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publications in refereed journals and books (1987-2004)
Submitted for publication
Chaline, N., Sandoz, J. C., Martin, S. J., Ratnieks, F. L. W., Jones, G. R. Learning and
discrimination of individual cuticular hydrocarbons by honey bees (Apis mellifera). (submitted)
Jensen, A. B., Palmer, K. A., Chaline, N., Raine. N. E., Tofilski, A., Pedersen, B. V., Boomsma, J. J.,
Ratnieks, F. L. W. Quantification of honey bee mating range and isolation in semi-isolated valleys in the
Peak District National Park, England, as revealed by paternity analysis using DNA microsatellites. (submitted).
Medina, L. M., Martin, S. J., Ratnieks, F. L. W. Reproduction of Varroa destructor
in drone brood cells of Africanized honey bees (Apis mellifera) in Yucatan, Mexico. (submitted)
Medina, L., Ratnieks, F. L. W. A scientific note on the occurence of hygienic behaviour in native British
honey bees (Apis mellifera mellifera). (submitted)
Martin, S. J., Ratnieks, F. L. W., Jones, G. Dufour's gland and egg recognition in honey bees. (submitted)
Paxton, R. J., Ruhnke, H., Ratnieks, F. L. W., Quezada-Euan, J. J. G. Queen monoandry yet lack of worker maternity
of males in the stingless bee Melipona beecheii (Apidae, Meliponini). (submitted)
Accepted for publication
Beekman, M., Seraphides N, Sumpter, D. J., Ratnieks, F. L. W. Patch use by honey bee colonies:
does colony size matter? (accepted)
Chaline, N., Martin, S. J., Ratnieks, F. L. W. Absence of nepotism towards imprisoned
young queens during swarming in the honey bee. Behavioral Ecology (accepted).
Ratnieks, F.L.W., Foster, K.R., Wenseleers, T. 2006. Conflict resolution in insect societies.
Annual Review of Entomology, in press.
(galleys, unrevised)
Wood, M. J., Ratnieks, F. L. W. Transfer of olfactory cues used in Vespula
wasp recognition by honey bee guards. Apidologie (accepted).
Published
Jackson, D.E., Ratnieks, F.L.W. 2006. Primer: Communication in Ants.
Current Biology, 16: R570-R574.
Tofilski, A., Ratnieks, F.L.W. 2006. Sand pile Formation in Dorymyrmex ants.
Journal of Insect Behaviour, 18: 505-512.
117. Robinson EJH, Jackson DE, Holcombe M, Ratnieks FLW. 2005. 'No entry' signal in ant foraging.
Nature, 438: 442.
Supplementary Information
116. Ratnieks, F.L.W., Wenseleers, T. 2005. Policing Insect Societies.
Science, 307: 54-56.
115. Wenseleers, T., Ratnieks, F.L.W., Ribeiro, M.F., Alves, D.A., Imperatriz-Fonseca, V.-L. Working-class royalty: bees beat the caste system. 2005.
Biology Letters, 1: 125-128.
114. Wenseleers, T. Tofilski, A., Ratnieks, F.L.W. 2005. Queen and worker policing in the tree wasp Dolichovespula sylvestris. Behavioral Ecology & Sociobiology, 58: 80-86.
113. Wenseleers, T., Badcock, N.S., Erven, K., Tofilski, A., Nascimento, F.S., Hart, A.G., Burke, T.A., Archer, M.E., Ratnieks, F.L.W. A test of worker policing theory in an advanced eusocial wasp, Vespula rufa. Evolution, 59: 1306-1314.
112. Chaline, N., Martin, S. J., Ratnieks, F. L. W. 2004. Worker policing persists in a hopelessly queenless honey bee colony (Apis mellifera).
Insectes Sociaux, 51: 113-116.
111. Chaline, N., Ratnieks, F. L. W., Raine, N. E., Badcock, N. S., Burke, T. 2004. Non-lethal sampling of honey bee, Apis mellifera, DNA using wing tips. Apidologie, 35: 311-318. 
110. D'Ettorre, P., Heinze, J., Ratnieks, F. L. W. 2004. Worker policing by egg eating in the ponerine ant Pachycondyla inversa. Proceedings of the Royal Society of London Series B-Biological Sciences 271: 1427-1434.
109. Jackson, D. E., Holcombe, M., Ratnieks, F. L. W. 2004. Geometry gives polarity to ant pheromone trails.
Nature 432: 907-909.
Supplement1
Supplement2
108. Martin, S. J., Chaline, N., Oldroyd, B. P., Jones, G. R., Ratnieks, F. L. W. 2004. Egg marking pheromones of anarchistic worker honeybees (Apis mellifera).
Behavioral Ecology 15: 839-844.
107. Martin, S.J., Jones, G. R., Chaline, N., Ratnieks F. L. W. 2004. Role of hydrocarbons in egg recognition in the honeybee.
Physiological Entomology 29: 395-399.
106. Wenseleers T., Hart A. G., Ratnieks F.L.W. 2004. When resistance is useless: policing and the evolution of reproductive acquiescence in insect societies.
American Naturalist 164: E154 -E167
105. Wenseleers, T., Hart, A. G., Ratnieks, F. L. W., Quezada-Euan, J. J. G. 2004. Queen execution and caste conflict in the stingless bee Melipona beecheii. Ethology, 110: 725-736.
104. Wenseleers, T., Helantera, H., Hart, A. G., Ratnieks, F. L. W. 2004. Worker reproduction and policing in insect societies: an ESS analysis. Journal of Evolutionary Biology 17: 1035-1047. 
103. Wenseleers, T., Ratnieks, F. L. W. 2004. Tragedy of the commons in Melipona bees. Proceedings of the Royal Society of London Series B-Biological Sciences 271: S310-312.
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102. Wood, M., Ratnieks, F.L.W. 2004. Olfactory cues and Vespula wasp recognition by honey bee guards. Apidologie 35: 461-468. 
101. Beekman, M., Komdeur, J., Ratnieks,
F. L. W. 2003. Reproductive conflicts in social animals -
who has power. Trends in Ecology and Evolution, 18: 277-282.

100. Beekman, M., Ratnieks, F. L. W.
2003. Power over reproduction in social Hymenoptera. Philosophical
Transactions of the Royal Society, 258: 1741-1753. 
99. Chaline N, Arnold G, Papin C, et
al. 2003. Patriline differences in emergency queen rearing
in the honey bee, Apis mellifera. Insectes Sociaux
50: 234-236.
98. Gregson, A. M., Hart, A. G., Holcombe,
M., Ratnieks, F. L. W. 2003. Partial nectar loads as a cause
of multiple nectar transfer in the honey bee (Apis mellifera):
a simulation model. Journal of Theoretical Biology 222: 1-8.
97.
Hart, A. G., Bowtell, R.
W., Kockenberger, W.,
Wenseleers, T., Ratnieks, F.
L. W. 2003. Magnetic
resonance imaging in entomology: a critical review. Journal
of Insect Science, 3.5. 9 pages. Available online: insectscience.org/3.5
96. Jeanson, R., Ratnieks, F. L. W.
& Deneubourg, J. L. 2003. Pheromone trail decay rates
on different substrates in the Pharaoh's ant, Monomorium
pharaonis. Physiological Entomology 28: 192-198. 
95. Monnin T., Ratnieks F. L. W., Brandão
C. R. F. 2003. Reproductive conflict in animal societies:
hierarchy length increases with colony size in queenless ponerine
ants. Behavioral Ecology & Sociobiology 54: 71-79. 
94. Pirk, C. A. A., Neumann, P., Ratnieks, F. L. W. 2003. Cape honeybees police worker-laid eggs despite
the absence of relatedness benefits. Behavioral Ecology, 14:
347-352. 
93.
Wenseleers, T., Ratnieks, F. L. W., Billen, J. 2003. Caste
fate conflict in swarm-founding social Hymenoptera: an inclusive
fitness analysis. Journal of Evolutionary Biology, 16: 647-658.

92. Beekman, M., Calis, J. N. M., Oldroyd,
B. P., Ratnieks, F. L. W. 2002. When do honeybee guards reject
their former nestmates after swarming? Insectes Sociaux 49:
56-61. 
91. Beekman, M., Good, G., Allsopp,
M. H., Radloff, S., Pirk, C. W. W., Ratnieks, F. L. W. 2002.
A non-policing honey bee colony (Apis mellifera capensis).
Naturwissenschaften 89: 479-482. 
90. Beekman, M., Wossler, T. C., Martin,
S. J., Ratnieks, F. L. W. 2002. Parasitic Cape honeybee workers
(Apis mellifera capensis) are not given differential
treatment by African guards (A. m. scuttelata). Insectes
Sociaux 49: 216-220. 
89. Chaline, N., Ratnieks, F. L. W.,
Burke, T. 2002. Anarchy in the UK: Detailed genetic analysis
of worker reproduction in a naturally occurring British anarchistic
honeybee, Apis mellifera, colony using DNA microsatellites.
Molecular Ecology 11: 1795-1803. 
88. Foster, K. R., Gulliver, J., Ratnieks,
F. L. W. 2002. Worker policing in the European hornet Vespa
crabro. Insectes Sociaux 49: 41-44. 
87. Hart, A. G., Anderson, C., Ratnieks,
F. L. W. 2002. Task partitioning in leafcutting ants. Acta
Ethologica 5:1-11. 
86. Hart, A. G. and Ratnieks,
F. L. W. 2002. Task partitioned nectar transfer in stingless
bees (Meliponini): work organisation in a phylogenetic context.
Ecological Entomology 27: 163-168. 
85. Hart, A. G. and Ratnieks,
F. L. W. 2002. Waste management in the leafcutting ant Atta
colombica. Behavioral Ecology 13: 224-231. 
84. Martin, S. J., Beekman, M., Wossler,
T. C., Ratnieks, F. L. W. 2002. Parasitic Cape honeybee workers,
Apis mellifera capensis, evade policing. Nature 415:
163-165.
83. Martin, S. J., Jones, G. R., Châline,
N., Middleton, H & Ratnieks, F. L. W. 2002. Reassessing
the role of the honeybee (Apis mellifera) Dufour's
gland in egg marking. Naturwissenschaften 89: 528-532.
82. Medina, L. M., Martin, S. J., Espinoza-Montaño,
L., Ratnieks, F. L. W. 2002. Reproduction of Varroa destructor
in worker brood of Africanized honey bees (Apis mellifera).
Experimental and Applied Acarology 27: 79-88. 
81. Monnin, T., Ratnieks, F. L. W.,
Jones, G. R., Beard, R. 2002. Pretender punishment induced
by chemical signalling in a queenless ant. Nature 419: 61-65.
80. Oldroyd, B. P., Ratnieks, F. L.
W., Wossler, T. C. 2002. Egg marking pheromones in honey-bees
Apis mellifera. Behavioural Ecology and Sociobiology
51: 590-591.
79. Wattanachaiyingcharoen, W., Oldroyd,
B. P., Good, G., Halling, L. Ratnieks, F. L. W., Wongsiri,
S. 2002. Lack of worker reproduction in the giant honey bee
Apis dorsata Fabricius. Insectes Sociaux 49: 80-85.
78. Barron, A. B., B. P. Oldroyd, and
F. L. W. Ratnieks. 2001. Worker reproduction in honey-bees
(Apis) and the anarchic syndrome: a review. Behavioral
Ecology and Sociobiology 50:199-208.
77. Beekman, M., D. J. T. Sumpter,
and F. L. W. Ratnieks. 2001. Phase transition between disordered
and ordered foraging in Pharaoh's ants. Proceedings of the
National Academy of Sciences of the United States of America
98:9703-9706.
76. Bourke, A. F. G., and F. L. W. Ratnieks.
2001. Kin-selected conflict in the bumble bee Bombus terrestris
(Hymenoptera : Apidae). Proceedings of the Royal Society of
London Series B-Biological Sciences 268:347-355.
75. Downs, S. G., F. L. W. Ratnieks,
N. S. Badcock, and A. Mynott. 2001. Honeybee guards do not
use food-derived odors to recognize non- nest mates: a test
of the Odor Convergence hypothesis. Behavioral Ecology 12:47-50.
74. Foster, K. R., and F. L. W. Ratnieks
2001. The effect of sex allocation biasing on the evolution
of worker policing. American Naturalist 158:615-623.
73. Foster, K. R., and F. L. W. Ratnieks.
2001. Convergent evolution of worker policing by egg eating
in the honeybee and common wasp. Proceedings of the Royal
Society of London Series B-Biological Sciences 268:169-174.
72. Foster, K. R., and F. L. W. Ratnieks.
2001. Paternity, reproduction and conflict in vespine wasps:
a model system for testing kin selection predictions. Behavioral
Ecology and Sociobiology 50:1-8.
71. Foster, K. R., F. L. W. Ratnieks,
N. Gyllenstrand, and P. A. Thoren. 2001. Colony kin structure
and male production in Dolichovespula wasps. Molecular
Ecology 10:1003-1010.
70. Foster, K. R., T. Wenseleers, and
F. L. W. Ratnieks. 2001. Spite: Hamilton's unproven theory.
Annales Zoologici Fennici 38:229-238.
69. Hart, A. G., and F. L. W. Ratnieks.
2001. Leaf caching in the leafcutting ant Atta colombica:
organizational shift, task partitioning and making the best
of a bad job. Animal Behaviour 62:227-234.
68. Hart, A. G., and F. L. W. Ratnieks.
2001. Task partitioning, division of labour and nest compartmentalisation
collectively isolate hazardous waste in the leafcutting ant
Atta cephalotes. Behavioral Ecology and Sociobiology
49:387-392.
67. Hart, A. G., and F. L. W. Ratnieks.
2001. Why do honey-bee (Apis mellifera) foragers transfer
nectar to several receivers? Information improvement through
multiple sampling in a biological system. Behavioral Ecology
and Sociobiology 49:244-250.
66. Miller, D. G., and F. L. W. Ratnieks.
2001. The timing of worker reproduction and breakdown of policing
behaviour in queenless honey bee (Apis mellifera L.)
societies. Insectes Sociaux 48:178-184.
65. Monnin, T., and F. L. W. Ratnieks.
2001. Policing in queenless ponerine ants. Behavioral Ecology
and Sociobiology 50:97-108.
64. Neumann, P., C. W. W. Pirk, H. R.
Hepburn, A. J. Solbrig, F. L. W. Ratnieks, P. J. Elzen, and
J. R. Baxter. 2001. Social encapsulation of beetle parasites
by Cape honeybee colonies (Apis mellifera capensis
Esch.). Naturwissenschaften 88:214-216.
63. Oldroyd, B. P., L. A. Halling, G.
Good, W. Wattanachaiyingcharoen, A. B. Barron, P. Nanork,
S. Wongsiri, and F. L. W. Ratnieks. 2001. Worker policing
and worker reproduction in Apis cerana. Behavioral
Ecology and Sociobiology 50:371-377.
62. Oldroyd, B. P., T. C. Wossler, and
F. L. W. Ratnieks. 2001. Regulation of ovary activation in
worker honey-bees (Apis mellifera): larval signal production
and adult response thresholds differ between anarchistic and
wild-type bees. Behavioral Ecology and Sociobiology 50:366-370.
61. Ratnieks, F. L. W. 2001. Heirs and
spares: caste conflict and excess queen production in Melipona
bees. Behavioral Ecology and Sociobiology 5:467-473; DOI 10.1007/s002650100388.
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talk on this (powerpoint)
60. Ratnieks, F. L. W., A. Lapish, and
L. Tann. 2001. Honey bee guards, Apis mellifera, accept
own subspecies non-nestmates more than other subspecies non-nestmates.
Insectes Sociaux 48:287-288.
59. Ratnieks, F. L. W., T. Monnin, and
K. R. Foster. 2001. Inclusive fitness theory: novel predictions
and tests in eusocial Hymenoptera. Annales Zoologici Fennici
38:201-214.
58. Anderson, C., and F. L. W. Ratnieks.
2000. Task partitioning in insect societies: novel situations.
Insectes Sociaux 47:198-199.
57. Beekman, M., and F. L. W. Ratnieks.
2000. Long-range foraging by the honey-bee, Apis mellifera
L. Functional Ecology 14:490-496.
56. Bernasconi, G., F. L. W. Ratnieks,
and E. Rand. 2000. Effect of "spraying" by fighting honey
bee queens (Apis mellifera L.) on the temporal structure
of fights. Insectes Sociaux 47:21-26.
55. Downs, S. G., and F. L. W. Ratnieks.
2000. Adaptive shifts in honey bee (Apis mellifera
L.) guarding behavior support predictions of the acceptance
threshold model. Behavioral Ecology 11:326-333.
54. Downs, S. G., F. L. W. Ratnieks,
S. L. Jefferies, and H. E. Rigby. 2000. The role of floral
oils in the nestmate recognition system of honey bees (Apis
mellifera L.). Apidologie 31:357-365. 
53. Foster, K. R., and F. L. W. Ratnieks.
2000. Facultative worker policing in a wasp. Nature 407:692-693.
52. Foster, K. R., F. L. W. Ratnieks,
and A. F. Raybould. 2000. Do hornets have zombie workers?
Molecular Ecology 9:735-742.
51. Foster, K. R., F. L. W. Ratnieks,
and T. Wenseleers. 2000. Spite in social insects. Trends in
Ecology and Evolution 15:469-470.
50. Hart, A. G., and F. L. W. Ratnieks.
2000. Leaf caching in Atta leafcutting ants: discrete
cache formation through positive feedback. Animal Behaviour
59:587-591. 
49. Oldroyd, B. P., and F. L. W. Ratnieks.
2000. Evolution of worker sterility in honey-bees (Apis
mellifera): how anarchistic workers evade policing by
laying eggs that have low removal rates. Behavioral Ecology
and Sociobiology 47:268-273.
48. Ratnieks, F. L. W. 2000. Worker
policing in the honey bee: basic facts and ideas. Insect Social
Life 3:3-10. 
47. Anderson, C., and F. L. W. Ratnieks.
1999. Task partitioning in insect societies. I. Effect of
colony size on queueing delay and colony ergonomic efficiency.
American Naturalist 154:521-535.
46. Anderson, C., and F. L. W. Ratnieks.
1999. Worker allocation in insect societies: coordination
of nectar foragers and nectar receivers in the honey bee.
Behavioral Ecology and Sociobiology 46:73-81.
45. Anderson, C., Ratnieks, F. L. W.
1999. Task partitioning in foraging: effect of colony size
on queueing times and information reliability. Pages 31-50
in: Detrain C, Deneubourg J L, Pasteels J M (eds.). Information
Processing in Social Insects. Birkhauser, Basel. 
44. Bernasconi, G., Bigler, L., Hesse,
M., Ratnieks, F. L. W. 1999. Characterization of queen specific
components of the fluid released by fighting honey bee queens.
Chemoecology 9: 161-167.
43. Bourke, A. F. G., and F. L. W. Ratnieks.
1999. Kin conflict over caste determination in social Hymenoptera.
Behavioral Ecology and Sociobiology 46:287-297.
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talk on this (powerpoint)
42. Downs, S. G., and F. L. W. Ratnieks.
1999. Recognition of conspecifics by honeybee guards uses
nonheritable cues acquired in the adult stage. Animal Behaviour
58:643-648.
41. Foster, K. R., P. Seppa, F. L. W.
Ratnieks, and P. A. Thorén. 1999. Low paternity in
the hornet Vespa crabro indicates that multiple mating by
queens is derived in vespine wasps. Behavioral Ecology and
Sociobiology 46:252-257.
40. Monnin, T., and F. L. W. Ratnieks.
1999. Reproduction versus work in queenless ants: when to
join a hierarchy of hopeful reproductives? Behavioral Ecology
and Sociobiology 46:413-422.
39. Ratnieks, F. L. W., Anderson, C.
1999. Task partitioning in insect societies: natural history,
ergonomics, and information. Actes Colloq. Insectes Sociaux.
12: 1-10.
38. Ratnieks, F. L. W., and C. Anderson.
1999. Task partitioning in insect societies. Insectes Sociaux
46:95-108.
37. Ratnieks, F. L. W., and C. Anderson.
1999. Task partitioning in insect societies. II. Use of queueing
delay information in recruitment. American Naturalist 154:536-548.
36. Ratnieks, F. L. W., Keller, L. 1998.
Queen control of egg fertilization in the honey bee. Behavioral
Ecology and Sociobiology 44: 57-61.
35. Sundström, L., Ratnieks, F. L. W.
1998. Sex ratio conflicts, mating frequency and queen fitness
in the ant Formica truncorum. Behavioral Ecology 9:116-121.
34. Ratnieks, F. L. W., Boomsma, J.
J. 1997. On the robustness of split sex ratio predictions
in monogynous social Hymenoptera. Journal of Theoretical Biology
185: 423-439.
33. Ratnieks, F. L. W. 1996. Evolution
of unstable and stable biparental care. Behavioral Ecology
: 490-493. 
32. Ratnieks, F. L. W., Visscher, P.
K. 1996. Living with the Africanized bee...Sinaloao beekeepers
adapt pollination to Africanized bees. California Agriculture.
50 (4) 24-28. 
31. Boomsma, J. J., Ratnieks, F. L.
W. 1996. Paternity in eusocial hymenoptera. Philosophical
Transactions of the Royal Society of London, B. 351: 947-975.

30. Ratnieks, F. L. W., Vetter, R.,
Visscher, P. K. 1996. A polygynous overwintered nest of Vespula
pensylvanica from southern California. Insectes Sociaux
43: 401-410. 
29. Ratnieks, F. L. W. 1995. Evidence
for a queen-produced egg-marking pheromone and its use in
worker policing in the honey bee. Journal of Apicultural Research
34: 31-37. 
28. Ratnieks, F. L. W., Boomsma, J.
J. 1995. Facultative sex allocation by workers and the evolution
of polyandry by queens in social Hymenoptera. American Naturalist
145: 969-993.
27. Ratnieks, F. L. W., Visscher, P.
K, Vetter, R. 1995. Treating American foulbrood using gamma
radiation: a case history. American Bee Journal 135: 557-561.
26. Ratnieks, F. L. W., Miller, D. G.
1993. Natural history of polygynous Vespula vulgaris
colonies in California. Psyche 100: 43-50.
25. Ratnieks, F. L. W., Miller, D. G.
1993. Division of honey bee drones during swarming. Animal
Behaviour 46: 803-805. 
24. Morse, R. A., Layne, J. N., Visscher,
P. K., Ratnieks, F. 1993. Selection of nest cavity volume
and entrance size by honey bees in Florida. Florida Scientist
56: 163-167.
23. Ratnieks, F. L. W. 1993. Egg-laying,
egg-removal, and ovary development by workers in queenright
honey bee colonies. Behavioral Ecology and Sociobiology 32:
191-198. 
22. Reeve, H. K., Ratnieks, F. L. W.
1993. Queen conflict in polygynous societies: mutual tolerance
and reproductive skew. In: Keller, L. (ed). Queen number and
sociality in insects. Oxford University Press, Oxford. 
21. Ratnieks, F. L. W., Reeve, H. K.
1992. The structure of conflict in hymenopteran societies:
processes that reduce conflict in advanced eusocial species.
Journal of Theoretical Biology 158: 33-65.

20. Getz, W. M., Kaitala, V., Ratnieks,
F. L. W. 1992. Invasion of sibmating genes in diploid and
haplodiploid populations. Evolutionary Ecology 6: 312-330.
19. Ratnieks, F. L.
W. 1992. American foulbrood: A review of an important honey
bee disease. Bee World 73: 177-191.
18. Ratnieks, F. L. W. 1991. Evolution
of discriminatory aggression in marine invertebrates. Journal
of Theoretical Biology 152: 557-565. 
17. Emlen, S. T., Reeve, H. K., Sherman,
P. W., Wrege, P. H., Ratnieks, F. L. W., Shellman-Reeve, J.
1991. Adaptive versus non-adaptive explanations of behavior:
the case of alloparental helping. American Naturalist 138:
259-270.
16. Ratnieks, F. L. W., Piery, M., Cuadriello,
I. 1991. The natural nest of the Africanized honey bee (Hymenoptera,
Apidae) near Tapachula, Chiapas, Mexico. Canadian Entomologist
123: 353-359. 
15. Ratnieks, F. L. W. 1991. Africanized
bees: Natural selection for colonizing ability. Pages 119-135.
In: Spivak, M., Breed, M. D., Fletcher, D. J. C. (eds) The
Africanized honey bee. Westview Press, Boulder, CO. 
14. Ratnieks, F. L. W. 1991. Facultative
sex allocation biasing by workers in social Hymenoptera. Evolution
45: 281-292. 
13. Ratnieks, F. L. W. 1991. The evolution
of genetic cue diversity in social Hymenoptera. American Naturalist
137: 202-226.
12. Ratnieks, F. L. W., Reeve, H. K.
1991. The evolution of queen-rearing nepotism in social Hymenoptera:
effects of discrimination costs in swarming species. Journal
of Evolutionary Biology 4: 93-115.
11. Ratnieks, F. L. W. 1990. The evolution
of polyandry by highly eusocial bee queens: The importance
of progressive versus mass provisioning and the timing of
removal of diploid males. Behavioral Ecology & Sociobiology
26: 342-348. 
10. Ratnieks, F. L. W. 1990. Assessment
of queen mating frequency by workers in social Hymenoptera.
Journal of Theoretical Biology 142: 87-94.

9. Morse, R. A., Camazine, S., Ferracane,
M., Minacci, P., Ratnieks, F. L. W., Spielholz, J., Underwood,
B.A. 1990. The population density of feral honey bee colonies
(Hymenoptera, Apidae) in the city of Oswego, New York. Journal
of Economic Entomology 83: 81-83.
8. Ratnieks, F. L. W., Visscher, P.
K. 1989. Worker policing in the honey bee. Nature 342: 796-797.
7. Ratnieks, F. L. W., Nowakowski, J.
1989. Honey bee swarms accept bait hives contaminated with
American foulbrood disease. Ecological Entomology 14: 475-478.
6. Mumme, R. L., Koenig, W. G., Ratnieks,
F. L. W. 1989. Helping behaviour, reproductive value, and
the future component of indirect fitness. Animal Behaviour
38: 331-343.
5. Ratnieks, F. L. W. 1989. Sterilization
of American foulbrood contaminated hive equipment using gamma
irradiation. American Bee Journal 129: 191-193.
4. Kukuk, P. F., Eickwort, G. C., Raveret-Richter,
M., Alexander, B., Gibson, R., Morse, R. A., Ratnieks, F.
L. W. 1989. The importance of the sting in the evolution of
sociality in the Hymenoptera. Annals of the Entomological
Society of America 82: 1-5. 
3. Ratnieks, F. L. W. 1988. Reproductive
harmony via mutual policing by workers in eusocial Hymenoptera.
American Naturalist 132: 217-236.
2. Ratnieks, F. L. W. 1988. Improved
bait hives. American Bee Journal 128: 125-7.
1. Robinson, G. E., Ratnieks,
F. L. W. 1987. Induction of premature foraging by orally and
topically applied juvenile hormone analogs in the honey bee
(Apis mellifera L.). Journal of Economic Entomology
80: 784-787. |
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articles in conference proceedings
21. Ratnieks,
F. L. W. 2002. Parasitism of African honey bee colonies by
cape honey bee workers. Pages 81-83 in Garófalo, C.
(ed.) V Encontro sobre abelhas. Universidade de São
Paulo, Ribeirão Preto. 
20. Ratnieks,
F. L. W. 2002. Cooperation through coercion: policing of male
production and female caste fate in honey bees and stingless
bee. Pages 10-14 in Garófalo, C. (ed.) V Encontro sobre
abelhas. Universidade de São Paulo, Ribeirão
Preto. 
19. Ratnieks, F. L. W., Monnin, T. 2002.
Dominance hierarchies and conflict: insights from inclusive
fitness theory. Page 74 in: Proceedings XIV International
Congress of IUSSI, Sapporo, Japan.
18. Wossler, T. C., Oldroyd, B. P.,
Ratnieks, F. L. W. 2002. Reproductive anarchy in the beehive:
the role of pheromones. Page 213 in: Proceedings XIV International
Congress of IUSSI, Sapporo, Japan.
17. Martin, S. J., Ratnieks, F. L.
W., Jones, G. R.. 2002. How do honeybee workers discriminate
between worker-laid and queen-laid eggs? Page 124 in: Proceedings
XIV International Congress of IUSSI, Sapporo, Japan.
16. D'Ettorre, P., Heinze, J., Ratnieks,
F. L. W. 2002. Policing by egg eating in the ponerine ant
Pachycondyla inversa. Page 77 in: Proceedings XIV International
Congress of IUSSI, Sapporo, Japan.
15. Monnin, T., Ratnieks, F. L. W.,
Jones, G. R., Beard, R. 2002. Page 76 in: Proceedings XIV
International Congress of IUSSI, Sapporo, Japan.
14. Beekman, M, Ratnieks, F. L. W.
2002. Power over reproduction in eusocial Hymenoptera. Page
74 in: Proceedings XIV International Congress of IUSSI, Sapporo,
Japan.
13. The Mathematical Modelling of Social
Behaviour. EU 'INSECTS' TMR meeting, Great Hucklow, UK, 14-18
Sept. 2001, 113 pages. 
12. Ratnieks, F. L. W. 2001. Por que
las abejas Melipona crian reinas en exceso? El conflicto de
castas y el poder de las larvas sobre su propio destino (Why
do Melipona bees rear excess queens? Caste fate conflict and
the power of larvae over their fate). Pages 102-103 in: Quezada-Euán,
J., May-Itzá, W., Moo-Valle, H., Medina, J. (eds) II
Seminario Mexicano sobre abejas sin aguijón.
11. Paxton, R., Ruhnke, H., Shah, M.,
Bego, L. R., Quezada-Euán, J. J. G, Ratnieks, F. L.
W.2001. Social evolution in stingless bees: are the workers
or is the queen in control of male parentage? Pages 108-107
in: Quezada-Euán, J., May-Itzá, W., Moo-Valle,
H., Medina, J. (eds) II Seminario Mexicano sobre abejas sin
aguijón.
10. Ratnieks, F. L. W. 2001. Our best
theory: Novel predictions and tests on socila insect biology
arising from W. D. Hamilton;s theory of inclusive fitness
35 years on. Page 16 in: Menzel, R., Rademacher, E. Proceedings
of the 2001 Berlin meeting of the European sections of IUSSI
(International Union for the Study of Social Insects).
9. Ratnieks, F. L. W. 2001. Conflict
and caste determination. Page 100a in: Menzel, R., Rademacher,
E. Proceedings of the 2001 Berlin meeting of the European
sections of IUSSI (International Union for the Study of Social
Insects).
8. Ruhnke, H., Shah, M., Mecke, R.,
Bego, L. R., Quezada-Euán, J., Ratnieks, F. L. W.,
Paxton, R. J. 2001. Kin conflict and the parentage of males
in stingless bees. Page 58 in: Menzel, R., Rademacher, E.
Proceedings of the 2001 Berlin meeting of the European sections
of IUSSI (International Union for the Study of Social Insects).
7. Ratnieks, F. L. W. 1998. Information
sources in reproductive manipulation. Page 387 in: Schwarz,
M. P., Hogendoorn, K. (eds) Proceedings of the 13th International
Congress of IUSSI. IUSSI.
6. Monnin, T., Peeters, C., Ratnieks,
F. L. W. 1998. "Queen" policing and worker policing
in the queenless ant Dinoponera quadriceps. Page 323 in: Schwarz,
M. P., Hogendoorn, K. (eds) Proceedings of the 13th International
Congress of IUSSI. IUSSI.
5. Foster, K. R., Seppä, P., Ratnieks,
F. L. W., Thorén, P. 1998. Low paternity and queen-worker
conflict in the hornet. Page 168 in: Schwarz, M. P., Hogendoorn,
K. (eds) Proceedings of the 13th International Congress of
IUSSI. IUSSI.
4. Ratnieks, F. L. W. 1996. Worker
reproduction and worker policing: how the balance is set.
p. 124, Abstracts of: Fifth International Conference of Systematic
and Evolutionary Biology.
3. Ratnieks, F. L. W. 1994. Worker policing
in honey bees. page 582 in: Lenoir, A., Arnold, G., Lepage,
M. (eds) Les Insectes Sociaux. (Proceedings of the 12th International
Congress of IUSSI). Université Paris Nord, Paris.
2. Ratnieks, F. L. W. 1992. Evidence
for an egg-marking pheromone in the honey bee. American Bee
Journal 132: 813.
1. Ratnieks, F. L. W. 1987. The adaptive
significance of nest site selection by honey bee swarms: calculation
of reproductive value of swarms using colony life table data.
Page 617 in: J. Eder and H. Rembold (eds) Chemistry and biology
of social insects. Verlag J. Peperny, Munich.
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