Insect Societies Outline

Insect of the Day: Myrmelachista ants
Insect Societies
Outline
•
•
Subsociality in insects
Eusociality in insects
Outline
• Inquilines & parasites of
social insects
• Evolution & maintenance
of eusociality
• Success of eusocial insects
Numerical dominance
of social insects
• Formica yessensis in 2.7 km
• 306 million workers
• 1 million queens
• 45,000 interconnected nests
2
“Siafu”
African Driver ant
(genus Dorylus)
Ant Colony Optimization
algorithms (ACOs)
Eusociality (“true social”)
• Division of labor with a caste system composed
of reproductive and non-reproductive individuals
• Cooperation in raising the young
• Overlapping generations
Eusociality (“true social”)
Subbelow
• Division of labor with a caste system
composed of reproductive and nonAll females are
reproductive individuals
able to lay eggs
Cooperation in raising the young
•
• Overlapping generations
Parental care with nesting
aggro aphid
Flavors of subsociality
behavior: communal nest with
• Quasisocial
females of same generation that care for
young and are all able to reproduce
behavior: communal nest with
• Semisocial
females of same generation that care for
young and are all able to reproduce - but
some act as workers and rarely lay eggs.
Quasisocial millipedes
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Insect of the day: Large Blue,
Phengaris arion
Outline
& parasites
• Inquilines
of social insects
&
• Evolution
maintenance of
eusociality
of eusocial
• Success
insects
Eusociality
•
Division of labor with a
caste system composed
of reproductive and nonreproductive individuals
•
Cooperation in raising
the young
•
Overlapping generations
Bald-faced hornet,
Dolichovespula maculata
Haplodiploidy
• Queens control the sex of their offspring
• Haplodiploid genetic system
• Fertilized haploid eggs develop into females
• Unfertilized haploid eggs develop into males
♀
Primitive eusociality &
nest founding
•
Nest founded by more
than one queen (gyne)
•
Rapidly goes from
polygynous to
monogynous
•
One queen establishes
dominance hierarchy
♂
Age polyethism
poly = many ethos = nature
• Individuals of eusocial insects in a caste
differ behaviorally and perform different
tasks as they age.
• Vespine wasps:
-
food distribution > cell cleaning > ventilation/wingfanning > nest defense > foraging > construction &
repair
Genomic-Based Phylogeny of Ants, Bees, and Wasps
2061
Figure 3. Evolution of the Aculeate Hymenoptera
Blue-green branches represent parasitoidism;
orange branches represent nest construction
and predation (with pollenivory and omnivory as
derivative states thereof). Asterisks designate lineages containing eusocial species. Ants are
entirely eusocial, but this is not true of all species
of Vespidae and Apoidea. Biological information
is from various sources, summarized in Gauld
and Bolton [2] and Huber [13]. Names of superfamilies are modified from Pilgrim et al. [8]. Placement of Rhopalosomatidae is based on Pilgrim
et al. [8] and Debevec et al. [11]. Images courtesy
of Alexander Wild and Kurt Schaefer.
2. Gauld, I.D., and Bolton, B., eds. (1988). The Hymenoptera (London:
with CAT-GTR as the amino acid replacement model, in an unpartitioned
Oxford University Press).
analysis of the 308-gene matrix. A species tree was estimated on the basis
of average ranks of gene coalescence events, as calculated in STAR [22].
3. Hölldobler, B., and Wilson, E.O. (1990). The Ants (Cambridge: Harvard
The input for this analysis was 100 bootstrap replicate trees of each of
University Press).
308 genes, built under maximum likelihood in RAxML. We also inferred a
4. Bradley, T.J., Briscoe, A.D., Brady, S.G., Contreras, H.L., Danforth, B.N.,
species tree with PhyloNet [33], which uses the parsimony-based criterion
Dudley, R., Grimaldi, D., Harrison, J.F., Kaiser, J.A., Merlin, C., et al.
of minimizing deep coalescences [34]. We used 308 input trees with boot(2009). Episodes in insect evolution. Integr. Comp. Biol. 49, 590–606.
strap support values generated in RAxML.
5. Hughes, W.O.H., Oldroyd, B.P., Beekman, M., and Ratnieks, F.L.W.
To evaluate alternate phylogenetic hypotheses against our best-scoring
(2008). Ancestral monogamy shows kin selection is key to the evolution
ML tree, we employed the Shimodaira-Hasegawa test [23]. Five constraints
of eusociality. Science 320, 1213–1216.
were considered (Table S3), and separate constrained partitioned analyses
6. Ronquist, F. (1999). Phylogeny of the Hymenoptera (Insecta): the state
were conducted using RAxML on the same 308-gene matrix used to
of the art. Zool. Scr. 28, 3–12.
generate our ML tree (Figure 2). The five best trees satisfying the respective
7. Brothers, D.J. (1999). Phylogeny and evolution of wasps, ants and bees
constraints were then subjected to the Shimodaira-Hasegawa test in
(Hymenoptera, Chrysidoidea, Vespoidea and Apoidea). Zool. Scr. 28,
RAxML (‘‘-f h’’ option) to determine whether they were significantly
worse
Genomic-Based Phylogeny of Ants,233–249.
Bees, and Wasps
2061
than our best unconstrained ML tree.
8. Pilgrim, E.M., von Dohlen, C.D., and Pitts, J.P. (2008). Molecular phylogenetics of Vespoidea indicate paraphyly of the superfamily and novel
Accession Numbers
relationships of its component families
and subfamilies. Zool. Scr. 37,
Figure 3. Evolution of the Aculeate Hymenoptera
539–560.
Blue-green branches represent parasitoidism;
Illumina reads have been deposited in the NCBI Sequence Read Archive
orange branches represent nest construction
9. Heraty, J., Ronquist, F., Carpenter, J.M.,
Hawks,
D., Schulmeister,
S.,
and predation
(with pollenivory
and omnivory as
with the accession number SRP020476. The matrices, partition files, and
derivative states thereof). Asterisks designate linDowling,
A.P.,
Murray,
D.,
Munro,
J.,
Wheeler,
W.C., Schiff, N., and
eages
containing eusocial species. Ants are
gene trees have been deposited in Dryad (http://doi.org/10.5061/dryad.
entirely
eusocial, but this is not true
of all species
Sharkey, M. (2011). Evolution of the
hymenopteran
megaradiation.
jt440).
of Vespidae and Apoidea. Biological information
Mol. Phylogenet. Evol. 60, 73–88.
is from various sources, summarized in Gauld
and Bolton [2] and Huber [13]. Names of super10. Sharkey, M.J., Carpenter, J.M., Vilhelmsen,
L., Heraty,
J.,et al.Liljeblad,
J.,
families are modified
from Pilgrim
[8]. PlaceSupplemental Information
ment of Rhopalosomatidae is based on Pilgrim
Dowling, A.P.G., Schulmeister, S., Murray,
D., Deans, A.R., Ronquist,
et al. [8] and Debevec et al. [11]. Images courtesy
of Alexander Wild
and Kurt Schaefer.
F., et al. (2012). Phylogenetic relationships
among
superfamilies of
Supplemental Information includes one figure, three tables, and SuppleHymenoptera. Cladistics 28, 80–112.
mental Experimental Procedures and can be found with this article online
11. Debevec, A.H., Cardinal, S., and Danforth, B.N. (2012). Identifying the
at http://dx.doi.org/10.1016/j.cub.2013.08.050.
sister group to the bees: a molecular phylogeny of Aculeata with an
emphasis on the superfamily Apoidea. Zool. Scr. 41, 527–535.
Acknowledgments
12. Wilson, J.S., von Dohlen, C.D., Forister, M.L., and Pitts, J.P. (2013).
Family-level divergences in the stinging wasps (Hymenoptera:
This work was funded by the University of California, Davis. We thank James
Aculeata), with correlations to angiosperm diversification. Evol. Biol.
Pitts for provision of the Apterogyna (Bradynobaenidae) specimens and
40, 101–107.
three anonymous reviewers for helpful comments thatwithimproved
the
2. Gauld, I.D., and Bolton, B., eds. (1988). The Hymenoptera (London:
CAT-GTR as the amino acid replacement model, in an unpartitioned
13.
Huber,
J.T. (2009).
Biodiversity
of Hymenoptera.
In Insect Biodiversity:
Oxford University
Press).
analysis
of
the
308-gene
matrix.
A
species
tree was estimated
on the basis
manuscript.
of average ranks of gene coalescence events, as calculated in STAR [22].
3. Hölldobler, B., and Wilson, E.O. (1990). The Ants (Cambridge: Harvard
Science
R.
Footit and P. Adler, eds. (Oxford: WileyThe input for this analysis was 100 bootstrap
replicateand
trees ofSociety,
each of
University Press).
308 genes, built under maximum likelihood
in RAxML. We also
a
4. Bradley, T.J., Briscoe, A.D., Brady, S.G., Contreras, H.L., Danforth, B.N.,
Blackwell),
pp.inferred
303–323.
species tree with PhyloNet [33], which uses the parsimony-based criterion
Received: June 26, 2013
Dudley, R., Grimaldi, D., Harrison, J.F., Kaiser, J.A., Merlin, C., et al.
of minimizing deep coalescences 14.
[34]. We
used 308 input
with boot- J., Akhter,
(2009). Episodes
insect evolution. Integr.
Comp.and
Biol. 49,Shultz,
590–606. J.W.
Hedin,
M.,trees
Starrett,
S.,inSchönhofer,
A.L.,
Revised: August 1, 2013
strap support values generated in RAxML.
5. Hughes, W.O.H., Oldroyd, B.P., Beekman, M., and Ratnieks, F.L.W.
(2012).
Phylogenomic
resolution
ofmonogamy
paleozoic
harvestTo evaluate alternate phylogenetic hypotheses
against
our best-scoring
(2008). Ancestral
shows kindivergences
selection is key to thein
evolution
Accepted: August 21, 2013
ML tree, we employed the Shimodaira-Hasegawa test [23]. Five constraints
of eusociality.
Science 320, 1213–1216.
(Arachnida,
Opiliones)
via analysis
of next-generation transcripwere considered (Table S3), and separatemen
constrained
partitioned analyses
Published: October 3, 2013
6. Ronquist, F. (1999). Phylogeny of the Hymenoptera (Insecta): the state
were conducted using RAxML on the tome
same 308-gene
usedONE
to
data.matrix
PLoS
7,ofe42888.
the art. Zool. Scr. 28, 3–12.
generate our ML tree (Figure 2). The five best trees satisfying the respective
7. Brothers, D.J. (1999). Phylogeny and evolution of wasps, ants and bees
constraints were then subjected15.
to the
Shimodaira-Hasegawa
test
in
Smith, S.A., Wilson, N.G.,
Goetz,Chrysidoidea,
F.E., Feehery,
C.,
Andrade,
(Hymenoptera,
Vespoidea and
Apoidea).
Zool. Scr. S.C.S.,
28,
References
RAxML (‘‘-f h’’ option) to determine whether they were significantly worse
233–249.
than our best unconstrained ML tree. Rouse, G.W., Giribet, G., and Dunn, C.W. (2011). Resolving the evolu8. Pilgrim, E.M., von Dohlen, C.D., and Pitts, J.P. (2008). Molecular phylo1. Wilson, E.O. (1971). The Insect Societies (Cambridge:
Harvard
tionary relationships of molluscs
with indicate
phylogenomic
tools. Nature
genetics of Vespoidea
paraphyly of the superfamily
and novel 480,
Accession Numbers
relationships of its component families and subfamilies. Zool. Scr. 37,
University Press).
364–367.
539–560.
Johnson, et al. 2013. Current Biology
Illumina reads have been deposited in the NCBI Sequence Read Archive
with the accession number SRP020476. The matrices, partition files, and
gene trees have been deposited in Dryad (http://doi.org/10.5061/dryad.
jt440).
Supplemental Information
Supplemental Information includes one figure, three tables, and Supplemental Experimental Procedures and can be found with this article online
at http://dx.doi.org/10.1016/j.cub.2013.08.050.
9. Heraty, J., Ronquist, F., Carpenter, J.M., Hawks, D., Schulmeister, S.,
Dowling, A.P., Murray, D., Munro, J., Wheeler, W.C., Schiff, N., and
Sharkey, M. (2011). Evolution of the hymenopteran megaradiation.
Mol. Phylogenet. Evol. 60, 73–88.
10. Sharkey, M.J., Carpenter, J.M., Vilhelmsen, L., Heraty, J., Liljeblad, J.,
Dowling, A.P.G., Schulmeister, S., Murray, D., Deans, A.R., Ronquist,
F., et al. (2012). Phylogenetic relationships among superfamilies of
Hymenoptera. Cladistics 28, 80–112.
11. Debevec, A.H., Cardinal, S., and Danforth, B.N. (2012). Identifying the
sister group to the bees: a molecular phylogeny of Aculeata with an
Kin selection
•
Why should some
individuals sacrifice their
reproduction for others?
•
•
Kin selection explanation
•
Contribution by a related
individual that assists
with the reproductive
success of its kin (but not
its own)
Darwinian fitness plus
kinship component
no caving, unless you’re kinfolk
Hamilton’s rule
•
•
Inclusive fitness
•
r relatedness, B altruism’s
benefits for recipient, C
altruism’s cost by donor
W.D. Hamilton suggested
that altruism is promoted
by: rB - C > 0
reduction of an organism’s
(parent’s) inclusive fitness
Photo credits
•
Formica yessensis, Gakken’s photo
encyclopedia ants,
www.ant.edb.miyakyo-u.ac.jp/
INTRODUCTION/Gakken79E/
Page_35.html
•
Ruby spotted swallowtail aggregation,
Jessika Canizalez,
www.whatsthatbug.com/2010/11/08/
unknown-caterpillar-aggregation-frommexico/
•
Morpho telemachus caterpillar
aggregation, Rich Hoyer,
www.birdingblogs.com/2011/
richhoyer/cristalino-montage-–-row-6/
morpho-telemachus-caterpillar
•
Ruby spotted swallowtail adult, Andy
Warren,
www.butterfliesofamerica.com/images/
Papilionidae/Papilioninae/
Papilio_anchisiades_idaeus/
Papilio_anchisiades_idaeus_F_MX_N
AY_Jumatan_360m_28-IX-1996-fd.jpg
•
European earwig, Nabokov,
www.entnemdept.ufl.edu/creatures/
veg/european_earwig.htm
•
Tarantula hawk & tarantula, Robyn
Waayer, www.sdrp.org/resources/
Ecology/Robyn%20Waayer/
GIIBA7tarantulahawkandtar.jpg
•
Tarantula hawk & tarantula,
Astrobradley,
www.commons.wikimedia.org/wiki/
File:Tarantula_hawk_cropped.JPG