Sperm competition and the coevolution of pre- and postcopulatory traits: Weapons evolve faster than testes among onthophagine dung beetles

Reproductive competition generates episodes of both pre- and postcopulatory sexual selection. Theoretical models of sperm competition predict that as the fitness gains from expenditure on the weapons of male combat increase, males should increase their expenditure on weapons and decrease their expen...

Full description

Saved in:
Bibliographic Details
Published inEvolution Vol. 70; no. 5; pp. 998 - 1008
Main Authors Simmons, Leigh W., Fitzpatrick, John L.
Format Journal Article
LanguageEnglish
Published United States Blackwell Publishing Ltd 01.05.2016
Society for the Study of Evolution
Oxford University Press
Subjects
Online AccessGet full text

Cover

Loading…
Abstract Reproductive competition generates episodes of both pre- and postcopulatory sexual selection. Theoretical models of sperm competition predict that as the fitness gains from expenditure on the weapons of male combat increase, males should increase their expenditure on weapons and decrease their expenditure on traits that contribute to competitive fertilization success. Although traits subject to sexual selection are known to have accelerated evolutionary rates of phenotypic divergence, it is not known whether the competing demands of investment into pre- and postcopulatory traits affect their relative rates of evolutionary divergence. We use a comparative approach to estimate the rates of divergence in pre- and postcopulatory traits among onthophagine dung beetles. Weapons evolved faster than body size while testes mass and sperm length evolved more slowly than body size, suggesting that precopulatory competition is the stronger episode of sexual selection acting on these beetles. Although horns evolved faster than testes, evolutionary increases in horn length were not associated with evolutionary reductions in testes mass. Our data for onthophagines support the notion that in taxa where males are unable to monopolize paternity, expenditure on both weapons and testes should both be favored.
AbstractList Reproductive competition generates episodes of both pre- and postcopulatory sexual selection. Theoretical models of sperm competition predict that as the fitness gains from expenditure on the weapons of male combat increase, males should increase their expenditure on weapons and decrease their expenditure on traits that contribute to competitive fertilization success. Although traits subject to sexual selection are known to have accelerated evolutionary rates of phenotypic divergence, it is not known whether the competing demands of investment into pre- and postcopulatory traits affect their relative rates of evolutionary divergence. We use a comparative approach to estimate the rates of divergence in pre-and postcopulatory traits among onthophagine dung beetles. Weapons evolved faster than body size while testes mass and sperm length evolved more slowly than body size, suggesting that precopulatory competition is the stronger episode of sexual selection acting on these beetles. Although horns evolved faster than testes, evolutionary increases in horn length were not associated with evolutionary reductions in testes mass. Our data for onthophagines support the notion that in taxa where males are unable to monopolize paternity, expenditure on both weapons and testes should both be favored.
Author Fitzpatrick, John L.
Simmons, Leigh W.
Author_xml – sequence: 1
  givenname: Leigh W.
  surname: Simmons
  fullname: Simmons, Leigh W.
  email: leigh.simmons@uwa.edu.au
  organization: Centre for Evolutionary Biology, School of Animal Biology (M092), The University of Western Australia, WA, 6009, Crawley, Australia
– sequence: 2
  givenname: John L.
  surname: Fitzpatrick
  fullname: Fitzpatrick, John L.
  organization: Department of Zoology/Ethology, Stockholm University, Stockholm, Sweden
BackLink https://www.ncbi.nlm.nih.gov/pubmed/27061413$$D View this record in MEDLINE/PubMed
https://urn.kb.se/resolve?urn=urn:nbn:se:su:diva-132514$$DView record from Swedish Publication Index
BookMark eNqFks1u1DAUhS1URKeFBQ8AssSGRdPa8W_YVf0DqdBFS7u0nMydGQ9JHGyn7bwCT407UwaJDd7Yuue7R746dw_t9L4HhN5SckjzOYJ7f0jLiooXaEKF0IWQXO6gCSGUF0yXZBftxbgkhFSCVq_QbqmIpJyyCfp1PUDocOO7AZJLzvfY9lOcFpBr2bcd1zU_w0OAYq0NPqbGD2Nrkw8rnIJ1KX7Cd2AH30f81HQPeGZjgpCNbI8T5HfEtvP9HPs-LfywsHPXA56OuVIDpBbia_RyZtsIb57vffT9_Ozm5HNxeXXx5eT4snBcCVGwsm6UmtXaghTEWlo20JRTC9xqrkCWXNBaCiGUrhotOWG8qqGccmgoKAZsHx1sfOMDDGNthuA6G1bGW2dO3e2x8WFu4mgoKwXlGf-4wYfgf455EtO52EDb2h78GA3VREtZUUX_j6qKcMkIJRn98A-69GPo89yZ0lWVIc0y9f6ZGusOptuv_skvA0cb4MG1sNrqlJinxTA5C7NeDHN2e7V-5I53m45lzPH9deRKllTLrBcb3eXMHre6DT-MVEwJc_ftwrDr06_nNyU1mv0GpfnGcA
ContentType Journal Article
Copyright Copyright © 2016 Society for the Study of Evolution
2016 The Author(s). © 2016 The Society for the Study of Evolution.
2016 The Author(s). Evolution © 2016 The Society for the Study of Evolution.
Copyright © 2016, Society for the Study of Evolution
Copyright_xml – notice: Copyright © 2016 Society for the Study of Evolution
– notice: 2016 The Author(s). © 2016 The Society for the Study of Evolution.
– notice: 2016 The Author(s). Evolution © 2016 The Society for the Study of Evolution.
– notice: Copyright © 2016, Society for the Study of Evolution
DBID BSCLL
CGR
CUY
CVF
ECM
EIF
NPM
7QG
7QL
7QP
7QR
7SN
7SS
7TK
7TM
7U9
8FD
C1K
FR3
H94
M7N
P64
RC3
7X8
ADTPV
AOWAS
DG7
DOI 10.1111/evo.12915
DatabaseName Istex
Medline
MEDLINE
MEDLINE (Ovid)
MEDLINE
MEDLINE
PubMed
Animal Behavior Abstracts
Bacteriology Abstracts (Microbiology B)
Calcium & Calcified Tissue Abstracts
Chemoreception Abstracts
Ecology Abstracts
Entomology Abstracts (Full archive)
Neurosciences Abstracts
Nucleic Acids Abstracts
Virology and AIDS Abstracts
Technology Research Database
Environmental Sciences and Pollution Management
Engineering Research Database
AIDS and Cancer Research Abstracts
Algology Mycology and Protozoology Abstracts (Microbiology C)
Biotechnology and BioEngineering Abstracts
Genetics Abstracts
MEDLINE - Academic
SwePub
SwePub Articles
SWEPUB Stockholms universitet
DatabaseTitle MEDLINE
Medline Complete
MEDLINE with Full Text
PubMed
MEDLINE (Ovid)
Virology and AIDS Abstracts
Technology Research Database
Nucleic Acids Abstracts
Ecology Abstracts
Neurosciences Abstracts
Biotechnology and BioEngineering Abstracts
Environmental Sciences and Pollution Management
Entomology Abstracts
Genetics Abstracts
Animal Behavior Abstracts
Bacteriology Abstracts (Microbiology B)
Algology Mycology and Protozoology Abstracts (Microbiology C)
AIDS and Cancer Research Abstracts
Chemoreception Abstracts
Engineering Research Database
Calcium & Calcified Tissue Abstracts
MEDLINE - Academic
DatabaseTitleList
Virology and AIDS Abstracts

Entomology Abstracts

MEDLINE - Academic
MEDLINE
Database_xml – sequence: 1
  dbid: NPM
  name: PubMed
  url: https://proxy.k.utb.cz/login?url=http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed
  sourceTypes: Index Database
– sequence: 2
  dbid: EIF
  name: MEDLINE
  url: https://proxy.k.utb.cz/login?url=https://www.webofscience.com/wos/medline/basic-search
  sourceTypes: Index Database
DeliveryMethod fulltext_linktorsrc
Discipline Biology
EISSN 1558-5646
EndPage 1008
ExternalDocumentID oai_DiVA_org_su_132514
4061409271
27061413
EVO12915
24762186
ark_67375_WNG_3SDMFT21_8
Genre article
Research Support, Non-U.S. Gov't
Journal Article
GroupedDBID ---
--Z
-JH
-~X
.3N
.GA
.Y3
05W
0R~
10A
1OC
29G
2AX
31~
33P
3O-
3SF
4.4
41~
42X
50Y
50Z
51W
51X
52M
52N
52O
52P
52S
52T
52U
52W
52X
53G
5GY
5HH
5LA
5VS
5WD
66C
702
79B
7PT
8-0
8-1
8-3
8-4
8-5
85S
8UM
930
A03
AAESR
AAEVG
AAHBH
AAHHS
AAHKG
AAISJ
AAKGQ
AAONW
AAPSS
AAPXW
AARHZ
AASGY
AAUAY
AAVAP
AAWDT
AAXRX
AAZKR
ABBHK
ABCQN
ABCUV
ABDPE
ABEJV
ABEML
ABJNI
ABLJU
ABMNT
ABPLY
ABPPZ
ABPTD
ABPVW
ABTLG
ABWJO
ABXSQ
ABXVV
ACAHQ
ACBWZ
ACCFJ
ACCZN
ACFBH
ACFRR
ACGFO
ACGFS
ACGOD
ACIPB
ACIWK
ACKIV
ACNCT
ACPOU
ACPRK
ACSCC
ACSTJ
ACUFI
ACUTJ
ACXBN
ACXQS
ACZBC
ADACV
ADBBV
ADEOM
ADHSS
ADIPN
ADIZJ
ADKYN
ADMGS
ADOZA
ADQBN
ADULT
ADXAS
ADZMN
ADZOD
AEEZP
AEGXH
AEIMD
AENEX
AEPYG
AEQDE
AEUPB
AEUQT
AFAZZ
AFBPY
AFFDN
AFFIJ
AFGKR
AFGWE
AFNWH
AFPWT
AFRAH
AFYAG
AFZJQ
AGMDO
AGUYK
AHXOZ
AI.
AIAGR
AILXY
AIURR
AIWBW
AJAOE
AJBDE
AJXKR
AKPMI
ALAGY
ALMA_UNASSIGNED_HOLDINGS
ALUQN
AMBMR
AMYDB
ANFBD
AQVQM
ASPBG
AS~
ATGXG
ATUGU
AUFTA
AVWKF
AZBYB
AZFZN
AZVAB
BAFTC
BCRHZ
BDRZF
BHBCM
BKOMP
BMNLL
BMXJE
BNHUX
BROTX
BRXPI
BSCLL
BY8
CAG
CBGCD
COF
CS3
CUYZI
D-E
D-F
D0L
D0S
DC7
DCZOG
DEVKO
DOOOF
DPXWK
DR2
DRFUL
DRSTM
DU5
EBS
ECGQY
EJD
ESX
F00
F01
F04
F5P
FAC
FAL
FAS
FD6
FEDTE
FJD
FJW
G-S
G.N
GODZA
GTFYD
H.T
H.X
H13
HF~
HGD
HQ2
HTVGU
HVGLF
HZ~
IAG
IAO
IEA
IEP
IOF
IPSME
ITC
IX1
J0M
JAAYA
JBMMH
JBS
JEB
JENOY
JHFFW
JKQEH
JLS
JLXEF
JPM
JSODD
JST
K48
KOP
LATKE
LC2
LC3
LEEKS
LH4
LITHE
LOXES
LP6
LP7
LUTES
LW6
LYRES
MK4
MRFUL
MRSTM
MSFUL
MSSTM
MVM
MXFUL
MXSTM
N04
N05
N9A
NEJ
NF~
NHB
NQS
O66
O9-
OBOKY
OIG
OJZSN
OK1
OVD
OWPYF
P-O
P2P
P2W
P2X
P4D
PQ0
PQQKQ
Q.N
Q11
Q5J
QB0
QN7
R.K
RBO
ROL
ROX
RWL
RX1
RXW
SA0
SJN
SUPJJ
TAE
TCN
TEORI
TN5
UB1
UBC
UHB
UQL
V8K
VH1
VJK
VQA
W8V
W99
WBKPD
WH7
WHG
WIH
WIK
WNSPC
WOHZO
WQJ
WRC
WYISQ
XG1
XOL
XSW
YXE
YYP
YZZ
ZCA
ZCG
ZZTAW
~02
~IA
~KM
~WT
1OB
AAMMB
ABDFA
ABGNP
ABSQW
ABXZS
ACHIC
ACSIT
ACYXJ
ADGKP
AEFGJ
AGORE
AGQPQ
AGXDD
AIDQK
AIDYY
AJBYB
AJNCP
NU-
AANHP
ACRPL
ADNMO
CGR
CUY
CVF
ECM
EIF
ISM
NPM
YIF
YIN
7QG
7QL
7QP
7QR
7SN
7SS
7TK
7TM
7U9
8FD
C1K
FR3
H94
M7N
P64
RC3
7X8
ABIME
ABPIB
ABZEO
ACVCV
ADTPV
ADXHL
AFKWF
AHGBF
AJDVS
AOWAS
APJGH
DG7
ID FETCH-LOGICAL-i4755-32bc77fb8ae650aa12cec2dae4a847e62451b6555789c8640349be2d4ec1e73e3
IEDL.DBID DR2
ISSN 0014-3820
1558-5646
IngestDate Thu Aug 21 06:50:47 EDT 2025
Fri Jul 11 15:04:55 EDT 2025
Fri Jul 11 16:11:47 EDT 2025
Fri Jul 25 10:41:43 EDT 2025
Wed Feb 19 02:40:28 EST 2025
Wed Jan 22 17:07:57 EST 2025
Sun Aug 24 12:10:44 EDT 2025
Wed Oct 30 09:51:47 EDT 2024
IsPeerReviewed true
IsScholarly true
Issue 5
Keywords evolutionary divergence
Ejaculate expenditure
male contest competition
horns
life-history trade-off
sperm length
Language English
License 2016 The Author(s). Evolution © 2016 The Society for the Study of Evolution.
LinkModel DirectLink
MergedId FETCHMERGED-LOGICAL-i4755-32bc77fb8ae650aa12cec2dae4a847e62451b6555789c8640349be2d4ec1e73e3
Notes istex:18299324B6C56A4843C48D8B5369E49E98A6E107
ArticleID:EVO12915
ark:/67375/WNG-3SDMFT21-8
ObjectType-Article-1
SourceType-Scholarly Journals-1
ObjectType-Feature-2
content type line 14
content type line 23
PMID 27061413
PQID 1789930183
PQPubID 42232
PageCount 11
ParticipantIDs swepub_primary_oai_DiVA_org_su_132514
proquest_miscellaneous_1808669171
proquest_miscellaneous_1790463010
proquest_journals_1789930183
pubmed_primary_27061413
wiley_primary_10_1111_evo_12915_EVO12915
jstor_primary_24762186
istex_primary_ark_67375_WNG_3SDMFT21_8
PublicationCentury 2000
PublicationDate May 2016
PublicationDateYYYYMMDD 2016-05-01
PublicationDate_xml – month: 05
  year: 2016
  text: May 2016
PublicationDecade 2010
PublicationPlace United States
PublicationPlace_xml – name: United States
– name: St. Louis
PublicationTitle Evolution
PublicationTitleAlternate Evolution
PublicationYear 2016
Publisher Blackwell Publishing Ltd
Society for the Study of Evolution
Oxford University Press
Publisher_xml – name: Blackwell Publishing Ltd
– name: Society for the Study of Evolution
– name: Oxford University Press
References Drummond, A. J., M. A. Suchard, D. Xie, and A. Rambaut. 2012. Baysesian phylogenetics with BEAUti and the BEAST 1.7. Mol. Biol. Evol. 29:1969-1973.
García-González, F., and L. W. Simmons. 2007. Shorter sperm confer higher competitive fertilization success. Evolution 61:816-824.
Tomkins, J. L., and L. W. Simmons. 2000. Sperm competition games played by dimorphic male beetles: fertilisation gains with equal mating access. Proc. R. Soc. Lond. B 267:1547-1553.
Mendelson, T. C., and K. L. Shaw. 2005. Rapid speciation in an arthropod. Nature 433:375-376.
Houle, D. 1992. Comparing evolvability and variability of quantitative traits. Genetics 130:195-204.
Wilkinson, G. S., and M. Taper. 1999. Evolution of genetic variation for condition-dependent traits in stalk-eyed flies. Proc. R. Soc. Lond. B 266:1685-1690.
Parker, G. A. 1970. Sperm competition and its evolutionary consequences in the insects. Biol. Rev. 45:525-567.
Hunt, J.. 2001. Status-dependent selection in the dimorphic beetle Onthophagus taurus. Proc. R. Soc. Lond. B 268:2409-2414.
Maddison, W. P., and D. R. Maddison. 1999. MacClade: analysis of phylogeny and character evolution. Version 3.08a. Sinauer, Sunderland, MA.
Pagel, M. 1999. Inferring the historical patterns of biological evolution. Nature 401:877-884.
Dines, J. P., S. L. Mesnick, K. Ralls, L. May-Collado, I. Agnarsson, and M. D. Dean. 2015. A trade-off between precopulatory and postcopulatory trait investment in male cetaceans. Evolution 69:1560-1572.
Simmons, L. W. 2001. Sperm competition and its evolutionary consequences in the insects. Princeton Univ. Press, Princeton.
Clutton-Brock, T. H., and G. A. Parker. 1992. Potential reproductive rates and the operation of sexual selection. Quart. Rev. Biol. 67:437-456.
Seddon, N., J. A. Botero, J. A. Tobias, P. O. Dunn, H. E. A. MacGregor, D. R. Rubenstein, J. A. C. Uy, J. T. Weir, L. A. Whittingham, and R. J. Safran. 2013. Sexual selection accelerates signal evolution during speciation in birds. Proc. R. Soc. Lond. B 280:20131065.
van Noordwijk, A. J., and G. de Jong. 1986. Acquisition and allocation of resources: their influence on variation in life history tactics. Am. Nat. 128:137-142.
Burnham K. P., and D. R. Anderson. 2002. Model selection and multimodel inference: A practical information-theoretic approach. 2nd ed.Springer-Verlag, New York.
Arnqvist, G. 1998. Comparative evidence for the evolution of genitalia by sexual selection. Nature 393:784-786.
Dunn, J. C., L. B. Halenar, T. G. Davies, J. Cristobal-Azkarate, D. Reby, D. Sykes, S. Dengg, W. T. Fitch, and L. A. Knapp. 2015. Evolutionary trade-off between vocal tract and testes dimensions in howler monkeys. Curr. Biol. 25:2839-2844.
Emlen, D. J., J. Marangelo, B. Ball, and C. W. Cunningham. 2005b. Diversity in the weapons of sexual selection: horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution 59:1060-1084.
Kruuk, L. E. B., J. Slate, J. M. Pemberton, S. Brotherstone, F. Guinness, and T. Clutton-Brock. 2002. Antler size in red deer: heritability and selection but no evolution. Evolution 56:1683-1695.
Kvarnemo, C., and L. W. Simmons. 2013. Polyandry as a mediator of sexual selection before and after mating. Philos. Trans. R Soc B Biol. Sci. 368:20120042.
Simmons, L. W., M. Beveridge, and S. Krauss. 2004. Genetic analysis of parentage within experimental populations of a male dimorphic beetle, Onthophagus taurus, using amplified fragment length polymorphism. Behav. Ecol. Sociobiol. 57:164-173.
Simmons, L. W., and D. J. Emlen. 2006. Evolutionary trade-off between weapons and testes. Proc. Natl. Acad. Sci. USA 103:16349-16351.
Kearse, M., R. Moir, A. Wilson, S. Stones-Havas, M. Cheung, S. Sturrock, S. Buxton, A. Cooper, S. Markowitz, C. Duran, et al. 2012. Geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28:1647-1649.
Price, J. J., and L. M. Whalen. 2009. Plumage evolution in the Oropendolas and Caciques: different divergence rates in polygynous and monogamous taxa. Evolution 63:2985-2998.
Emlen, D. J., J. Hunt, and L. W. Simmons. 2005a. Evolution of sexual dimorphism in the expression of beetle horns: phylogenetic evidence for modularity, evolutionary lability, and constraint. Am. Nat. 166:S42-S68.
Hunt, J., and L. W. Simmons. 1997. Patterns of fluctuating asymmetry in beetle horns: an experimental examination of the honest signalling hypothesis. Behav. Ecol. Sociobiol. 41:109-114.
Cotter, S. C., M. Beveridge, and L. W. Simmons. 2008. Male morph predicts investment in larval immune function in the dung beetle, Onthophagus taurus. Behav. Ecol. 19:331-337.
Emlen, D. J. 1997. Alternative reproductive tactics and male-dimorphism in the horned beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Behav. Ecol. Sociobiol. 41:335-342.
Kokko, H., H. Klug, and M. D. Jennions. 2012. Unifying cornerstones of sexual selection: operational sex ratio, Bateman gradient and the scope for competitive investment. Ecol. Let. 15:1340-1351.
Simmons, L. W., and B. A. Buzatto. 2014. Contrasting responses of pre- and post-copulatory traits to variation in mating competition. Funct. Ecol. 28:494-499.
Moczek, A. P., and D. J. Emlen. 2000. Male horn dimorphism in the scarab beetle Onthophagus taurus: do alternative reproductive tactics favor alternative phenotypes? Anim. Behav. 59:459-466.
Simmons, L. W.. 2007. Quantitative genetic correlation between trait and preference supports a sexually selected sperm process. Proc. Natnl. Acad. Sci. USA 104:16604-16608.
Freckleton, R. P., P. H. Harvey, and M. Pagel. 2002. Phylogenetic analysis and comparative data: a test and review of evidence. Am. Nat. 160:712-726.
Parker, G. A., and T. Pizzari. 2010. Sperm competition and ejaculate economics. Biol. Rev. 85:897-934.
Werner, M., and L. W. Simmons. 2011. Ultrastructure of spermatozoa of Onthophagus taurus (Coleoptera, Scarabaeidae) exhibits heritable variation. Naturewissenschaften 98:213-223.
Parker, G. A., C. M. Lessells, and L. W. Simmons. 2013. Sperm competition games: a general model for pre-copulatory male-male competition. Evolution 67:95-109.
Cooper, N., G.A. Thomas, C. Venditta, A. Meade, and R.P. Freckleton. 2015. A cautionary note on the use of Ornstein Uhlenbeck models in macroevolutionary studies. Biol. J. Linn. Soc. 118:64-77.
Darriba, D., G. L. Taboada, R. Doallo, and D. Posada. 2012. jModelTest 2: more models, new heuristics and parallel computing. Nat. Methods 9:772.
Simmons, L. W., and J. L. Tomkins. 1996. Sexual selection and the allometry of earwig forceps. Evol. Ecol. 10:97-104.
Lüpold, S., J. L. Tomkins, L. W. Simmons, and J. L. Fitzpatrick. 2014. Female monopolization mediates the relationship between pre- and postcopulatory sexual traits. Nat. Comm. 5:3184. DOI: 3110.1038/ncomms4184.
Perry, J. C., and L. Rowe. 2010. Condition-dependent ejaculate size and composition in a ladybird beetle. Proc. R. Soc. Lond. B 277:3639-3648.
Emlen, D. J.. 2008. The evolution of animal weapons. Ann. Rev. Ecol. Evol. Syst. 39:387-413.
Rowe, M., T. Albrecht, E. R. A. Cramer, A. Johnsen, T. Laskemoen, J. T. Weir, and J. T. Lifjeld. 2015. Postcopulatory sexual selection is associated with accelerated evolution of sperm morphology. Evolution 69:1044-1052.
Parker, G. A., and L. W. Simmons. 1996. Parental investment and the control of sexual selection: predicting the direction of sexual competition. Proc. R. Soc. Lond. B 263:315-321.
Simmons, L. W., D. J. Emlen, and J. L. Tomkins. 2007. Sperm competition games between sneaks and guards: a comparative analysis using dimorphic male beetles. Evolution 61:2684-2692.
delBarco-Trillo, J. 2011. Adjustment of sperm allocation under high risk of sperm competition across taxa: a meta-analysis. J. Evol. Biol. 24:1706-1714.
Revell, L. J. 2012. Phytools: an R package for phylogenetic comparative biology (and other things). Methods Ecol. Evol. 3:217-223.
Simmons, L. W., and J. L. Fitzpatrick. 2012. Sperm wars and the evolution of male fertility. Reproduction 144:519-534.
Fromhage, L., and H. Kokko. 2014. Sexually selected traits evolve positive allometry when some matings occur irrespective of the trait. Evolution 68:1332-1338.
Darwin, C. 1871. The descent of man and selection in relation to sex. John Murray, London.
Gonzalez-Voyer, A., and N. Kolm. 2011. Rates of phenotypic evolution of ecological characters and sexual traits during the Tanganyikan cichlid adaptive radiation. J. Evol. Biol. 24:2378-2388.
Fitzpatrick, J. L., M. Almbro, A. Gonzalez-Voyer, N. Kolm, and L. W. Simmons. 2012. Male contest competition and the coevolution of weaponry and testes in pinnipeds. Evolution 66:3595-3604.
Kelly, C. D., and M. D. Jennions. 2011. Sexual selection and sperm quantity: meta-analyses of strategic ejaculation. Biol. Rev. 86:863-884.
Blomberg, S. P., T. Garland, and A. R. Ives. 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57:717-745.
Caro, T. M., C. M. Graham, C. J. Stoner, and M. M. Flores. 2003. Correlates of horn and antler shape in bovids and cervids. Behav. Ecol. Sociobiol. 55:32-41.
Emlen, D. J.. 2001. Costs and the diversification of exaggerated animal structures. Science 291:1534-1536.
Bro-Jørgensen, J. 2007. The intensity of sexual selection predicts weapon size in male bovids. Evolution 61:1316-1326.
Knell, R. J., and L. W. Simmons. 2010. Mating tactics determine patterns of condition dependence in a dimorphic horned beetle. Proc. R. Soc. Lond. B 277:2347-2353.
Harmon, L. J., J. T. Weir, C. D. Brock, R. E. Glor, and W. Challenger. 2008. GEIGER: investigating evolutionary radiations. Bioinformatics 24:129-131.
Amitin, E. G., and S. Pitnick. 2007. Influence of developmental environment on male- and female-mediated sperm precedence in Drosophila melanogaster. J. Evol. Biol. 20:381-391.
Edgar, R. C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acid Res. 32:1792-1797.
Adams, D. C. 2013. Comparing evolutionary rates for different phenoty
2007; 104
1997; 41
2013; 67
2013; 62
2002; 56
2013; 368
2008; 39
2003; 57
2014; 68
2011; 98
1996; 263
2012; 15
2014; 28
2013; 280
2005b; 59
1999; 401
1998; 393
2001; 268
2003; 55
2004; 32
2014; 5
2001
2000; 59
2001; 291
2010; 277
2008; 24
2011; 24
2012; 29
1992; 46
2007; 61
2012; 28
2007; 20
2008; 62
2012; 66
2009; 63
2012; 144
2005; 433
2008; 19
1994
1871
2002
1999; 266
1996; 58
2010; 85
1996; 10
1999
1986; 128
2015; 69
2015; 25
2002; 160
2012; 3
1992; 130
2000; 267
2005a; 166
2004; 58
2004; 57
2011; 86
1970; 45
2015; 118
1992; 67
2006; 103
2012; 9
References_xml – reference: Simmons, L. W., D. J. Emlen, and J. L. Tomkins. 2007. Sperm competition games between sneaks and guards: a comparative analysis using dimorphic male beetles. Evolution 61:2684-2692.
– reference: delBarco-Trillo, J. 2011. Adjustment of sperm allocation under high risk of sperm competition across taxa: a meta-analysis. J. Evol. Biol. 24:1706-1714.
– reference: Simmons, L. W., and G. A. Parker. 1992. Individual variation in sperm competition success of yellow dung flies, Scatophaga stercoraria. Evolution 46:366-375.
– reference: Adams, D. C. 2013. Comparing evolutionary rates for different phenotypic traits on a phylogeny using likelihood. Syst. Biol. 62:181-192.
– reference: Gonzalez-Voyer, A., and N. Kolm. 2011. Rates of phenotypic evolution of ecological characters and sexual traits during the Tanganyikan cichlid adaptive radiation. J. Evol. Biol. 24:2378-2388.
– reference: Fitzpatrick, J. L., M. Almbro, A. Gonzalez-Voyer, N. Kolm, and L. W. Simmons. 2012. Male contest competition and the coevolution of weaponry and testes in pinnipeds. Evolution 66:3595-3604.
– reference: Burnham K. P., and D. R. Anderson. 2002. Model selection and multimodel inference: A practical information-theoretic approach. 2nd ed.Springer-Verlag, New York.
– reference: Blomberg, S. P., T. Garland, and A. R. Ives. 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57:717-745.
– reference: Parker, G. A., C. M. Lessells, and L. W. Simmons. 2013. Sperm competition games: a general model for pre-copulatory male-male competition. Evolution 67:95-109.
– reference: Simmons, L. W., and D. J. Emlen. 2006. Evolutionary trade-off between weapons and testes. Proc. Natl. Acad. Sci. USA 103:16349-16351.
– reference: Perry, J. C., and L. Rowe. 2010. Condition-dependent ejaculate size and composition in a ladybird beetle. Proc. R. Soc. Lond. B 277:3639-3648.
– reference: Edgar, R. C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acid Res. 32:1792-1797.
– reference: Rowe, M., T. Albrecht, E. R. A. Cramer, A. Johnsen, T. Laskemoen, J. T. Weir, and J. T. Lifjeld. 2015. Postcopulatory sexual selection is associated with accelerated evolution of sperm morphology. Evolution 69:1044-1052.
– reference: Bro-Jørgensen, J. 2007. The intensity of sexual selection predicts weapon size in male bovids. Evolution 61:1316-1326.
– reference: Moczek, A. P., and D. J. Emlen. 2000. Male horn dimorphism in the scarab beetle Onthophagus taurus: do alternative reproductive tactics favor alternative phenotypes? Anim. Behav. 59:459-466.
– reference: Parker, G. A. 1970. Sperm competition and its evolutionary consequences in the insects. Biol. Rev. 45:525-567.
– reference: Kruuk, L. E. B., J. Slate, J. M. Pemberton, S. Brotherstone, F. Guinness, and T. Clutton-Brock. 2002. Antler size in red deer: heritability and selection but no evolution. Evolution 56:1683-1695.
– reference: Seddon, N., J. A. Botero, J. A. Tobias, P. O. Dunn, H. E. A. MacGregor, D. R. Rubenstein, J. A. C. Uy, J. T. Weir, L. A. Whittingham, and R. J. Safran. 2013. Sexual selection accelerates signal evolution during speciation in birds. Proc. R. Soc. Lond. B 280:20131065.
– reference: Price, J. J., and L. M. Whalen. 2009. Plumage evolution in the Oropendolas and Caciques: different divergence rates in polygynous and monogamous taxa. Evolution 63:2985-2998.
– reference: Caro, T. M., C. M. Graham, C. J. Stoner, and M. M. Flores. 2003. Correlates of horn and antler shape in bovids and cervids. Behav. Ecol. Sociobiol. 55:32-41.
– reference: Cotton, S., K. Fowler, and A. Pomiankowski. 2004. Condition dependence of sexual ornament size and variation in the stalk-eyed fly Cyrtodiopsis dalmanni (Diptera: Diopsidae). Evolution 58:1038-1046.
– reference: Simmons, L. W., and J. L. Tomkins. 1996. Sexual selection and the allometry of earwig forceps. Evol. Ecol. 10:97-104.
– reference: Parker, G. A., and L. W. Simmons. 1996. Parental investment and the control of sexual selection: predicting the direction of sexual competition. Proc. R. Soc. Lond. B 263:315-321.
– reference: Darwin, C. 1871. The descent of man and selection in relation to sex. John Murray, London.
– reference: Amitin, E. G., and S. Pitnick. 2007. Influence of developmental environment on male- and female-mediated sperm precedence in Drosophila melanogaster. J. Evol. Biol. 20:381-391.
– reference: Cooper, N., G.A. Thomas, C. Venditta, A. Meade, and R.P. Freckleton. 2015. A cautionary note on the use of Ornstein Uhlenbeck models in macroevolutionary studies. Biol. J. Linn. Soc. 118:64-77.
– reference: Cotter, S. C., M. Beveridge, and L. W. Simmons. 2008. Male morph predicts investment in larval immune function in the dung beetle, Onthophagus taurus. Behav. Ecol. 19:331-337.
– reference: Dines, J. P., S. L. Mesnick, K. Ralls, L. May-Collado, I. Agnarsson, and M. D. Dean. 2015. A trade-off between precopulatory and postcopulatory trait investment in male cetaceans. Evolution 69:1560-1572.
– reference: Emlen, D. J., J. Marangelo, B. Ball, and C. W. Cunningham. 2005b. Diversity in the weapons of sexual selection: horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution 59:1060-1084.
– reference: García-González, F., and L. W. Simmons. 2007. Shorter sperm confer higher competitive fertilization success. Evolution 61:816-824.
– reference: Revell, L. J. 2012. Phytools: an R package for phylogenetic comparative biology (and other things). Methods Ecol. Evol. 3:217-223.
– reference: Kokko, H., H. Klug, and M. D. Jennions. 2012. Unifying cornerstones of sexual selection: operational sex ratio, Bateman gradient and the scope for competitive investment. Ecol. Let. 15:1340-1351.
– reference: Emlen, D. J. 1997. Alternative reproductive tactics and male-dimorphism in the horned beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Behav. Ecol. Sociobiol. 41:335-342.
– reference: Simmons, L. W.. 2007. Quantitative genetic correlation between trait and preference supports a sexually selected sperm process. Proc. Natnl. Acad. Sci. USA 104:16604-16608.
– reference: Kelly, C. D., and M. D. Jennions. 2011. Sexual selection and sperm quantity: meta-analyses of strategic ejaculation. Biol. Rev. 86:863-884.
– reference: Pagel, M. 1999. Inferring the historical patterns of biological evolution. Nature 401:877-884.
– reference: Hunt, J., and L. W. Simmons. 1997. Patterns of fluctuating asymmetry in beetle horns: an experimental examination of the honest signalling hypothesis. Behav. Ecol. Sociobiol. 41:109-114.
– reference: Emlen, D. J.. 2008. The evolution of animal weapons. Ann. Rev. Ecol. Evol. Syst. 39:387-413.
– reference: Knell, R. J., and L. W. Simmons. 2010. Mating tactics determine patterns of condition dependence in a dimorphic horned beetle. Proc. R. Soc. Lond. B 277:2347-2353.
– reference: Parker, G. A., and T. Pizzari. 2010. Sperm competition and ejaculate economics. Biol. Rev. 85:897-934.
– reference: Simmons, L. W., and J. S. Kotiaho. 2002. Evolution of ejaculates: patterns of phenotypic and genotypic variation and condition dependence in sperm competition traits. Evolution 56:1622-1631.
– reference: Emlen, D. J., J. Hunt, and L. W. Simmons. 2005a. Evolution of sexual dimorphism in the expression of beetle horns: phylogenetic evidence for modularity, evolutionary lability, and constraint. Am. Nat. 166:S42-S68.
– reference: Emlen, D. J.. 2001. Costs and the diversification of exaggerated animal structures. Science 291:1534-1536.
– reference: Kearse, M., R. Moir, A. Wilson, S. Stones-Havas, M. Cheung, S. Sturrock, S. Buxton, A. Cooper, S. Markowitz, C. Duran, et al. 2012. Geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28:1647-1649.
– reference: Werner, M., and L. W. Simmons. 2011. Ultrastructure of spermatozoa of Onthophagus taurus (Coleoptera, Scarabaeidae) exhibits heritable variation. Naturewissenschaften 98:213-223.
– reference: Drummond, A. J., M. A. Suchard, D. Xie, and A. Rambaut. 2012. Baysesian phylogenetics with BEAUti and the BEAST 1.7. Mol. Biol. Evol. 29:1969-1973.
– reference: Darriba, D., G. L. Taboada, R. Doallo, and D. Posada. 2012. jModelTest 2: more models, new heuristics and parallel computing. Nat. Methods 9:772.
– reference: Mendelson, T. C., and K. L. Shaw. 2005. Rapid speciation in an arthropod. Nature 433:375-376.
– reference: Simmons, L. W., and J. L. Fitzpatrick. 2012. Sperm wars and the evolution of male fertility. Reproduction 144:519-534.
– reference: Houle, D. 1992. Comparing evolvability and variability of quantitative traits. Genetics 130:195-204.
– reference: Kvarnemo, C., and L. W. Simmons. 2013. Polyandry as a mediator of sexual selection before and after mating. Philos. Trans. R Soc B Biol. Sci. 368:20120042.
– reference: Andersson, M. 1994. Sexual selection. Princeton Univ. Press, Princeton.
– reference: Fromhage, L., and H. Kokko. 2014. Sexually selected traits evolve positive allometry when some matings occur irrespective of the trait. Evolution 68:1332-1338.
– reference: Tomkins, J. L., and L. W. Simmons. 2000. Sperm competition games played by dimorphic male beetles: fertilisation gains with equal mating access. Proc. R. Soc. Lond. B 267:1547-1553.
– reference: Dunn, J. C., L. B. Halenar, T. G. Davies, J. Cristobal-Azkarate, D. Reby, D. Sykes, S. Dengg, W. T. Fitch, and L. A. Knapp. 2015. Evolutionary trade-off between vocal tract and testes dimensions in howler monkeys. Curr. Biol. 25:2839-2844.
– reference: Hunt, J.. 2001. Status-dependent selection in the dimorphic beetle Onthophagus taurus. Proc. R. Soc. Lond. B 268:2409-2414.
– reference: Arnqvist, G. 1998. Comparative evidence for the evolution of genitalia by sexual selection. Nature 393:784-786.
– reference: Berglund, A., A. Bisazza, and A. Pilastro. 1996. Armaments and ornaments: an evolutionary explanation of traits of dual utility. Biol. J. Linn. Soc. 58:385-399.
– reference: Maddison, W. P., and D. R. Maddison. 1999. MacClade: analysis of phylogeny and character evolution. Version 3.08a. Sinauer, Sunderland, MA.
– reference: Lüpold, S., J. L. Tomkins, L. W. Simmons, and J. L. Fitzpatrick. 2014. Female monopolization mediates the relationship between pre- and postcopulatory sexual traits. Nat. Comm. 5:3184. DOI: 3110.1038/ncomms4184.
– reference: van Noordwijk, A. J., and G. de Jong. 1986. Acquisition and allocation of resources: their influence on variation in life history tactics. Am. Nat. 128:137-142.
– reference: Clutton-Brock, T. H., and G. A. Parker. 1992. Potential reproductive rates and the operation of sexual selection. Quart. Rev. Biol. 67:437-456.
– reference: Simmons, L. W., M. Beveridge, and S. Krauss. 2004. Genetic analysis of parentage within experimental populations of a male dimorphic beetle, Onthophagus taurus, using amplified fragment length polymorphism. Behav. Ecol. Sociobiol. 57:164-173.
– reference: Freckleton, R. P., P. H. Harvey, and M. Pagel. 2002. Phylogenetic analysis and comparative data: a test and review of evidence. Am. Nat. 160:712-726.
– reference: Harmon, L. J., J. T. Weir, C. D. Brock, R. E. Glor, and W. Challenger. 2008. GEIGER: investigating evolutionary radiations. Bioinformatics 24:129-131.
– reference: Wilkinson, G. S., and M. Taper. 1999. Evolution of genetic variation for condition-dependent traits in stalk-eyed flies. Proc. R. Soc. Lond. B 266:1685-1690.
– reference: Simmons, L. W., and B. A. Buzatto. 2014. Contrasting responses of pre- and post-copulatory traits to variation in mating competition. Funct. Ecol. 28:494-499.
– reference: Simmons, L. W., and F. García-González. 2008. Evolutionary reduction in testes size and competitive fertilization success in response to the experimental removal of sexual selection in dung beetles. Evolution 62:2580-2591.
– reference: Simmons, L. W. 2001. Sperm competition and its evolutionary consequences in the insects. Princeton Univ. Press, Princeton.
– volume: 69
  start-page: 2613
  year: 2015
  end-page: 2624
– volume: 67
  start-page: 437
  year: 1992
  end-page: 456
  article-title: Potential reproductive rates and the operation of sexual selection
  publication-title: Quart. Rev. Biol.
– volume: 57
  start-page: 717
  year: 2003
  end-page: 745
  article-title: Testing for phylogenetic signal in comparative data: behavioral traits are more labile
  publication-title: Evolution
– volume: 68
  start-page: 1332
  year: 2014
  end-page: 1338
  article-title: Sexually selected traits evolve positive allometry when some matings occur irrespective of the trait
  publication-title: Evolution
– volume: 46
  start-page: 366
  year: 1992
  end-page: 375
  article-title: Individual variation in sperm competition success of yellow dung flies,
  publication-title: Evolution
– volume: 268
  start-page: 2409
  year: 2001
  end-page: 2414
  article-title: Status‐dependent selection in the dimorphic beetle
  publication-title: Proc. R. Soc. Lond. B
– volume: 86
  start-page: 863
  year: 2011
  end-page: 884
  article-title: Sexual selection and sperm quantity: meta‐analyses of strategic ejaculation
  publication-title: Biol. Rev.
– volume: 61
  start-page: 1316
  year: 2007
  end-page: 1326
  article-title: The intensity of sexual selection predicts weapon size in male bovids
  publication-title: Evolution
– year: 2001
– volume: 85
  start-page: 897
  year: 2010
  end-page: 934
  article-title: Sperm competition and ejaculate economics
  publication-title: Biol. Rev.
– volume: 59
  start-page: 459
  year: 2000
  end-page: 466
  article-title: Male horn dimorphism in the scarab beetle : do alternative reproductive tactics favor alternative phenotypes?
  publication-title: Anim. Behav
– volume: 67
  start-page: 95
  year: 2013
  end-page: 109
  article-title: Sperm competition games: a general model for pre‐copulatory male–male competition
  publication-title: Evolution
– volume: 263
  start-page: 315
  year: 1996
  end-page: 321
  article-title: Parental investment and the control of sexual selection: predicting the direction of sexual competition
  publication-title: Proc. R. Soc. Lond. B
– volume: 62
  start-page: 181
  year: 2013
  end-page: 192
  article-title: Comparing evolutionary rates for different phenotypic traits on a phylogeny using likelihood
  publication-title: Syst. Biol.
– volume: 32
  start-page: 1792
  year: 2004
  end-page: 1797
  article-title: MUSCLE: multiple sequence alignment with high accuracy and high throughput
  publication-title: Nucleic Acid Res.
– volume: 130
  start-page: 195
  year: 1992
  end-page: 204
  article-title: Comparing evolvability and variability of quantitative traits
  publication-title: Genetics
– volume: 128
  start-page: 137
  year: 1986
  end-page: 142
  article-title: Acquisition and allocation of resources: their influence on variation in life history tactics
  publication-title: Am. Nat.
– volume: 291
  start-page: 1534
  year: 2001
  end-page: 1536
  article-title: Costs and the diversification of exaggerated animal structures
  publication-title: Science
– volume: 39
  start-page: 387
  year: 2008
  end-page: 413
  article-title: The evolution of animal weapons
  publication-title: Ann. Rev. Ecol. Evol. Syst.
– volume: 103
  start-page: 16349
  year: 2006
  end-page: 16351
  article-title: Evolutionary trade‐off between weapons and testes
  publication-title: Proc. Natl. Acad. Sci. USA
– year: 1994
– volume: 368
  start-page: 20120042
  year: 2013
  article-title: Polyandry as a mediator of sexual selection before and after mating
  publication-title: Philos. Trans. R Soc B Biol. Sci.
– volume: 160
  start-page: 712
  year: 2002
  end-page: 726
  article-title: Phylogenetic analysis and comparative data: a test and review of evidence
  publication-title: Am. Nat.
– volume: 433
  start-page: 375
  year: 2005
  end-page: 376
  article-title: Rapid speciation in an arthropod
  publication-title: Nature
– volume: 62
  start-page: 2580
  year: 2008
  end-page: 2591
  article-title: Evolutionary reduction in testes size and competitive fertilization success in response to the experimental removal of sexual selection in dung beetles
  publication-title: Evolution
– volume: 69
  start-page: 1044
  year: 2015
  end-page: 1052
  article-title: Postcopulatory sexual selection is associated with accelerated evolution of sperm morphology
  publication-title: Evolution
– volume: 28
  start-page: 494
  year: 2014
  end-page: 499
  article-title: Contrasting responses of pre‐ and post‐copulatory traits to variation in mating competition
  publication-title: Funct. Ecol.
– volume: 5
  start-page: 3184
  year: 2014
  article-title: Female monopolization mediates the relationship between pre‐ and postcopulatory sexual traits
  publication-title: Nat. Comm.
– volume: 144
  start-page: 519
  year: 2012
  end-page: 534
  article-title: Sperm wars and the evolution of male fertility
  publication-title: Reproduction
– volume: 61
  start-page: 816
  year: 2007
  end-page: 824
  article-title: Shorter sperm confer higher competitive fertilization success
  publication-title: Evolution
– volume: 56
  start-page: 1622
  year: 2002
  end-page: 1631
  article-title: Evolution of ejaculates: patterns of phenotypic and genotypic variation and condition dependence in sperm competition traits
  publication-title: Evolution
– volume: 10
  start-page: 97
  year: 1996
  end-page: 104
  article-title: Sexual selection and the allometry of earwig forceps
  publication-title: Evol. Ecol.
– volume: 266
  start-page: 1685
  year: 1999
  end-page: 1690
  article-title: Evolution of genetic variation for condition‐dependent traits in stalk‐eyed flies
  publication-title: Proc. R. Soc. Lond. B
– volume: 61
  start-page: 2684
  year: 2007
  end-page: 2692
  article-title: Sperm competition games between sneaks and guards: a comparative analysis using dimorphic male beetles
  publication-title: Evolution
– volume: 55
  start-page: 32
  year: 2003
  end-page: 41
  article-title: Correlates of horn and antler shape in bovids and cervids
  publication-title: Behav. Ecol. Sociobiol.
– volume: 41
  start-page: 109
  year: 1997
  end-page: 114
  article-title: Patterns of fluctuating asymmetry in beetle horns: an experimental examination of the honest signalling hypothesis
  publication-title: Behav. Ecol. Sociobiol.
– volume: 104
  start-page: 16604
  year: 2007
  end-page: 16608
  article-title: Quantitative genetic correlation between trait and preference supports a sexually selected sperm process
  publication-title: Proc. Natnl. Acad. Sci. USA
– volume: 24
  start-page: 2378
  year: 2011
  end-page: 2388
  article-title: Rates of phenotypic evolution of ecological characters and sexual traits during the adaptive radiation
  publication-title: J. Evol. Biol.
– volume: 69
  start-page: 1560
  year: 2015
  end-page: 1572
  article-title: A trade‐off between precopulatory and postcopulatory trait investment in male cetaceans
  publication-title: Evolution
– volume: 41
  start-page: 335
  year: 1997
  end-page: 342
  article-title: Alternative reproductive tactics and male‐dimorphism in the horned beetle (Coleoptera: Scarabaeidae)
  publication-title: Behav. Ecol. Sociobiol
– volume: 9
  start-page: 772
  year: 2012
  article-title: jModelTest 2: more models, new heuristics and parallel computing
  publication-title: Nat. Methods
– volume: 24
  start-page: 1706
  year: 2011
  end-page: 1714
  article-title: Adjustment of sperm allocation under high risk of sperm competition across taxa: a meta‐analysis
  publication-title: J. Evol. Biol.
– volume: 280
  start-page: 20131065
  year: 2013
  article-title: Sexual selection accelerates signal evolution during speciation in birds
  publication-title: Proc. R. Soc. Lond. B
– volume: 66
  start-page: 3595
  year: 2012
  end-page: 3604
  article-title: Male contest competition and the coevolution of weaponry and testes in pinnipeds
  publication-title: Evolution
– volume: 277
  start-page: 3639
  year: 2010
  end-page: 3648
  article-title: Condition‐dependent ejaculate size and composition in a ladybird beetle
  publication-title: Proc. R. Soc. Lond. B
– volume: 267
  start-page: 1547
  year: 2000
  end-page: 1553
  article-title: Sperm competition games played by dimorphic male beetles: fertilisation gains with equal mating access
  publication-title: Proc. R. Soc. Lond. B
– volume: 58
  start-page: 1038
  year: 2004
  end-page: 1046
  article-title: Condition dependence of sexual ornament size and variation in the stalk‐eyed fly (Diptera: Diopsidae)
  publication-title: Evolution
– volume: 58
  start-page: 385
  year: 1996
  end-page: 399
  article-title: Armaments and ornaments: an evolutionary explanation of traits of dual utility
  publication-title: Biol. J. Linn. Soc.
– volume: 25
  start-page: 2839
  year: 2015
  end-page: 2844
  article-title: Evolutionary trade‐off between vocal tract and testes dimensions in howler monkeys
  publication-title: Curr. Biol.
– volume: 166
  start-page: S42
  year: 2005a
  end-page: S68
  article-title: Evolution of sexual dimorphism in the expression of beetle horns: phylogenetic evidence for modularity, evolutionary lability, and constraint
  publication-title: Am. Nat.
– volume: 29
  start-page: 1969
  year: 2012
  end-page: 1973
  article-title: Baysesian phylogenetics with BEAUti and the BEAST 1.7
  publication-title: Mol. Biol. Evol.
– volume: 19
  start-page: 331
  year: 2008
  end-page: 337
  article-title: Male morph predicts investment in larval immune function in the dung beetle,
  publication-title: Behav. Ecol
– volume: 63
  start-page: 2985
  year: 2009
  end-page: 2998
  article-title: Plumage evolution in the Oropendolas and Caciques: different divergence rates in polygynous and monogamous taxa
  publication-title: Evolution
– volume: 15
  start-page: 1340
  year: 2012
  end-page: 1351
  article-title: Unifying cornerstones of sexual selection: operational sex ratio, Bateman gradient and the scope for competitive investment
  publication-title: Ecol. Let.
– volume: 20
  start-page: 381
  year: 2007
  end-page: 391
  article-title: Influence of developmental environment on male‐ and female‐mediated sperm precedence in
  publication-title: J. Evol. Biol
– volume: 401
  start-page: 877
  year: 1999
  end-page: 884
  article-title: Inferring the historical patterns of biological evolution
  publication-title: Nature
– volume: 393
  start-page: 784
  year: 1998
  end-page: 786
  article-title: Comparative evidence for the evolution of genitalia by sexual selection
  publication-title: Nature
– year: 1871
– volume: 118
  start-page: 64
  year: 2015
  end-page: 77
  article-title: A cautionary note on the use of Ornstein Uhlenbeck models in macroevolutionary studies
  publication-title: Biol. J. Linn. Soc.
– year: 2002
– volume: 56
  start-page: 1683
  year: 2002
  end-page: 1695
  article-title: Antler size in red deer: heritability and selection but no evolution
  publication-title: Evolution
– volume: 3
  start-page: 217
  year: 2012
  end-page: 223
  article-title: Phytools: an R package for phylogenetic comparative biology (and other things)
  publication-title: Methods Ecol. Evol.
– volume: 57
  start-page: 164
  year: 2004
  end-page: 173
  article-title: Genetic analysis of parentage within experimental populations of a male dimorphic beetle, , using amplified fragment length polymorphism
  publication-title: Behav. Ecol. Sociobiol
– volume: 45
  start-page: 525
  year: 1970
  end-page: 567
  article-title: Sperm competition and its evolutionary consequences in the insects
  publication-title: Biol. Rev.
– volume: 59
  start-page: 1060
  year: 2005b
  end-page: 1084
  article-title: Diversity in the weapons of sexual selection: horn evolution in the beetle genus (Coleoptera: Scarabaeidae)
  publication-title: Evolution
– volume: 28
  start-page: 1647
  year: 2012
  end-page: 1649
  article-title: Geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data
  publication-title: Bioinformatics
– volume: 24
  start-page: 129
  year: 2008
  end-page: 131
  article-title: GEIGER: investigating evolutionary radiations
  publication-title: Bioinformatics
– volume: 277
  start-page: 2347
  year: 2010
  end-page: 2353
  article-title: Mating tactics determine patterns of condition dependence in a dimorphic horned beetle
  publication-title: Proc. R. Soc. Lond. B
– year: 1999
– volume: 98
  start-page: 213
  year: 2011
  end-page: 223
  article-title: Ultrastructure of spermatozoa of (Coleoptera, Scarabaeidae) exhibits heritable variation
  publication-title: Naturewissenschaften
SSID ssj0009519
Score 2.2881613
Snippet Reproductive competition generates episodes of both pre- and postcopulatory sexual selection. Theoretical models of sperm competition predict that as the...
Reproductive competition generates episodes of both pre‐ and postcopulatory sexual selection. Theoretical models of sperm competition predict that as the...
SourceID swepub
proquest
pubmed
wiley
jstor
istex
SourceType Open Access Repository
Aggregation Database
Index Database
Publisher
StartPage 998
SubjectTerms Animal reproduction
Animals
Body size
Coleoptera - genetics
Coleoptera - physiology
Competition
Copulation
Dung
Ejaculate expenditure
Evolution
Evolution, Molecular
evolutionary divergence
Female
Fertilization
horns
Insects
life-history trade-off
Male
male contest competition
Mating Preference, Animal
Organ Size
Sperm
sperm length
Spermatozoa - cytology
Spermatozoa - physiology
Testis - anatomy & histology
Testis - growth & development
Weapons
Title Sperm competition and the coevolution of pre- and postcopulatory traits: Weapons evolve faster than testes among onthophagine dung beetles
URI https://api.istex.fr/ark:/67375/WNG-3SDMFT21-8/fulltext.pdf
https://www.jstor.org/stable/24762186
https://onlinelibrary.wiley.com/doi/abs/10.1111%2Fevo.12915
https://www.ncbi.nlm.nih.gov/pubmed/27061413
https://www.proquest.com/docview/1789930183
https://www.proquest.com/docview/1790463010
https://www.proquest.com/docview/1808669171
https://urn.kb.se/resolve?urn=urn:nbn:se:su:diva-132514
Volume 70
hasFullText 1
inHoldings 1
isFullTextHit
isPrint
link http://utb.summon.serialssolutions.com/2.0.0/link/0/eLvHCXMwnV1bi9QwFA7LguCL99XqKhFUfOkwSdObPi3ujouwK7jXByEk6akuC21pO4Prkz_AB3-jv8Rz0pm6ioj4NkxOmHRyLt9JT77D2BOjIIlUnIepS6ahKqUJbaogxNhWSOmscoYuJ-_tJ7tH6s1pfLrGXq7uwgz8EOOBG1mG99dk4MZ2l4wcFvUEg5W_YE61WgSI3slLhLtigL5ChRGGuSWrEFXxjDMRkNJ_-WlVi_gnlDlSiP6KXn34mV1n71cLH6pOzifz3k7c5984Hf_zyW6wa0tYyrcGPbrJ1qC6xa4MjSovbrOvBw16cO48yPZFXtxUBUfwiN_BYqm-vC5508L3L9_8aFN3vRv6g9XtBadmFH33gp-AoapcTtMWwEtDVA2cTvB5T4evHfcNkHhNtAbNR-qiBLxAn8QtAGp2d4cdzXYOX-2Gyz4O4ZlK4ziMpHVpWtrMAOJBY4R04GRhQBmMjZBIFQubxDE6j9xliSLKHAuyUOAEpBFEG2y9qiu4x3gR5TCVpcmJVR6xoHU2clMBpVJFjp46YM_8jupm4OrQpj2n0rU01if7r3V0sL03O5RCZwHb8Fs-CkqFYUFkScA2Vzqgl-bcaYEry9EVZlHAHo_DaIj0dsVUUM9JJif2Ncxv_yKTYQaZYIYsAnZ30K-fC0gpORf4C08HhRtHiAF8--x4S9ftB93NtYgQlKqAPfdKNIqt0jncPu3VR-8cv_Uf7v-76AN2FSFhMpR0brL1vp3DQ4RdvX3k7esHa0srXA
linkProvider Wiley-Blackwell
linkToHtml http://utb.summon.serialssolutions.com/2.0.0/link/0/eLvHCXMwnV1Lb9NAEF5VRQguvAuGAosEiIuj7Hr9Qlwq2hCgCRJNH5dqtV6PoapkR7ETUU78AA78Rn4JM-vEFIQQ4hZlZ5V1dh7frGe_YeyxURAFKkz92EZ9XxXS-FmswMfYlktpM2UNXU4ejaPhvnpzFB6tsReruzAtP0R34EaW4fw1GTgdSJ-zclhUPYxWdMP8AnX0dgnVe3mOcle04FcoP8BAt-QVojqebipCUvo3P62qEf-EMzsS0V_xqwtAg6vseLX0tu7ktDdvsp79_Bur4_8-2zV2ZYlM-VarStfZGpQ32MW2V-XZTfZ1b4pOnFuHs12dFzdlzhE_4newWGowrwo-ncH3L9_c6LSqG9u2CKtmZ5z6UTT1c34IhgpzOU1bAC8MsTVwOsTnDZ2_1tz1QOIVMRtMP1IjJeA5uiWeAaBy17fY_mBn8nLoL1s5-CcqDkM_kJmN4yJLDCAkNEZIC1bmBpTB8AiRVKHIojBE_5HaJFLEmpOBzBVYAXEAwQZbL6sS7jCeByn0ZWFSIpZHOJjZLLB9AYVSeYrO2mNP3ZbqaUvXoc3slKrX4lAfjl_pYG97NJhIoROPbbg97wSlwsggkshjmysl0EuLrrXAlaXoDZPAY4-6YbRFesFiSqjmJJMSARumuH-RSTCJjDBJFh673SrYzwXElJ8L_IUnrcZ1I0QCvn1ysKWr2Qddz7UIEJcqjz1zWtSJrTI63D7t1EfvHLxzH-7-u-hDdmk4Ge3q3dfjt_fYZUSIUVvhucnWm9kc7iMKa7IHzth-APZ3L3c
linkToPdf http://utb.summon.serialssolutions.com/2.0.0/link/0/eLvHCXMwnV1Lj9MwELZWi0BceC8EFjASIC6pasd5wWlFtyyPLYh9HlayHGcCq5WSqEkrlhM_gAO_kV_CjNOGBSGEuFX1WHXqeXzjjL9h7KFREAUqTP3YRkNfFdL4WazAx9iWS2kzZQ1dTt6eRFt76tVheLjCni3vwnT8EP2BG1mG89dk4HVenDFymFcDDFZ0wfycioYJqfTovTzDuCs67CuUH2CcW9AKURlPPxURKf2Zn5bFiH-CmT2H6K_w1cWf8WV2tFx5V3ZyMpi12cB-_o3U8T8f7Qq7tMClfKNTpKtsBcpr7HzXqfL0Ovu6U6ML59ahbFflxU2Zc0SP-B3MF_rLq4LXU_j-5ZsbraumtV2DsGp6yqkbRds85QdgqCyX07Q58MIQVwOnI3ze0ulrw10HJF4Rr0H9kdooAc_RKfEMAFW7ucH2xpu7z7f8RSMH_1jFYegHMrNxXGSJAQSExghpwcrcgDIYHCGSKhRZFIboPVKbRIo4czKQuQIrIA4gWGOrZVXCLcbzIIWhLExKtPIIBjObBXYooFAqT9FVe-yx21Fdd2Qd2kxPqHYtDvXB5IUOdkbb410pdOKxNbflvaBUGBdEEnlsfakDemHPjRa4shR9YRJ47EE_jJZIr1dMCdWMZFKiX8ME9y8yCaaQEabIwmM3O_36uYCYsnOBv_CoU7h-hCjAR8f7G7qaftDNTIsAUany2BOnRL3YMp_D7dNOffTm_lv34fa_i95nF96NxvrNy8nrO-wiwsOoK-9cZ6vtdAZ3EYK12T1naj8AbmQuLw
openUrl ctx_ver=Z39.88-2004&ctx_enc=info%3Aofi%2Fenc%3AUTF-8&rfr_id=info%3Asid%2Fsummon.serialssolutions.com&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Sperm+competition+and+the+coevolution+of+pre%E2%80%90+and+postcopulatory+traits%3A+Weapons+evolve+faster+than+testes+among+onthophagine+dung+beetles&rft.jtitle=Evolution&rft.au=Simmons%2C+Leigh+W.&rft.au=Fitzpatrick%2C+John+L.&rft.date=2016-05-01&rft.issn=0014-3820&rft.eissn=1558-5646&rft.volume=70&rft.issue=5&rft.spage=998&rft.epage=1008&rft_id=info:doi/10.1111%2Fevo.12915&rft.externalDBID=10.1111%252Fevo.12915&rft.externalDocID=EVO12915
thumbnail_l http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/lc.gif&issn=0014-3820&client=summon
thumbnail_m http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/mc.gif&issn=0014-3820&client=summon
thumbnail_s http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/sc.gif&issn=0014-3820&client=summon