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Diversification of Spider Silk
Biomechanics
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Orb weaving spiders
spin silks from seven different glands, all of which are used for
different functions. Some of these silks are as strong as steel, while
others are as stretchy as rubber. These silks are constructed from a small
array of proteins, providing unique opportunities to link external selective
pressures on the mechanical performance of spider silks to evolutionary
changes in silk genes. Based upon research that I began in the laboratory of
Cheryl
Hayashi, I am currently characterizing the biomechanical
properties of this diverse tool kit of silks for many spiders to study how
the mechanical performance of silks changes during behavioral shifts in how
spiders construct webs, such as the evolution of capture webs and the
development of aerial orb webs (Blackledge et al. 2005a, b). Furthermore, I am studying how spiders can
manipulate the mechanical properties of silk in behavioral contexts such as
responding to different types of prey or constructing different types of
webs. This research is part of two NSF funded projects:
"Araneid phylogeny and evolution of spider
silk phenotypes" and
"Acquisition of a dynamic
nano-force tensile test system for ultrathin fibers with environmental
control and integrated image analysis" in collaboration with
Nikolaj Scharff
and John Wenzel.
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Evolution of prey specialization in spiders
The Araneidae is the largest family of spiders that construct
wheel-shaped two-dimensional orb webs. Yet, many araneids have evolved
highly specialized web construction behaviors that result in webs unlike
typical orb webs. I am currently working to establish a generic level
phylogeny for relationships within this group to investigate how and why
these behavioral specializations have evolved, how they contribute to the
dominance of araneid spiders as predators of flying insects, and how the
evolution of behavior and silk biomechanics interact with one another. This
project includes molecular data from multiple nuclear and mitochondrial
loci, as well as morphological data in collaboration with
Dr. Nikolaj
Scharff and Mark Stowe.
The extreme web
reduction found in bolas spiders is of particular interest to me. Bolas
spiders have reduced the orb web to a single sticky thread, which they throw
at moth prey. These moths are attracted through aggressive chemical mimicry
of female moth sex pheromones. Our phylogenetic research is currently testing
the hypothesis that this unique hunting behavior evolved through a series of
web reductions in a number of closely related species of spiders that
display lesser degrees of specialization upon moths. Once we have established
an historical framework for the evolution of web building behaviors within
these moth specialists, we will then examine changes in the functional
properties of the unique capture silks used by these spiders. This will
provide insight into how evolution has acted upon the genes coding for silks
during extreme behavioral specialization.

Behavioral innovation and biodiversity:
adaptive radiations in Hawai'i
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 Adaptive
radiation is a major theme in the evolution of the world’s biodiversity, yet
in non-adaptive radiations, speciation occurs with minor
change in the ecologies of species. Little is known about whether lineages
in adaptive radiations exploit more resources than non-adaptive radiations,
divide resources more finely, or both? Furthermore, does adaptive radiation
result in predictable evolutionary patterns of resource exploitation, or is
historical contingency so great that, despite selective pressures in common
between communities, evolution of resource use within radiations is largely
unpredictable? I have found that within an endemic radiation of Hawaiian
spiders, sympatric species of Tetragnatha display an extraordinary
diversification in how and where they construct webs (Blackledge, Binford &
Gillespie 2003). Remarkably similar web building behaviors, or “ethotypes”,
have evolved independently in groups of species on different islands,
suggesting a deterministic pattern to the behavioral diversification of
endemic Hawaiian Tetragnatha (Blackledge & Gillespie, 2004).
Future research will examine how these web ethotypes function in prey
capture, to test whether convergence in web architecture is related to
convergent selective pressures for prey capture behavior (e.g. Blackledge,
Hang & Gillespie, in prep). Furthermore, I will compare the biomechanical
properties of the capture silks used to construct these diverse webs, as a
framework to study how silk gene evolution is related to behavioral
diversification within this adaptive radiation.
Cylcosa (Araneidae)
is the only other group of orb-weaving spiders, besides Tetragnatha (Tetragnathidae),
to have dispersed to and speciated across the isolated Hawaiian Islands. In
contrast to Tetragnatha, Cyclosa comprises a relatively
species poor radiation. These two groups of spiders provide a unique
opportunity to study why speciation is higher in one lineage than in
another. Both genera are sympatric, found within the same habitats, but are
segregated temporally because Cyclosa is diurnal while Tetragnatha
is nocturnal. Thus, competition does not explain this difference in
diversification. Preliminary data also suggest that the Hawaiian Cyclosa
are not especially younger than Hawaiian Tetragnatha. Instead,
comparison of resource use between these two radiations may give insight
into the causes of adaptive radiation. By quantifying variation in the web
architectures and resource use of endemic Hawaiian spiders and
reconstructing their phylogenetic relationships I can answer two hypotheses
about diversification. Do species diversify in adaptive radiations because
they exploit greater ecological opportunity? Or, do species divide resources
more narrowly, or tolerate greater niche overlap, so that adaptive
radiations can occur regardless of ecological opportunity?
Learn more about Hawaiian spiders at Rosemary
Gillespie's website.
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Tetragnatha hawaiensis web from Hawai'i |

Tetragnatha stelarobusta from Maui |

Cyclosa simplicicauda rufescens from Hawai'i |

Predator-prey conflict and sensory drive: stabilimentum web decorations
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My research focuses on
the conflict that arises, between signaling presence of webs to predators
and to prey, when spiders include stabilimenta in their webs. These
conspicuous silk lines, crosses and spirals may have several defensive
functions including camouflage of spiders, startling predators, and acting
as aposematic warnings for the presence of webs (Blackledge & Wenzel 1999,
2001). However, my research indicates that insect prey can also use
stabilimenta as a signal in avoidance of webs, indicating that there should
be selection against the use of stabilimenta in web avoidance by insects
(Blackledge & Wenzel 1999). The reflectance spectrum of the silk used to
build stabilimenta suggests that the silk is cryptic to insects, unlike more
primitive spider silks (Blackledge 1998b). This is supported by my
experiments demonstrating that honey bees can learn to forage at targets
made from primitive spider silks but not targets made from stabilimentum
silk (Blackledge & Wenzel 2000). I suggest that the evolution of silk
coloration has occurred through a process termed sensory drive, where innate
biases in the color vision of insects has selected for the cryptic
properties of stabilimenta. This system is unusual because most examples of
sensory drive involve sexually selected signals but spiders’ silks have
evolved under natural selection from predators and prey. |
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Argiope aurantia is common around Akron |

immature Argiope
often construct doily-like designs |

mud-dauber wasps are
important predators of spiders |
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