argiope species. Specific
stabilimenta shapes are considered more attractive to UV sensitive insects, such as certain bee
species, which seem to approach cruciate stabilimenta much faster than linear stabilimenta88.
It has also been suggested that there is a trade-off between constructing decorations to attract
12
prey and reducing detectability by visually acute predators that may be attracted to the
structure89.
However, not all studies are coherent with the UV reflectance theory. Orthopterans, which are
the main prey for Argiope bruennichi in their grassland habitat, do not respond to stabilimenta
but flying insects, such as Hymenopterans and Dipterans, are more likely to respond due to
their sensitivity to UV90, 91.
1.12 - Web damage prevention function of stabilimenta
Another theory that is often mentioned in the literature, but has little supporting evidence, is
the web advertisement theory. This theory suggests that some orb weaving spiders adorn their
webs with a stabilimentum to signal the web to larger vertebrates, such as birds, to prevent web
damage92. Earlier studies monitored Argiope webs in the field, testing the effect of adding
artificial paper decorations to undecorated webs on the overall web damage at the end of each
day. This test resulted in decorated webs remaining more intact than undecorated webs, which
may have reflected on the benefit of the signalling function78. However, this field study did not
take into consideration the damage caused by prey capture and other environmental
biotic/abiotic factors. It is possible that insects perceived the paper decorations as different from
silken structures and avoided the webs more often, thus reducing web damage.
However, a later field study found that 16.4% of Argiope appensa on the island of Guam, where
native bird species have been eradicated due to predation by the invasive Brown Tree Snake,
adorned their webs with stabilimenta. This was then compared to A. appensa on three
neighbouring islands where the native avian fauna remains intact. 41.9-56.9% of spiders on
these neighbouring islands constructed stabilimenta, perhaps to advertise their webs in an
environment with a higher density of bird species93. If web decorations have an antipredator
function (see section 1.14), then these results may reflect the lack of predators on Guam.
13
Experimentally damaging the web has recently been shown to encourage Argiope to increase
the area and conspicuousness of web decorations over time94. These results have been
associated with spiders learning that their environment causes a lot of web damage that is not
caused by prey, but perhaps larger vertebrates, triggering this web building response to
advertise the web more rigorously. However, these findings may be supportive of the
stabilizing function, as spiders may have been enhancing stabilimenta in response to frequent
web damage to strengthen the web in anticipation of damaging factors occurring again. Having
said that, a mechanical function of the stabilimentum is unlikely as it tends to be placed loosely
over web.
1.13 - Stabilimenta for predator avoidance
Antipredator functions have been applied to stabilimenta. Certain stabilimenta morphologies
may serve different antipredator defences, as cruciate web decorations, which are built by
certain Argiope species, are thought to make the spider seem larger to exceed the gape size of
predators as the spider’s legs are often aligned with the ‘arms’ of the cross shaped structure95.
Linear stabilimenta also appear to deter predators, as Blue Jays are known to, when given the
choice, attack spiders in non-decorated webs more frequently that spiders in decorated webs96.
However, it is difficult to interpret exactly what is it about the stabilimenta that deterred the
birds. Juvenile Argiope tend to build a disc shaped stabilimentum, which has been associated
with protection from predators. Juvenile Argiope versicolor move more frequently between
both sides of the discoid stabilimentum in the presence of a predator, potentially using the
structure as ‘shield’ for protection from attack. This was compared to the response of adult
females of the same species, which do not always build decorations. Adult females seem more
likely to drop from undecorated webs than from decorated webs in response to a predator,
potentially supporting the antipredator function97. As web decorations are optionally
constructed, there could be a cost-benefit scenario in response to the environment that
determines stabilimenta construction.
14
1.14 - Unexplained stabilimenta variation
In addition to these contradictory findings, there has been a focus on stabilimenta construction
in a constant environment. Decoration construction in Argiope mascordi can be highly variable
in a constant environment, alternating between linear, circular and cruciate shapes98.
Construction of web decorations has also been associated with individual spiders that have
consumed more prey biomass and are well nourished, as they can afford to invest in more silk99.
These findings suggest that stabilimenta may not have an adaptive function, but that
construction is an intrinsic behaviour.
1.15 - Aim of study
In this study, the effect of contrasting environmental light intensities on Zygiella x-notata web
construction will be investigated. It is hypothesised that Z. x-notata will use light intensity as a
proxy for anticipating levels of prey activity. Insects tend to be more active in high light
intensities100, 101 and it is predicted that Z. x-notata will reduce the prey capture aspects (capture
area, radii number and mesh spacing) of the web in high light intensities. This strategy would
be silk-efficient, as less silk is used to build a smaller web while the probability of catching
prey in a brighter environment is higher. This strategy would also reduce visibility to
predators38. These prey capture aspects are expected to become enhanced, with the formation
of an overall larger web, in dark conditions thus to increase the chance of catching prey when
it would be less active. Fine detail changes in orbs web in response to light intensity are rarely
studied in the literature and there is also a lack of focus on this topic in a controlled environment
where external factors that may affect web building are limited. This study intends to solely
focus on the effect of the abiotic environment on web building.
The extent of web building variation in Zygiella x-notata and Argiope bruennichi adults and
juveniles under constant abiotic conditions will also be investigated. It is predicted that there
will be no significant variation in any of the web characteristics in Z. x-notata and A.
bruennichi, as the abiotic environment will remain the same with no changes in environmental
cues that could be detected by the spiders. This is the first in depth study investigating web
building variation in the fine aspects of the orb web, other than stabilimenta, under constant
15
abiotic conditions. This study aims to improve the current limited understanding of the degree
of web building variation in a constant environment, providing insights into the extent of
influence that abiotic factors have on web building or whether there is any influence at all. This
will fill significant gaps in the current literature on this topic.
1.16 - Study species’
Argiope bruennichi is a large, strikingly marked orb weaving spider that originates from the
Mediterranean region. The species’ rapid expansion into cooler northerly climates since the
20th century has led to colonisation of the United Kingdom, with the first record occurring in
1922 in Rye, East Sussex102. It is thought that the spider arrived in the UK via ballooning, where
spiderlings deliberately produce a thread of silk to catch on wind currents that can then cause
the animals to drift, while airborne, up to hundreds of kilometres103. There is also the possibility
of introduction via artificial transport, such as cargo ships. Wasp Spiders in the UK mainly
occur throughout Southern England in habitats such as meadows, grassland and woodland edge
where they mainly catch Hymenopterans, Dipterans and Orthopterans via the use of their large
elaborate orb webs104. Maturity occurs in mid-summer for both males (44.5mm body length)
and females (11-15mm body length), but adult females are also present throughout autumn
unlike the males that die during mating, due to female cannibalism, or soon after mating105.
Zygiella x-notata builds an orb web with, among British spiders, a distinctive missing sector106.
A signal thread is constructed through this missing sector and leads to the spider’s retreat, where
it stays in contact with one of the spider’s front legs107. The thread is used to detect vibrations
of prey that are caught in the web, while the spider remains concealed108. Z. x-notata is often
found in urban environments on artificial structures, such as window frames and fences where
they tend to construct new webs daily. Mature males tend to live from late summer to autumn,
whereas females can be seen all year round109.
16
Image 1.2 – The orb web of Argiope bruennichi with vertical stabilimentum.
Image 1.1 – Adult female Argiope bruennichi..
17
Image 1.4 – The orb web of Zygiella x-notata with distinctive, but not always present, missing sector.
Image 1.3 – Adult female Zygiella x-notata.
18
Chapter 2 – Methodology
2.1 - Argiope bruennichi and Zygiella x-notata collection
Juvenile A. bruennichi were collected from April to June 2017 at Monks Brook Meadows in
Southampton, UK (Grid reference: 50°56’49.3”N°122’16.4”W / 50.947033, -1.371220). All
target individuals found while walking in suitable habitat were collected from their orb webs,
which were constructed in low, dense vegetation. Collecting all individuals provided a
representative samples size of juveniles and adults. Only adult females were collected between
July and August. Mature males tend to not build webs at this life stage, as they are usually
found near or within the webs of females. The habitat at this location is semi-improved
mesotrophic grassland110.
Adult female Zygiella x-notata were collected from metal railings around the outside of the
Life Sciences building in Bristol, UK. (51°27’31.4”N2°36’029”W / 51.458707, -2.600794) and
back gardens in Bristol (51°30'12.2"N 2°35'56.5"W / 51.503349, -2.598969) and Southampton
(50°56'17.4"N 1°22'46.2"W / 50.938167, -1.379500). Z. x-notata were collected from their
webs at night as this species tends to be nocturnal and can be found in the hub of the web in
darkness. Additional specimens were collected during daylight by use of an electronic
toothbrush to lure spiders out from their retreats. Collection took place from February-
September 2017.
2.2 - Maintenance
Juveniles were kept in small transparent Perspex® 7cm x 7cm x 3cm frames which were custom
built (Figure 2.2). One side of the frames were enclosed with Perspex® covered in petroleum
gel and the other side was closed off with Parafilm® covered in petroleum gel. The Parafilm®
had holes pierced through it to allow ventilation within the enclosed space and exposure to the
ambient environmental conditions. Adults were kept in larger 30cm x 30cm x 4cm transparent
Perspex® frames divided by 35cm x 35cm PVC sheets covered in petroleum gel (Figure 2.1).
19
All spiders were kept in the laboratory for seven days, enabling them to build three
acclimatisation webs before being used for data collection. Spiders that did not build three webs
within this time period were released back into the wild. The petroleum gel was used to prevent
spiders attaching silk threads to the dividers so that webs were not destroyed when taking
frames out of the set up. This arrangement encourages the spiders to build an orb web perfectly
within the frame111.
2.3 – Morphometrics
A Leica M205c microscope was used with the Leica application suite software to digitally
measure the cephalothorax of each spider. Each spider was restrained in a small, flat petri dish
while a photograph of each individual’s cephalothorax was taken together with a scale bar for
future reference. Cephalothorax width was measured in millimetres and rounded to one decimal
place (e.g. 3.84mm rounded to 3.8mm). Cephalothorax measurements were taken as not only
does this body feature remain consistent in size until the spider moults, but it is this
measurement that is also considered reliable when predicting web dimensions in orb weavers
at all life stages112. The live mass of each spider was measured in milligrams to two decimal
places. The live mass of spiders was measured because, as well as cephalothorax width, this
has also been shown to influence aspects of orb web construction, such mesh spacing112, and
potentially other orb web aspects. These measurements were repeated each time the spider
moulted owing to significant increase in overall size, which is likely to affect web architecture.
20
Figure 2.1 – A side view of the frame set up for adult spiders based on Zschokke & Herberstein (2005)113.
Dividers were translucent and spiders were exposed to light coming from the bulbs on the sides of the chamber,
which could penetrate the transparent sides of the Perspex® frames (labelled ‘frame’). Heavy weighted objects
were placed at both ends of the set-up, holding the frames together tightly to prevent escapes. There were small
gaps between dividers and frames that enable ventilation, but were not large enough for spiders to fit through.
Figure 2.2 – Individual frame for adult spiders outside of the divider set up. Spiders built webs within the open
space of the frames.
Divider Divider Divider Divider Divider Divider
30cm
4 cm
30 cm
Frame Frame Frame Frame
5 cm
21
Figure 2.3 – Individual frame for juvenile A. bruennichi. No dividers were necessary due to Parafilm®
on one
side of the frame and fixed Perspex®
enclosing the other side.
Juveniles were fed one Drosophila fruit fly and adults were fed one Diptera species of 10-
15mm body length per newly constructed web. Adult webs were sprayed with water after
photographs of webs were taken in order to hydrate the spiders. For juveniles, a pipette was
used to place drops of water on the hub (centre) of the web, which the juveniles could drink
from.
2.4 - Environmental conditions
Spiders were kept within their frame set up in two environmental chambers (Micro
ClimaSeriesTM Premium Ich Insect Chamber). In these chambers, the abiotic environment could
be controlled. In this chamber, day time conditions were set at 20°C and 40% relative humidity
(RH). The juveniles were exposed to a light intensity of 137.5µmol m-2 (195mm from light
source). Adults were exposed to 54.8µmol m-2 from the left side of the frames (510mm from
light source) and 137.47µmol m-2 from the right side of the frames (180mm from light source).
Night time conditions were 16°C, 40% RH and total darkness. Spiders were exposed to a 15
hour light/ 9 hour darkness cycle to mimic UK summer time daylight hours and to not interfere
3 cm
Parafilm screen
7 cm
Ventilation holes
22
with the circadian clocks of these organisms. Such interference could have an effect on foraging
behaviour and ultimately web building in orb weaving spiders114. The webs built by 17
individual A. bruennichi (10 juveniles and 7 adults) and 24 Zygiella x-notata were exposed to
these conditions, which were kept constant in order to investigate the degree of web building
under constant abiotic conditions.
2.5 - Light intensity experiment (Zygiella x-notata)
In order to investigate the effect of lowering light intensity on orb web building, 14 Zygiella x-
notata individuals were exposed to two different environmental light intensities for two weeks
each. The day time conditions in the control cabinet (bright condition) consisted of 157.3µmol
m-² light intensity, 20°C and 40% RH. Night conditions consisted of 16°C, 40% RH and total
darkness. Spiders were exposed to a 15h light/ 9h darkness cycle. The conditions in the
experimental cabinet (dark condition) were the same, except that the day time light intensity
was significantly lower at 29.6µmol m-². Webs built by each spider within a two-week period
in each condition were measured and compared.
2.6 - Web measurements
All webs built by spiders in both environmental conditions were photographed against a black
paper background. Bright lights were aimed at the webs from behind in order to illuminate the
silk for easier photography and clearer images. The webs were photographed adjacent to a ruler
within the plane of the web in order to set a scale on the software ImageJ, where the web aspects
of interest could be measured accurately. To ensure that A. bruennichi individuals were
rebuilding their webs each time they were sampled, the webs were dusted with crushed pollen
granules after the photographs were taken. If the yellow pollen dust had disappeared the next
day and a web was present, it was assumed that this was a new web as spiders take down their
webs before building a new one. The new web could also be compared to the image of the
previous web to ensure that it was definitely different and newly built. This dusting method
was used as a means of avoiding web destruction, as previous research suggests that web
23
damage affects stabilimenta morphology115. The webs of Z. x-notata were destroyed after
photographs were taken and the spiders were fed and watered. Only the anchor, bridge and
frame threads were left intact to promote subsequent web building (Image 2.1).
Image 2.1 – Diagram to show location of bridge (yellow arrows), frame (red arrow) and anchor (blue arrow)
threads outer edges of the orb web. These were left intact, while all other threads were destroyed after each web
was sampled.
The web aspects of interest were radii number, centre area (mm2) (hub area and free zone area
combined), catchment area (mm2) (area covered by flagelliform spirals), average upper mesh
spacing (mm), lower mesh spacing (mm), right mesh spacing (mm), left mesh spacing (mm),
upper and lower linear stabilimentum area (mm2) and stabilimentum length (mm) (Image 2.3).
Mesh spacing was measured by dividing the length of the radial thread between in the inner-
most spiral to the outer-most spiral (upper, lower, right or left portions of the web) by the
number of flagelliform spirals that pass through this length measurement116. The radial thread
measured for mesh spacing in each web portion was chosen as the most vertical radial thread
in the upper and lower portions of the web (upper and lower mesh spacing) and the most
horizontal radial thread in the right and left portions of the web (right and left mesh spacing).
For discoid stabilimenta that were built by some juveniles, only area was measured due to the
irregular shape of the structure and difficulty in identifying an accurate diameter. Linear
24
stabilimenta lengths were measured from the upper-most points of the upper and lower
stabilimenta to the lower most point.
Image 2.2 – Photographs of juvenile discoid stabilimentum (left) and adult vertical linear stabilimentum (right).
Location of the lower and upper portions of linear stabilimenta in the webs of Argiope bruennichi labelled in
right image. Blue bars represent a scale of 1cm.
Image 2.3 – Diagram to show web aspects of interest. Capture area is the area covered by sticky spirals. The
measurement of length between the inner-most spiral and outer-most spiral along a radial thread is divided by
the number of spirals that pass through this length in order to obtain the average mesh spacing. A ruler is placed
adjacent to the web in order to set a scale on Image J before digitally measuring webs. Blue scale bar represents
2cm.
Upper stabilimentum
Lower stabilimentum
Discoid stabilimentum
Hub
Free zone
Radial thread
Mesh spacing
Capture spiral
25
The area of the characteristic missing sector in the webs of Z. x-notata was measured to
investigate the plasticity of this structure under the environmental conditions provided. The
area of the space between the signal thread both adjacent radial threads (left and right of signal
thread) within the missing sector was also measured to explore plasticity in the symmetry of
this structure. These area measurements were taken within the two outer most spirals adjacent
to the signal thread, as shown in Image 2.4. If the signal thread was constructed outside of the
vertical plain, it was only possible to obtain the area of the whole sector and not the area either
side of the signal thread.
Image 2.4 – Orb web of Zygiella x-notata with the missing sector outlined in yellow. The red line shows the
limit of area measurement in the missing sector, as area measurements were taken within both of the outer most
spirals adjacent to the signal thread. The area of the space between the signal thread of both adjacent radial
threads (green) was also measured, again, within the red line. Blue bar represents 2cm.
Signal thread
26
2.7 - Statistical methods
With the use of SPSS statistical software, a Friedman’s test was used to investigate the extent
of variation in the web building of juvenile A. bruennichi, adult A. bruennichi and Z. x-notata
over time under constant abiotic conditions. The assumptions of the Friedman’s test are that
one group is measured on three or more occasions (the orb webs of spiders over 3 or more
days), the group is a random sample from the population, the dependant variable (web aspects)
is measured at the continuous or ordinal level and data are not normally distributed. The
Friedman’s test is often considered the non-parametric version of the one-way ANOVA. The
significance level chosen was 0.05.
A Wilcoxon signed-rank test was used to compare web construction by Z. x-notata in both
bright and dark conditions in order to compare differences in the construction of web aspects
in both conditions. For this test, the dependant variable is also measured at the continuous or
ordinal level (web aspects), there must be at least one independent variable (light intensity)
arranged into two or more categories (light and dark condition) to allow comparison and again,
this test is non-parametric as data are not normally distributed. This test takes into account that
the samples are paired and compares the difference in web aspect-building within each spider
in the light and dark condition. The significance level chosen was 0.05.
Chapter 3 – Results
In total, 261 webs were built by all 41 individual spiders from both Argiope bruennichi and
Zygiella x-notata in all experiments overall (Appendix – Table A1). 138 webs were built by
both species in the experiments that investigated web building variation under a constant abiotic
environment. 32 webs of these webs were built by all adult A.bruennichi, with 18 webs between
6 individuals used for analysis; juvenile A. bruennichi built 56 webs overall, with 40 webs
between 10 individuals used for analysis and Z. x-notata built 50 webs overall, with 33 webs
built by 11 individuals used for analysis. In the light intensity experiment, 123 webs were built
by 14 individual Z. x-notata overall with 58 webs built in light condition (3-6 webs built by
each individual) and 65 webs built in dark condition (3-8 webs built by each individual). These
27
subsections of webs used by each sample group were based on the number of webs built by
spiders after the three acclimatisation webs. Post-acclimatisation webs used were then rounded
down to the number of webs most frequently constructed among all spiders in each group for
data analysis. This ensured that the same number of post-acclimatisation webs were analysed
for each spider.
3.1 - Web building variation under constant abiotic conditions
Web building variation in a constant abiotic environment was investigated to test the null
hypothesis that all of the web aspects measured would not significantly vary over time under
constant conditions. The Friedman’s tests show that the null hypothesis can be accepted for
Zygiella x-notata as none of the web aspects measured significantly varied. However, it can be
rejected for adult and juvenile Argiope bruennichi for some specific web aspects, as there was
significant variation in the construction of upper mesh spacing in the juvenile A. bruennichi
webs and radii number in adult A.bruennichi webs over time.
3.2 - Juvenile Argiope bruennichi
Starting with juvenile A. bruennichi, only upper mesh spacing showed marginally significant
variation (N=10, χ²=8.040, p=0.045) (Figure 3.1) under constant environmental abiotic
conditions (20°C daytime, 40% RH day and night, 137.5 micro moles per squared, 16°C night
time, 15 hours light/ 9 hours dark). Juvenile A. bruennichi webs aspects that did not
significantly vary over time include radii number (N=10. χ²=4.355, p=0.226); lower mesh
spacing (N=10, χ²=4.394, p=0.222), left mesh spacing (N=10, χ²=6.184, p=0.103), right mesh
spacing (N=10, χ²=5.880, p=0.118), capture area (N=10, χ²=1.800, p=0.615) and centre area
(N=10, χ²=5.160, p=0.16) (Table 3.1).
28
Table 3.1 – Variability of the web parameters of 10 juvenile A. bruennichi over time. Asterisk denotes
significant variation over time, as determined by the Friedman’s test (p<0.05).
Web parameter Chi-square Days/webs No. of individuals P value
Radii no. 4.355 4 10 0.226
Lower mesh
spacing
4.394 4 10 0.222
Upper mesh
spacing 8.040 4 10 0.045*
Left mesh spacing 6.184 4 10 0.103
Right mesh spacing 5.880 4 10 0.118
Capture area 1.800 4 10 0.615
Centre area 5.160 4 10 0.16
Figure 3.1 – Scatter-line plot to show significant variation in upper mesh spacing within and between 10 individual
juvenile Argiope bruennichi over a 4-day period under constant abiotic conditions. Each colour represents an
individual spider.
29
Figure 3.2 – Bar graph to show mean upper mesh spacing measurements (y-axis) for each juvenile A. bruennichi
individual over a four-day period. Standard deviations, as represented by error bars, show variation in this web
parameter within individuals. Each colour represents an individual spider.
Although the Friedman’s test results show that centre area, radii number and capture area did
not significantly vary in the webs of juvenile A. bruennichi over a 4-day period (Table 3.1),
variation in these web characteristic measurements within each individual spider (6% - 18.5%
coefficient of variance range; Table 3.2; Fig. 3.3, 3.5 and 3.7) is lower than variation of these
web characteristic measurements between individuals (24.03% - 28.1% CV range) on each day
(although narrower CV range 24.03% - 28.1%; Table 3.3; Fig. 3.4, 3.6 and 3.8) when
comparing coefficient of variance values (Table 3.2 and 3.3).
0.98(Av)
1.13(Av)
1.08(Av)
1.16(Av)
1.05(Av)
1.76(Av)
1.36(Av)
1.22(Av)
1.09(Av)
1.05(Av)
0
0.5
1
1.5
2
2.5
Up
per
mes
h s
pac
ing
(mm
) 1
7
5
4
6
25
23
15
3
2
30
Table 3.2 - Coefficient of variance, expressed as a percentage, within each individual juvenile A. bruennichi for
centre area, radii number and capture area measurements over the 4-day period (n=4 for each cell).
Spider 1 7 5 4 6 25 23 15 3 2 Average
Centre
area
15.4% 22.4% 6.3% 11.3% 5% 22% 11.8% 10.1% 22.7% 22.2% 14.92%
Radii
no.
10.3% 13.2% 8.4% 6.6% 7.6% 7.3% 8.9% 4.4% 12% 11.3% 9%
Capture
area
19.7% 17.8% 7.9% 17.4% 9.3% 18.5% 9.9% 3.5% 16.9% 22% 14.29%
Average 15.13% 17.8% 7.53% 11.77% 7.3% 15.93% 10.2% 6% 17.2% 18.5%
Table 3.3 - Coefficient of variance percentages among all individual juvenile A. bruennichi centre area, radii
number and capture area measurements on each of the 4 days (n=10 for each cell).
Day 1 2 3 4 Av
Centre area 49.7% 43% 40.8% 51.3% 46.2
Radii no. 10.4% 18.8% 16% 18.1% 15.83
Capture area 22.7% 21.7% 16.1% 14.9% 18.85
Av 27.6 27.83 24.03 28.1
31
Figure 3.3 – Whisker plot to show considerable centre area variation within each individual juvenile A.
bruennichi over a 4-day period (n=4). Each colour represents an individual spider.
Figure 3.4 – Whisker plot to show centre area variation among all individual juvenile A. bruennichi on each of
the 4 days (n=10).
32
Figure 3.5 – Whisker plot to show considerable radii number variation within each individual juvenile A.
bruennichi over a 4-day period. Each colour represents an individual spider.
Figure 3.6 – Whisker plot to show radii number variation among all individual juvenile A. bruennichi on each of
the 4 days appears to be less considerable than variation within individuals.
33
Figure 3.7 – Whisker plot to show considerable capture area variation within each individual juvenile A.
bruennichi over a 4-day period. Each colour represents an individual spider.
Figure 3.8 – Whisker plot to show capture area variation among all individual juvenile A. bruennichi on each of
the 4 days appears to be less considerable than variation within individuals.
34
3.3 - Adult Argiope bruennichi
Adult female A. bruennichi showed slightly different variations in web building over time, as
radii number significantly varied over time (N=6, χ²=8.667, p=0.013) (Figure 9) and upper
mesh spacing did not (N=6, χ²=0.273, p=0.873). However, like juvenile A. bruennichi, adult
female A. bruennichi did not show any significant changes over time in lower mesh spacing
(N=6, χ²=0.667, p=0.717); left mesh spacing (N=6, χ²=1, p=0.607), right mesh spacing (N=6,
χ²=0.261, p=0.878), capture area (N=6, χ²=4, p=0.135) and centre area (N=6, χ²=4.333,
p=0.115) (Table 3.4).
Table 3.4 – Variability of the web parameters of 6 adult A. bruennichi over time. Asterisk denotes significant
variation over time, as determined by the Friedman’s test (p<0.05).
Web
parameter
Species Chi-square Days/webs No, of
individuals
P value
Radii no. Argiope bruennichi 8.667 3 6 0.013*
Lower mesh
spacing Argiope bruennichi 0.667 3 6 0.717
Upper mesh
spacing
Argiope bruennichi 0.273 3 6 0.873
Left mesh
spacing
Argiope bruennichi 1.000 3 6 0.607
Right mesh
spacing
Argiope bruennichi 0.261 3 6 0.878
Capture area Argiope bruennichi 4.000 3 6 0.135
Centre area Argiope bruennichi 4.333 3 6 0.115
35
Figure 3.9 – Scatter plot to show significant adult female A. bruennichi radii number variation within and between
6 individuals over a 3-day period under constant abiotic conditions. Each colour represents an individual spider.
Plots for spider 22 and 30 are identical.
Figure 3.10 – Bar graph to show averages (y-axis) of adult A.bruennichi radii number counts over a 3-day period.
Standard deviations, as represented by error bars, show variation in number of radial thread built by 6 individuals
over 3 days. Each colour represents an individual spider.
Although the Friedman’s test results show no significant variation in upper mesh spacing,
centre area and capture area between adult A. bruennichi over a 3-day period (Table 3.4),
20
22
24
26
28
30
32
34
36
0 0.5 1 1.5 2 2.5 3 3.5 4
Day/Web
22
17
30
32
19
16
27.3 Av
31.7 Av
27.3 Av
26.3 Av
3.6 Av
29.67 Av
0
5
10
15
20
25
30
35
40
22
17
30
32
19
16
36
variation in these web characteristics appear to be low overall within the web building of each
individual spider. However, some spiders show much higher CV values than others according
to the coefficient of variation values for these web aspects (CV range: 9.77% - 53.87; Figures
3.11, 3.13 and 3.15). Variation in these web parameters among all individuals over the 3-day
period appears to be higher overall with a narrower CV range (CV range: 34.07% - 44.43%;
figures 3.12, 3.14 and 3.16). Upper mesh spacing and centre area variation is at its highest on
day 2 (Figures 3.12 and 3.14).
Table 3.5 - Coefficient of variance percentages within individual adult A. bruennichi for upper mesh spacing,
centre area and capture area measurements over the 3-day period.
Spider 22 17 30 32 19 16 Average
Upper
mesh
10.2% 24.1% 14% 19.5% 6.6% 22.2% 16.1%
Centre
area
26.8% 20.9% 10.3% 13.9% 19.8% 16.6% 18.1%
Capture
area
18.1% 3.8% 5% 36.7% 8.3% 45.2% 19.5%
Average 18.4% 16.3% 9.8% 23.4% 34.7% 53.9
Table 3.6 - Coefficient of variance percentages between all individual A. bruennichi upper mesh spacing, centre
area and capture area measurements on each of the 3 days.
Day 1 2 3 Average
Upper mesh 13.8% 29.8% 29.5% 24.4%
Centre area 53.8% 29.8% 62.7% 48.8%
Capture area 50.4% 42.6% 41.1% 44.7%
Average 39.3% 34.1% 44.4%
37
Figure 3.11 – Upper mesh spacing variation within each individual adult A. bruennichi over a 3-day period.
Each colour represents an individual spider.
Figure 3.12 – Upper mesh spacing variation among all adult A. bruennichi over the 3-day period appears to be
less considerable than variation within individuals, with most variation occurring on day 2.
38
Figure 3.13 – Centre area variation within each individual adult A. bruennichi over a 3-day period. Each colour
represents an individual spider.
Figure 3.14 – Centre area variation among adult A. bruennichi over the 3-day period appears to be less
considerable than variation within individuals, with most variation occurring on day 2.
39
Figure 3.15 - Capture area variation within each individual adult A. bruennichi over a 3-day period. Each colour
represents an individual spider.
Figure 3.16 – Capture area variation among all adult A. bruennichi over the 3-day period appears to be less
considerable than variation within individuals.
40
3.4 - Stabilimenta building
Stabilimenta building in juvenile and adult A. bruennichi was scarce, occurring only 16 times
between juveniles and adults within 8 individuals (4 times in adults between 2 individual adult
A. bruennichi and 12 times between 6 juveniles).
3.5 - Zygiella x-notata
Zygiella x-notata adult females showed no significant variation in any of the web parameters
measured over time: Radii number (N=11, χ²=3.659, p=0.161); lower mesh spacing (N=11,
χ²=3.455, p=0.178), upper mesh spacing (N=11, χ²=2.182, p=0.336), left mesh spacing (N=11,
χ²=3.818, p=0.148), right mesh spacing (N=11, χ²=5.091, p=0.078), capture area (N=11,
χ²=0.727, p=0.695), centre area (N=11, χ²=3.455, p=0.178), sector area (N=11, χ²=1.333,
p=0.513) (Table 3.7).
Table 3.7 – Friedman’s test results show no significant web building variation over time for adult female
Zygiella x-notata web parameters (p>0.05).
Web
parameter
Species Chi-square Days/webs No. of
individuals
P value
Radii no. Zygiella x-notata 3.659 3 11 0.161
Lower mesh
spacing
Zygiella x-notata 3.455 3 11 0.178
Upper mesh
spacing Zygiella x-notata 2.182 3 11 0.336
Left mesh
spacing
Zygiella x-notata 3.818 3 11 0.148
Right mesh
spacing
Zygiella x-notata 5.091 3 11 0.078
Capture area Zygiella x-notata 0.727 3 11 0.695
Centre area Zygiella x-notata 3.455 3 11 0.178
Sector area Zygiella x-notata 1.333 3 11 0.513
Although the Friedman’s test results show no significant variation in any of the Zygiella xnotata
web characteristics measured over time, there is low overall variation in majority of web
building characteristics within individual spiders over a 3-day period, with some individuals
41
showing much higher variation (Table 3.8, CV range: 7.09% - 45.5%). These web
characteristics include centre area (CV range for centre area: 4.4% - 39.6%; Figure 3.17) and
capture area (CV range for capture area: 5.2% - 50%; Figure 3.19), which are also low in overall
variation within individual adult and juvenile A. bruennichi. Radii number also has low
variability within individual Z. x-notata, (CV range for radii number: 2.7% – 20%, Figure 3.21)
and upper mesh spacing has low overall variability with much larger CV values shown in some
spiders (CV range for upper mesh: 7.6% - 43.8%, Figure 3.23), which is the same for adult A.
bruennichi (Figure 3.11) within individuals. Z. x-notata showed low overall variation within
individual left, right and lower mesh spacing over the 3 days with some individuals showing
much high variation than others (appendix: Figure A1, A3, A5). The unique Z. x-notata web
feature, the missing sector, showed very high area variation within some individual spiders,
while other individuals sustain more consistent sector areas (CV range for sector area: 10.5% -
173.2%, figure 3.25). The variation in all of these web characteristic measurements between
individuals on each of the 3 days is high (CV range: 32.63% - 52.96%; Figures 3.18, 3.20, 3.22,
3.24, 3.26 and appendix: Figure A2, A4, A6 respectively). These observations are particularly
apparent when comparing coefficient of variance values (Table 3.8 and 3.9).
Table 3.8 – Coefficient of variance percentage values for radii, left mesh, right mesh, lower mesh, upper mesh,
capture area, centre area and sector area measurements within individual Z. x-notata over the 3-day period.
Spider 40 34 30 32 17 13 24 31 33 42 66 Average
Radii no. 7.1 13. 6 13. 2 10.2
4.5 5.8 8.3 20 4.2 2.7 4.7 8.57
Left mesh 9.1 32. 3 28. 8 28.6
44.6
9.8 20.2
30.2 30.2
4.3 10.1 22.65
Right
mesh
3.6 8.1
21. 3 47.9
26.2
8.8 11 30.9 11.5
15.7 9.9 17.72
Lower
mesh
7.3 19
41. 7 29.4
10.2
15.4
11.7
33.75
4.5 24.6 10.4 18.90
Upper
mesh
7.6 25. 6 43. 8 35.2
41.2
17.3
18.6
26.9 10.6
11.6 24 23.85
Capture
area
7.1 8.2
50
27.9
26.6
21.5
5.2 46.7 40.3
33.8 8.9 25.11
Centre
area
4.4 25. 9 49. 4 11.9
20.4
7.2 20.2
20.4 26.8
39.6 36.5 23.88
Sector
area
10.5
33. 2 44. 6 173.2
57.7
31.7
86.6
19.3 33.1
173.2
118.1
71.02
Average 7.09
20. 74
36.
60
45.54
28.9
3
14.6
9
22.7
3
28.52
20.1
5
38.18
27.83
42
Table 3.9 – Coefficient of variance percentage values for web characteristic measurement between individual Z. x-
notata on each of the 3 days.
Day 1 2 3 Average
Radii no. 21.3 18 19.4 19.57
Left mesh 22.9 37.2 17.7 25.93
Right mesh 24.3 56.2 33.6 38.03
Lower mesh 17.2 40 32.3 29.83
Upper mesh 39 37.2 28.8 35
Capture area 33.1 71.2 54.3 52.87
Centre area 31.8 47 55.1 44.63
Sector area 71.4 116.9 58.8 82.37
Average 32.63 52.96 37.5
Figure 3.17 – Considerable centre area variation within each individual adult Z. x-notata over a 3-day period.
Each colour represents an individual spider.
43
Figure 3.18 – Centre area variation among all adult Z. x-notata on each of the 3 days is less considerable than
variation within individuals.
Figure 3.19 – Considerable capture area variation within each individual adult Z. x-notata over a 3-day period.
Each colour represents an individual spider.
44
Figure 3.20 - Capture area variation among all adult Z. x-notata on each of the 3 days is less considerable than
variation within individuals.
Figure 3.21 - Considerable radii number variation within each individual adult Z. x-notata over a 3-day period.
Each colour represents an individual spider.
45
Figure 3.22 – Radii number variation among all adult Z. x-notata on each of the 3 days is less considerable than
variation within individuals.
Figure 3.23 - Considerable upper mesh spacing variation within each individual adult Z. x-notata over a 3-day
period. Each colour represents an individual spider.
46
Figure 3.24 – Upper mesh spacing variation among all adult Z. x-notata on each of the 3 days is less
considerable than variation within individuals.
Figure 3.25 - Considerable sector area variation within each individual adult Z. x-notata over a 3-day period.
Each colour represents an individual spider.
47
Figure 3.26 – Sector area variation among all adult Z. x-notata on each of the 3 days is less considerable than
variation within individuals.
3.6 - Light intensity experiment
A Wilcoxon signed-rank test was used to investigate differences in web building in response to
different environmental light intensities. The results show that there was no significant
difference between the web parameter measurements in the webs of 14 individual Z. x-notata
in the light condition versus the dark condition, except for lower mesh spacing (Table 3.10).
This rejects the proposed hypothesis that capture area, radii number and all mesh spacing in the
webs of Z. x-notata would be reduced in the high light intensity condition and enhanced in the
low light intensity condition and the null hypothesis can be accepted as light intensity had no
effect on web construction apart from lower mesh spacing. However, 9 out of 14 spiders,
although marginally, significantly increased the lower mesh spacing in the brighter condition
(Table 3.10; Figure 3.27).
48
Table 3.10 – Differences in web parameter measurement from 14 individual Zygiella x-notata adult female webs
in both the light and dark condition. Results of the Wilcoxon Signed Rank test show no significant differences in
between the web parameters built in the light and dark condition, except for lower mesh spacing (p=>0.05 for all
web parameters except lower mesh, which shows p=0.048*). Overall mesh spacing is the mean of upper, lower,
right and left mesh spacing measurements for each spider in the light and dark condition.
Web
parameter
Species Test
statistic
(Z)
N P value Mean (L) Mean (D) SD (L) SD (D)
Radii no. Zygiella
x-notata -1.224b 14 0.221 26.7 28.47 5.58 3.77
Lower mesh
spacing (mm)
Zygiella
x-notata
-1.977b 14 0.048* 2.71 2.43 0.62 0.37
Upper mesh
spacing (mm)
Zygiella
x-notata
-8.47b 14 0.397 3 2.96 0.7 0.8
Left mesh
spacing (mm) Zygiella
x-notata -1.287b 14 0.198 2.64 2.42 0.71 0.46
Right mesh
spacing (mm)
Zygiella
x-notata
-0.910b 14 0.363 2.61 2.48 0.51 0.369
Overall mesh
spacing (mm)
Zygiella
x-notata
-1.601b 14 0.109 2.74 2.57 0.57 0.43
Capture area
(mm2)
Zygiella
x-notata
-0.659b 14 0.510 10577 11199.58 4439.61 3294.99
Centre area
(mm2)
Zygiella
x-notata
-.910b 14 0.363 895.95 984.19 324.78 312.68
Sector area
(mm2) Zygiella
x-notata -1.475b 14 0.140 694.35 590 333.92 257.26
49
Figure 3.27 – The average values of lower mesh spacing measurements in the light intensity experiment. 9 out of 14
Z. x-notata built larger lower mesh spacing in bright light intensity, although to no significant extent. ‘L’ represents
the lighter conditions and ‘D’ represents the darker condition. Each colour represents an individual spider.
Chapter 4 - Discussion
Web building in juvenile A. bruennichi, adult A. bruennichi and Zygiella x-notata was mostly
consistent over time under constant abiotic conditions and remained mostly consistent in
Zygiella x-notata, apart from lower mesh spacing, when exposed to different environmental
light intensities. The only web parameters that significantly varied over time under constant
conditions were radii number in adult A. bruennichi webs and upper mesh spacing in juvenile
A. bruennichi webs. This variation in A. bruennichi upper mesh spacing and radii number
rejects the null hypothesis that web building by both species would remain totally consistent
under the same abiotic conditions over time. The mostly insignificant changes in web building
by Z. x-notata in low and high light intensity environments somewhat accepts the other null
hypothesis, thus spiders do not invest less in the prey capture aspects (smaller capture area,
fewer radial threads and increased overall mesh spacing) in bright conditions when prey is
L
D
L
D
L
D
L
D
L
DL
D
LD
L
DL D
L
D
L
D
LD
L D
LD
0
0.5
1
1.5
2
2.5
3
3.5
4
4.5Lo
we
r m
esh
he
igh
t (m
m)
The affect of light intensity on lower mesh construction by Zygiella x-notata
69
71
72
73
74
77
79
83
89
90
92
84
85
50
likely to be more active. However, the decrease in mesh spacing shown by spiders in the dark
condition was marginally significant.
In order to balance foraging success and acclimatisation to laboratory conditions between
spiders, three webs were allowed to be built by each spider in the frame set up before
experiments took place. Allowing spiders to build three or more acclimatisation webs is
considered the best practice in preparation for laboratory-based experiments in previous
studies34.
4.1 - Juvenile upper mesh variation and overall mesh consistency
Orb web asymmetry has been observed in various spider species, which has led to the proposal
of multiple adaptive hypotheses. The sticky capture spirals of the orb web have been shown to
be more abundant and to cover a larger area below the hub in the lower portion of vertically-
built orb webs in most cases117, 118. Uncommon exceptions are represented by spiders in the
Scloderus genus as they tend to drastically enhance the area of their orb webs above the hub in
the upper portion of the web, where more spirals reside119. This type of asymmetry shown by
Scloderus is hypothesised to be an adaptation for catching moths, as a moth will often tumble
down the web and lose protective wing scales when its flight is intercepted. Once enough wing
scales are lost, the silk can adhere to the moth’s body and ensnare it120. Aside from area, some
genera, such as Micrathena and Gasteracantha, build non-vertical orb webs with a roughly
equal number of spirals in the lower and upper portions of the web121, suggesting that spiral
distribution may be genera-specific. In terms of mesh spacing, earlier studies describe the mesh
spacing of orb webs to be narrower in the lower portion of the web122, while later observations
indicate that there is no difference between mesh spacing in the upper and lower areas of the
web123. Some spiders, mostly those in the families Tetragnathidae and Uloboridae, built non-
vertical webs124. Mesh spacing in the non-vertical orb webs of Micrathena gracilis tend to be
smaller in the lower web region, while mesh is narrower in the upper region of Leucage
mariana, suggesting that mesh spacing could be interspecifically determined among different
species125. Mesh spacing has also been shown to be more uniform in the lower section of the
web126. Furthermore, arrangement of radial threads is often asymmetric. The angle between
51
individual radial threads tends to be smaller in the lower and often larger part of the web as this
would reduce the likelihood of sticky spirals contacting each other and fusing when displaced
by wind, which prevents web damage125. Orb weavers have also been known to ‘double’ radial
threads, by splitting one radial thread into two, in the upper portion of the web to structurally
cope with larger forces in this area127, 128.
Although not all aspects of asymmetry were measured in this study, the significant variation in
upper mesh spacing over time in the webs of juvenile A. bruennichi may relate to the
‘top/bottom’ asymmetry hypothesis, which proposes that the habit of orb weavers building a
larger capture area in the lower region of the web promotes the capture of prey there. This
would enable the spider to move downwards with gravity to catch prey, which is less
energetically costly than moving upwards against gravity129. Juvenile A. bruennichi may not
prioritise consistent upper mesh building as it is not a priority for prey capture129. Further
investigation, with both asymmetry and mesh spacing being measured simultaneously over
time, with similar methodology to this study, is required to understand the relationship between
upper mesh spacing and the top/bottom asymmetry in the webs of orb weavers and how this
relationship changes as a spider moults and grows.
The interspecific differences in the construction of mesh spacing shown in previous studies125
may support the results in this study, as no significant variation in upper mesh spacing over
time occurred in the webs of Zygiella x-notata, but did occur in the webs of juvenile A.
bruennichi. Perhaps these results suggest that Z. x-notata is a more consistent web-builder than
Argiope bruennichi under constant conditions. The uniformity of lower mesh building observed
in previous studies125 is also coherent with the results as adult A. bruennichi, juvenile A.
bruennichi and Zygiella x-notata all showed insignificant variation in lower mesh construction
over time. Future research should focus on differences in mesh spacing consistency between
various species in order to better understand interspecific patterns in the construction of this
web aspect and the possible functions that differing mesh sizes serve for different orb weavers
in contrasting habitats.
52
4.2 - Spatial constraints
In contrast to juvenile A. bruennichi, adult A. bruennichi did not show significant variation in
upper mesh spacing over time. This may be due to the restricted size of the smaller frames that
juveniles were kept in, as Araneids, including species in the Argiope genus, have been shown
to alter the geometry of their webs drastically to conform to the space available within different
sized frames130. Other studies have shown that Araneus diadematus elongates its web vertically
when exposed to narrower web-building spaces131. This treatment also caused A. diadematus
to reduce mesh size on the shorter horizontal radial threads near the left and right areas of the
hub, but mesh was larger along longer, vertical radial threads above and below the hub,
indicating that radii position and length influences the attachment points of the sticky spirals.
Furthermore, another Araneid, Leucage argyra, is capable of constructing extremely reduced
webs when kept in small frames that are only 7% of the mean span of webs in the wild for this
species132. L. argyra also demonstrated changes in orb web design at different stages of web
building, including radial thread and frame connection, building of the hub, layout of the sticky
spirals and termination of sticky spirals. Other orb weavers, such as the ladder web spider
Telaprocera maudae, significantly elongated their webs when kept in a narrow, horizontal
frame space and built more circular webs when kept in frames with an increased diameter in
web-building space133. This species positions its web against tree trunks in the wild and this
web-building plasticity would enable any tree of any trunk size to be inhabited throughout the
spider’s development and dispersal from tree to tree59. This behavioural flexibility may also
aid in maximising prey capture with the available space64.
The extent of building plasticity may be species-dependant, as Cyclosa caroli showed very low
web-building frequency when kept in smaller, elongated frames, whereas Eustala ilicita
showed higher web-building frequency and was quick to adapt its webs to the limited space,
indicating that behavioural plasticity between these two phylogenetically close species is
influenced by the microstructure of the web-building area134. E. ilicita may require building
flexibility, as it inhabits specific vegetation where it occurs in high densities135, whereas C.
caroli occurs in low density in various understory vegetation, where it would naturally have
more space and would not be accustomed to building in restricted spaces136. In summary,
perhaps juvenile A. bruennichi would have kept upper mesh spacing more consistent if
53
provided with more space. It is interesting to note that lower mesh was kept consistent, which
may indicate prioritisation as catching prey below the hub is more energy efficient129.
The consistent building environment of the frames may also explain the lack of variation in the
majority of web parameters, apart from adult A. bruennichi radii number and juvenile A.
bruennichi upper mesh spacing, over time among all sample spiders from both species under
constant abiotic conditions as orb weavers can use anchor points from previously built webs at
the same site as a reference for building new webs, reducing cognitive demand137. The frame
size and shape remained constant throughout experiments and although webs of Z. x-notata
were destroyed after they were photographed, the frame, bridge and anchor threads remained
intact to promote web repair. This would have enabled the spiders to build webs in exactly the
same building space where they were likely to use the frame, bridge and anchor threads from
their previous webs as reference points. This may explain the overall lack of web variation in
both species used in this study in laboratory conditions, but these findings may not reflect web
variation in the wild, as spiders often disperse or are displaced by environmental factors in
nature, increasing cognitive demand to build webs at new sites133. Furthermore, this could be
why there were almost no significant differences in web building shown by Zygiella x-notata
in the light intensity experiment when exposed to a high light intensity environment and a low
light intensity environment as perhaps the spatial aspects of the environment, which were
constant, are more influential than the abiotic conditions on web building behaviour.
Although the effect of varying web-building space on orb web geometry has been well
documented, the diversity of sample species in previous studies is relatively small as there are
approximately 4,500 orb weaving spider species known worldwide138. This provides vast
opportunity in future research to investigate how the many different orb weaving species, which
have not been used in previous experiments, adapt their web building to spatial constraints and
how the extent of building plasticity over time changes between phylogenetically distant and
close species that differ in their ecology.
54
4.3 - Web building experience
Juvenile orb weavers are known to build circular, more symmetrical webs139, which challenges
the previously mentioned ‘top/bottom asymmetry’ hypothesis. Larinoides sclopetarius adults
that are deprived of web building experience have subsequently been observed building more
circular, symmetrical webs much like those of many juvenile orb weavers73. Some A.
bruennichi adults and juveniles in this study built webs readily and more frequently than other
individuals in the frame set up. There was variation in the number of webs built between
individual spiders, as although four webs from each juvenile and three webs from each adult
were used for analysis, some built up to seven webs (Appendix – Table 1). Perhaps the
infrequent web-builders lacked the experience required to maintain consistent web geometry
and this may have influenced increased radii variation in adult webs and upper mesh variation
in juvenile webs. Adult A. bruennichi that built infrequently may have shown this previously
observed tendency to build more symmetrical webs when deprived of web building experience,
while frequent web builders built asymmetrical webs. However, this does not explain the upper
mesh variation in juvenile A. bruennichi and further investigation into mesh variation in the
upper and lower regions of the orb web in experienced and inexperienced spiders is required to
understand this phenomenon more thoroughly.
4.4 - Size limitation hypothesis
Another hypothesis, the ‘size limitation hypothesis’, may provide an alternative explanation for
the lack of web building variation in most of the web aspects measured over time under
consistent abiotic conditions in juvenile A. bruennichi, as it suggests that young animals with
underdeveloped brains are limited in their behavioural flexibility140. This hypothesis also
suggests that young animals, including juvenile spiders, may be more prone to be making
‘errors’ when building multiple webs over time, limiting the consistency of web geometry. This
seems logical, as smaller neurones in juvenile spiders would presumably limit the complexity
of behaviours141. However, it has been difficult to show convincing experimental support for
this142. These previous findings may be reflected in the results from juvenile A. bruennichi
building, as keeping upper mesh spacing consistent, along with the rest of the web, may have
been beyond their cognitive ability while their central nervous systems are still developing.
However, future experiments with similar methodology, a larger sample size of juvenile spiders
55
and a wider range of species would investigate mesh ‘errors’ in more detail and could
potentially show how these errors differ interspecifically, as different species may show
inconsistencies in different web characteristics according to their ecology and habitat.
4.5 - Adult Argiope bruennichi radii variation
The number of radial threads in the webs of adult A. bruennichi varied significantly over time,
whereas juvenile radial threads did not significantly vary. This may be due to the general
increase in the volume and the strength of the silk in adult spiders143, 64, which reduces the need
for consistent radial thread construction to structurally support the web144. The fact that some
webs would have been more frequently destroyed by prey impact could have influenced
variation in radii number. Radial threads provide the majority of structural support in the
web144and due to the difference in web building and web destruction frequency between
individuals, perhaps spiders that experienced more web damage constructed more radial
threads, whereas infrequent web-builders added fewer radii to their webs. Previous studies on
the effect of web damage on web-building have found spiders that experience a lot of web
damage tend to travel greater distances in search of new web building sites when compared to
spiders that did not experience any web damage145, 146. Furthermore, web damage is highly
energetically costly for spiders due to a loss of silk and effectiveness of the prey trap147. The
lack of web building performed by some individual A. bruennichi may simply be due to the
conservation of silk in an environment where web damage by prey was frequent. Alternatively,
energetic strain of prey impact rates on webs may have been influential. One prey item was
placed in the web every time a new web was built, but webs were rebuilt at different frequencies
between individual spiders as some adults built up to seven webs within the same time frame
as other individuals that only built three to four webs (the first three experimental webs from
each individual adult were used for analysis). Therefore, it could also be the number of prey
impact events on the webs of adult A. bruennichi that caused radii number to significantly vary
over time, as spiders that experienced high rates of prey impact would, in theory, increase radii
number over time to cope with the higher kinetic energy strain on the web57.
56
However, a larger sample than six individual adults would have shown more representative
results. Furthermore, prey was placed in the webs of spiders and was not entered into the webs
with force, which would presumably make prey impact on the web an unlikely influence.
Further investigation into radial thread construction in the subsequent webs of spiders that have
experienced frequent significant web damage is needed to understand the importance of radii
as a supportive aspect of the orb web in more detail. Further research on this topic may also
improve knowledge on the web-building decisions made by spiders in an environment where
damage to a valuable resource is frequent.
4.6 - Biotic factors more influential on web construction?
4.6.1 - Effect of food intake
In the experiment to investigate web building variation under constant abiotic conditions in
A.bruennichi and Zygiella x-notata, keeping biotic conditions consistent throughout the course
of web building was more difficult than keeping abiotic conditions consistent. The
cephalothorax widths (mm) did not vary much within each group (adult A. bruennichi
CV=10.98%, juvenile A. bruennichi CV=24.48%, Zygiella x-notata CV=10.44%), but the mass
(mg) of each spider was more variable within each group (adult A. bruennichi CV=31%,
juvenile A. bruennichi CV=69%, Zygiella x-notata CV=35%). Adult A. bruennichi were
consistently fed on various Diptera species of similar size (10-15mm body length) and juvenile
A. bruennichi and Z. x-notata were consistently fed Drosophila. However, spiders were only
fed when they had built an orb web and some individuals built more frequently in the frame
set-up than others, resulting in an irregularity of food intake among individuals.
Orb weavers have been known to increase the frequency of their web building and to increase
variability in mesh and capture area construction in response to decreased prey capture, while
showing a reduction in web building frequency and an increase in capture area over time when
prey availability is increased148. This may relate to the significant variation in juvenile A.
bruennichi upper mesh spacing, as these spiders were collected from the field and may have
57
had different foraging success in the wild before they were collected. Perhaps juvenile A.
bruennichi with low foraging success prior to collection exhibited high variability in mesh
spacing. However, this does not provide a potential explanation for left, right and lower mesh
remaining consistent throughout the experiment. In addition, spiders that are in better
nutritional condition have been shown to use less silk in their web building25, so perhaps radial
thread investment was variable between spiders of differing nutritional states, which could have
caused the significant variation in radii number over time in adult A. bruennichi webs. Again,
the sample size of adult A.bruennichi was small (six individuals) and a larger sample may have
shown different results. Also, this does not explain the overall consistency in other web
characteristics shown by adults under constant abiotic conditions and the total consistency in
web building by adult Z. x-notata. Some of the adult A. bruennichi may have had more foraging
success and had obtained more nutrients than other individuals before they were collected from
the field, which could be indicated in the variation in mass, which was high among the three
sample groups.
4.6.2 - Prey type
Prey characteristics have been shown to influence web building with convincing supporting
evidence15, 18. A more recent study observed Nephila pilipes investing more silk in a large
capture area, more radial threads and longer spiral threads when fed on live Diptera compared
to being fed on crickets, which triggers a reduction in radii number, radii length and enlarged
mesh spacing to create a stiffer, more tense web that would be more ideal for catching larger
heavier prey with a greater kinetic energy output149. In this study, all spiders were fed the same
type of prey throughout their web building time apart from slight differences in prey offered to
adult A. bruennichi, which may explain the lack of variation and the mostly consistent building
by Z. x-notata in light and dark conditions. As mentioned before, prey was placed in the webs
of spiders in this study rather than entered with force, which would presumably have a low
impact on the web and a low kinetic energy output. As previously mentioned, the presence of
visually acute predators can also be influential on an orb weaver’s web construction, but this
was not present in this study22, 23. However, there are contradicting data, as orb weaver species
exposed to Odonata and Hymenoptera prey in two separate treatments showed no changes in
58
mesh spacing in their webs, although one of the species, Larinoides cornutus, delayed its web
building in the presence of the potentially dangerous Hympentoperan prey150.
4.6.3 - Prey size
The Dipteran prey size that was available for adult A. bruennichi ranged from 10-15mm in body
length and although this seems like a small size range, to the spiders it may be a large enough
range to influence the number of radial threads that were constructed. Larger prey tends to
produce a larger kinetic energy output151, which would require more radii to increase
dissipation of this energy. Perhaps only offering prey that was a more consistent size would
have shown less variation in radii number between individuals. Spiders that have less frequently
eaten are also known to spread their silk resources more sparingly over a web with a larger
area, where radii number is reduced and mesh spacing is increased. This web design may
potentially aim for the interception of larger prey, while smaller prey pass through the web, in
times of hunger while investing less energy in silk152. Various studies have investigated the
effect of prey type on web building15, 18, but future research should investigate how small/large
the range of prey size captured by spiders must be to cause significant variation in web building
over time. This will also provide valuable information on how consistent prey size must be kept
in studies that are aiming to test the effects of other treatments on web building.
4.6.4 - Hydration
Intake of water is considered to be vitally important for producing silk153 and spiders need to
drink loose water as well as obtaining water from their prey154. Water has been observed
collecting in stabilimenta in the webs of Argiope and these spiders actively search for water on
these web decorations155. The webs of adult A. bruennichi and Z. x-notata were sprayed with
water at the same time as feeding. Again, there was likely to be an imbalance of hydration
levels between spiders due to irregular web building, with hindered silk production in
infrequently hydrated spiders.
59
4.6.5 - Effect of aging
Aging can negatively impact the performance of silk glands, causing older spiders that are
coming to the end of their life to invest less energy in constructing the orb web156, 157. Although
all adult A. bruennichi in this study were collected as adult females, they may have slightly
varied in their time of maturation in the field and this information was unknown. This may be
reflected in the significant variation in radii number constructed by adult A. bruennichi over
time, as the older spiders may have invested less in radial threads due to energy constraints and
hindered performance of the major ampullate silk glands. However, adult A. bruennichi were
collected at the same time of year, mostly in late July, to reduce the chance of significant age
differences.
4.6.6 - Pollen consumption
A. bruennichi webs were dusted with pollen in order to indicate a newly built web when a
pollen-free web had been constructed. Pollen ingestion by young Araneus diamematus has been
shown to double the lifespan and significantly increase web building frequency when compared
to spiders that were fed on fungi or were starved, suggesting that pollen is a major source of
nutrients for juvenile orb weavers158. As webs among juvenile A. bruennichi were not always
built in synchronisation, some spiders would have likely ingested more pollen than others when
recycling and rebuilding the web. This may have cause subsequent web-building to be more
frequent in some individuals and could have possibly influenced the significant variation in
upper mesh over time.
4.6.7 - Effect of reproduction on web construction
Although this study did not investigate the effect of gravidness on web building, visually it
appears that there could be a reduction of energy investment in the webs of three individual Z.
x-notata (spiders 30, 72 and 73) that produced egg sacs during the light intensity experiment.
The webs appeared to become smaller and more deformed in the build up to the day when the
egg sacs were laid and appeared to contain a lower density of glue droplets on the flagelliform
60
spirals after the day of egg sac production. This was particularly clear to see in the web building
of spider 73 (Images 4.1—4.4). Capture area and total thread length has been previously shown
to decrease over the five webs prior to egg sac production, becoming most reduced the day
before eggs were laid13. These previous observations also show that after egg sac production,
mesh height stayed larger while overall thread length gradually increased over five subsequent
webs, indicating that investment in the web is reduced in preparation for egg sac production13.
However, Z. x-notata web building did not significantly change at all throughout the light
intensity experiment.
Image 4.1 – Normal web constructed by Z. x-notata 73 the day before the egg sac was laid.
61
Image 4.2 – Reduced and deformed web constructed when egg sac was found in the enclosure.
Image 4.3 – Web lacking flagelliform glue droplets constructed after the day of egg sac production. The web
constructed the day after looked similar.
62
Image 4.4 – Third web constructed since egg sac was laid. The web has returned to its normal state, with
increased visibility due to more glue droplets on flagelliform silk.
These images appear to show reduced investment in flagelliform silk (sticky spirals) on the day
and the two subsequent days after the egg sac was produced. Perhaps if future studies focused
on the webs of a larger sample of gravid Z. x-notata individuals prior to production of egg sacs
under constant conditions, variation patterns in web aspects would be more obvious. It is also
important for subsequent research to observe changes in web building post egg production to
see if web geometry consistently returns to an original state or is permanently affected by egg-
laying.
4.7 - Light intensity experiment (Zygiella x-notata)
Reduction in web area in high light intensities, where insects tend to be more active, has been
observed previously36. Building smaller webs in bright conditions may also have an
antipredator function, to avoid prolonged periods of activity and exposure to predators42.
However, again, results from this study show the contrary as there were mostly no significant
63
differences in the web geometry of Z. x-notata webs in lighter and darker conditions in the light
intensity experiment.
However, the 14 Z. x-notata did build significantly larger lower mesh spacing in brighter
conditions than they did in darker conditions. The reason for this may relate to silk investment
and prey activity. Presumably, silk investment would be reduced when Z. x-notata built webs
with larger mesh spacing in the bright condition16, 17. This could be due to prey tending to be
more active in brighter conditions100, 101 as spiders can afford to invest less energy in the web
due to the chance of catching prey being higher in a brighter environment. Why change just
lower mesh spacing? Referring back to the ‘top/bottom’ asymmetry hypothesis, it is more
energy efficient for orb weaving spiders that build webs vertically to capture prey in the lower
portion of the web. This is so that the spider can travel downwards with gravity to capture the
prey, rather than against gravity129. This is thought to be why many orb weavers show increased
capture area below the hub117, 118. Z. x-notata may have reduced lower mesh spacing in the dark
condition to prevent smaller prey passing through the web and so that, importantly, prey of
almost any shape or size can be caught in conditions when prey is more scarce15. Potentially,
these spiders could be using light intensity as a proxy for predicting prey activity, but as silk
investment was not measured and quantified it is not possible to come to this conclusion with
confidence. However, it does provide an opportunity for future research to focus not only on
web geometry differences in different light intensities, but also silk investment and energy
expenditure. It is also important to note that this difference in lower mesh spacing in the light
and dark condition is only marginally significant (p=0.048) and a larger sample of spiders may
produce different results.
Z. x-notata, from personal observation, tend to occupy the hub at night and are more active at
this time, while they use their silken retreat during the daylight hours. In fact, many of the
individuals collected in the field were found at night on their webs. They also tend to rebuild
their webs in the early hours of morning, when there is minimal light. Perhaps night time
temperature and humidity are more influential on this species’ web building, as they would be
foraging/constructing webs at this time. However, it appears that day time light intensity does
not influence the majority of web building of Z. x-notata, apart from lower mesh spacing
potentially. It would be ideal if impending research investigated the effect of varying
64
temperature and humidity at night on the web building of Z. x-notata. This may provide the
spiders with more environmental cues that would influence web construction decisions when
the spider is more active and closer to the time of web rebuilding.
It would have been ideal to investigate the effect of different light intensities on the web
building of Argiope bruennichi, especially with the stabilimenta aspect. However due to the
seasonality of this species and the time available, this was not possible. I would encourage
future research into the effect of light intensity on orb web structure to focus on Argiope species
as well.
4.8 - Hierarchy of variation
Previous studies have mostly focused on the influence of a changing environment on the
construction of orb webs by spiders, but there is a distinct lack of focus on how web geometry
varies over time in a constant environment. In this study, there was low variation within the
web building of individuals over each day from all sample groups (low CV values), although
the CV values for each individual Z. x-notata differed greatly (Tables 3.2, 3.5 & 3.8). Variation
in geometry of webs among all individual spiders on each day was greater (Tables 3.3, 3.6 &
3.9), but the CV values on each day are more similar to each other. Web variation between
different individuals makes sense in regard to the previously mentioned factors that naturally
differ from one individual to the next, such as age, individual condition and web building
experience. Age may be a contributing factor to the web building variation within and between
individuals over time in adult spiders as the production of silk generally decreases as time until
natural death decreases156, 157. This unclear trend in response to abiotic factors may be due to
the biotic environment having a stronger influence on web building behaviour.
Referring back to the ‘size limitation hypothesis’, it may be that variation within individual A.
bruennichi juveniles is due to ‘errors’ in web building while the central nervous system is still
developing140, but this does not provide an explanation for a similar pattern in the webs of adult.
A. bruennichi and Zygiella x-notata. Further research is needed to explore web building
65
variation within individuals in a constant environment to generate hypotheses on the adaptive
function of changing web structure frequently over time and what mechanisms direct this
variation.
4.9 - Lack of stabilimenta
Stabilimenta only occurred 16 times out of the 88 webs built by all adult and juvenile A.
bruennichi and the lack of these web decorations is unclear. However, this could be explained
by findings in previous studies on stabilimenta. Wind is one of the main abiotic factors that
would occur in nature, but it was not taken into consideration in this study due to the absence
of wind in the laboratory set up. Orb weaving spiders have been observed enhancing
stabilimenta as environmental wind speed increases159. In regard to the theory of a signalling
function of stabilimenta, this could possibly be explained by strong winds increasing the
likelihood of birds contacting the web, making a conspicuous signal on the web more necessary
to avoid web damage, or it could be the wind generated by birds flying close that triggers an
antipredator stabilimenta-building response160. The area of stabilimenta has also been shown
to increase in response to airborne vibrational stimuli that mimics the approach of a predatory
Hymenopteran/Dipteran, supporting an antipredator function161. Perhaps the spiders in this
study would have built stabilimenta more frequently if exposed to wind or other airborne
stimuli in the environment, which would occur in the wild.
The physical condition of a spider has been considered a key factor in determining stabilimenta
construction. Body mass and abdomen length have been shown to positively correlate with
stabilimenta area, but smaller, lighter spiders are more likely to choose to add decorations in
the first place162. This is consistent with observations in this study, as the presence of
decorations was more frequent in the webs of juvenile A. bruennichi (12 out of the total 16
stabilimenta webs recorded). The general lack of stabilimenta may be due to reduced food
consumption in the lab (one prey item per newly built web), as web-building spiders in the field
would capture more prey items during peak insect activity163 and some of these prey items may
be ~200% of the spider’s body size164, although optimal prey length usually ranges from 50%
- 80% of the spider’s body length in most species165. Therefore, spiders in the field are expected
66
to be heavier, longer in body length and better nourished than in the lab. A. bruennichi in this
study rarely built webs in synchronisation and webs were built at different frequencies between
individuals. This combination of reduced food consumption and varying web building
frequency may be reflected by the irregularity of stabilimenta construction. However, the
decision of stabilimenta construction by healthy spiders may be based on other factors as
stabilimenta only accounts for approximately 10% of the dry weight of the web, which would
presumably be energetically inexpensive to build166.
Stabilimenta construction may also be influenced by the abundance of prey in the environment.
Positive associations have been made between stabilimenta building of Argiope argentata and
the increased abundance of its main prey, stingless bees167. The lack of stabilimenta building
by A. bruennichi in this study may due to the lack of prey in the spiders’ environment within
the laboratory frame set up. Spiders only experienced the presence of prey when flies were
placed into their webs. In regard to the prey attraction theory, perhaps detection of prey
abundance in the environment, before it is captured, is a key trigger for web decoration-
building.
4.10 - Preliminary observations of stabilimenta
In a preliminary experiment to test to effect of temperature on the web building of Argiope
bruennichi, the presence of stabilimenta was noticeably different between the cooler control
condition (day: 157.3µmol m-² light intensity, 20°C and 40% RH. Night: 16°C, 40% RH and
total darkness. 15h light/ 9h darkness) and the warmer experimental condition (day: 77.3µmol
m-2, light intensity 25°C and 40% RH, 15h light/ 9h darkness) with stabilimenta occurring 16
times in the control condition (97 webs built by 6 sdults and 10 juveniles, with stabilimenta
occurring 12 times in juvenile webs) and only once in the experimental condition (41 webs
built by 3 adults and 6 juveniles with stabilimenta occurring once in a juvenile web). However,
due to difficulties keeping the light intensity equal in each condition due to faults in the
chambers, it is not possible to determine whether it was temperature or light intensity triggering
the building of stabilimenta. There were also fewer and different individual spiders in the
67
experimental condition, meaning that the populations used were not independent or paired in
this preliminary experiment. That said, this does provide a platform for further research into
the affect that light intensity and/or temperature has on stabilimenta building decisions by
Argiope bruennichi and the ontogenetic influence on stabilimenta building.
4.11 - Conclusion
The aim of this study was to investigate the influence of change in environmental light intensity
on the web building behaviour of an orb weaver and to show the degree of web building
variation under constant abiotic conditions. It was predicted that Z. x-notata would enhance the
prey capture aspects of the orb web in lower light intensities, as in theory, insect activity would
be lower in these conditions. Web building variation by Z. x-notata and A. bruennichi was
expected to be insignificant in a constant abiotic environment due to the lack of environmental
cues that could potentially trigger a change in web building behaviour.
In conclusion, radii number in the webs of adult A. bruennichi and upper mesh spacing in the
webs of juvenile A. bruennichi significantly vary over time in a constant abiotic environment.
However, the majority of web parameters remained consistent over time. Upper mesh variation
over time in juveniles may be influenced by the ‘top/down asymmetry’ hypothesis, spatial
constraints of the frames and/or ontogeny. Radii variation over time may be influenced by slight
variations over time in prey size, web building frequency, feeding, drinking and web damage
by prey. However, the exact cause of these variations remains unclear and further research is
required to isolate each of these potential effects in order to observe their impact on web
building in A. bruennichi.
There have not been many previous studies that have investigated web building variation under
constant abiotic conditions, but the existing studies have shown that building variation in a
constant environment can be significantly variable98. Here, the results mostly show the
opposite, with an overall lack of variation in the webs of A. bruennichi apart from radii in adult
webs and upper mesh in juvenile webs, and no significant variation in the webs of Z. x-notata.
In general, individual spiders from both species vary in their web building over time (shown
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by coefficient of variation within individuals), but a population of spiders shows no overall
trend over time. This indicates that web building variation within individuals could be random,
as the results could possibly suggest that differences in web building among a population are
not influenced by the abiotic environment, nor does a population vary the majority of web
aspects in a constant environment. However, further investigation is required to support this.
There were almost no differences in the geometry of Z. x-notata webs in both treatments in the
light intensity experiment, which may be due this species’ habit of being more active at night
and less sensitive to day time light intensities. However, lower mesh spacing was marginally
significantly reduced in the dark condition, which may relate to lower prey abundance in a
darker environment and investment in silk production. Further investigation on the effect of
night time abiotic factors, such as humidity and temperature, on Z. x-notata web building may
be more applicable to this species. However, it could also be that the biotic environment is
more influential on web building than the abiotic environment, which is supported by findings
from various studies in the literature13, 15, 18, 22, 23, 25.
These findings have enhanced the limited understanding of adaptive orb web construction in
response to changing abiotic factors by an orb weaver, which has proved to be mostly lacking.
Variation in the fine details of the orb web over time in a constant abiotic environment is shown
to be totally absent, filling important gaps in the literature on this rarely studied topic.
69
Appendix
Table A1 – Total number of webs built by juvenile A. bruennichi, adult A. bruennichi and Zygiella x-notata over
time. Numbered green boxes represent number of webs built by each spider. The first four webs built by each
juvenile A. bruennichi were used for analysis. The first three webs built by adult A. bruennichi and Z. xnotata
were used for analysis.
Spider Juvenile A. bruennichi
1 1 2 3 4 5 6 7
2 1 2 3 4 5 6
3 1 2 3 4 5 6
4 1 2 3 4 5 6
5 1 2 3 4 5 6
6 1 2 3 4 5 6
7 1 2 3 4 5
23 1 2 3 4
15 1 2 3 4 5 6
25 1 2 3 4
Adult A. bruennichi
22 1 2 3 4 5 6
17 1 2 3 4 5
30 1 2 3 4 5 6 7
32 1 2 3 4 5 6 7
19 1 2 3 4
16 1 2 3
Z. x-notata
40 1 2 3 4
34 1 2 3 4 5 6
30 1 2 3 4 5 6 7 8
32 1 2 3 4 5
13 1 2 3 4 5
70
17 1 2 3 4 5
24 1 2 3 4 5
31 1 2 3
33 1 2 3
42 1 2 3
66 1 2 3
Figure A1 - Considerable left mesh spacing variation within each individual Z. x-notata over
a 3-day period. Each colour represents an individual spider.
71
Figure A2 – Left mesh spacing variation between adult Z. x-notata on each of the 3 days is less
considerable than variation within individuals.
Figure A3 - Considerable right mesh spacing variation within individual adult Z. x-notata over
a 3-day period. Each colour represents an individual spider.
72
Figure A4 - Right mesh spacing variation between adult Z. x-notata on each of the 3 days is
less considerable than variation within individuals.
Figure A5 - Considerable lower mesh spacing variation within individual adult Z. x-notata
over a 3-day period. Each colour represents an individual spider.
73
Figure A6 - Lower mesh spacing variation between adult Z. x-notata on each of the 3 days is
less considerable than variation within individuals.
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