Professor Stephen Martin

Chair in Social Entomology

  • Peel Building Room G48
  • T: +44 (0)161 295 2476
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Office Times

Thursday 2pm - 4pm 


My first studies in biology included bird-ringing and collecting insects in the UK and China. I completed my BSc at Bangor University in Wales, followed by MSc from Shinshu University in Japan on beetles as environmental indicators of habitat disturbance. I also studied hornets which led onto my PhD where I developed my interests in social insects, which was furthered when I returned to Japan to conduct research into the population dynamics of hornets on a small sub-tropical island. On returning to the UK I worked for the National Bee Unit for seven years researching the pests and pathogens of honeybees, where I became internationally known for my work in the parasitic Varroa mite.

I then returned to University and spent 12 years at Sheffield elucidation chemical recognition systems in social insects and continuing my work into honeybee pathogens mainly in Hawaii. In 2012, I moved over to the University of Salford in order to continue my research and help educate the next generation of scientists.


My teaching is in the area of zoology, animal behaviour and chemical ecology. It is closely linked to and driven by my research. I see teaching is a two-way process allowing for transfer of knowledge and expertise to the next generation from which the birth of new ideas and continually questioning current ideas is generated. I am very active outside university giving talks to beekeeper groups and other stakeholders in industry all around the world.

Research Interests

Invertebrates among most diverse and important group of animals. I have always had a deep interest in this group and their ecological importance in ecosystems. Of these, the social insects (bees, wasps, ants and termites) are the most biologically important due to numerical dominance.  My research focuses on various aspects of their biology since social insects provide excellent model systems with which to test a wide range of behaviours from conflict to cooperation. 

Currently there is a worldwide decline in pollinating insects and I am looking at the molecular changes associated with increased virulence of an emerging viral pathogen using the honeybee, Varroa mite and deformed wing virus as a model system. Recently we have shown in Hawaii that the new viral transmission route caused by the parasitic Varroa mite has selected for a viral strain that has led to the global death of millions of honeybee colonies. The impact of this study has helped stakeholders understand the problem and has influenced policy at the European Parliament. Currently we are looking at ways mite tolerance has arisen in various honey bee populations.

I am also investigating the underlying mechanisms of phenotypic plasticity in chemical recognition systems in ants, using Formcia ants as the model system. Understanding how phenotypic variation is generated and then maintained is one of the major challenges currently facing biologists.

We have developed over the past 8 years a detailed understanding of the chemical recognition system in the ant Formcia exsecta and are currently working out the molecular mechanisms that generate phenotypic variation, using the latest ideas and tools in molecular biology.

Qualifications and Memberships

BSc (Zoology inc. Applied Zoology, 2i) University of Wales 1983

MSc (by research) Shinshu University Japan 1987

PhD University of Wales 1989

DSc University of Wales, 2006

Fellow of the Royal Entomological Society F.R.E.S.

Executive Board member of International Bee Research Association

NERC peer review collage member

Editorial Board Member of Applied and Environmental Microbiology

Editorial Board Member of Plos One


137.    Brettell, LE., Mordecai, GJ.,Pachori, P., Martin SJ .(2017) Novel RNA virus genome discovered in Ghost ants (Tapinoma melanocephalum) from Hawaii.. Genome Announcements             (genomeA00669-17R1)

136.     Brettell LE, Mordecai GJ, Schroeder DC, Jones IM, da Silva, JR, Vicente-Rubiano M, Martin SJ  (2017) A Comparison of Deformed Wing Virus in Deformed and Asymptomatic Honey Bees Insects, 8, 28; doi:10.3390/insects8010028

135.     Brettell LE, and Martin SJ (2017) Oldest Varroa tolerant honey bee population provides insight into the origins of the global decline of honey bees. Scientific Reports (in press).

134.  Hall JL, Alpers K, Bown KJ, Martin SJ, Birtles RJ (2017) Epidemiological crowdsourcing; using mass-participation outdoor events to assess human exposure to tick-borne pathogens. Emerg            Infect Dis (in press).

133.    Benaets K, Van Geystelen A, Cardoen D, De Smet L, de Graaf  DC, Schoofs L, Larmuseau MHD, Brettell LE, Martin SJ, Wenseleers T (2017) Covert deformed wing virus infections have long-term deleterious effects on honeybee foraging and survival. Proceedings of the Royal Society B (in press)

132.    Martin SJ, and Correia-Oliveira ME (2016) The occurrence of ecto-parasitic Leptus sp. mites on        Africanised honey bees. Journal of Apicultural Research 55, 243-246.

131.     Mordecai GJ, Brettell L, Pachori P, Villalobos E, Martin SJ, Jones IM, & Schroeder, DC (2016) A new Iflavirus, Moku Virus, found in wasps, honey bees and Varroa. Scientific Reports, 6:34983

130.    Guillem, R. M., Drijfhout, F. P. and Martin, S. J. (2016) Species-specific cuticular hydrocarbon stability within European Myrmica ants. Journal of Chemical Ecology 42, 1052–1062.

129.    Wilfert, L., Long, G., Leggett H.C., Schmid-Hempel. P., Butlin, R., Martin, SJ and Boots, M (2016) Deformed Wing Virus is a Recent Global Epidemic in Honeybees driven by Varroa Mites. Science 351: 594-597

128. * Mordecai GJ, Brettell L, Martin SJ, Dixon D, Jones IM, & Schroeder, DC (2016) Superinfection exclusion and the long-term survival honey bees in Varroa-infested colonies. ISEM J 10, 1182-1191 doi: 10.1038/ismej.2015.186

127.  *Mordecai GJ, Wilfert L, Martin SJ, Jones IM &  Schroeder, DC (2016) Diversity in a honey bee pathogen: first report of a third master variant of the  Deformed Wing Virus quasispecies ISME J 10, 1264-1273  doi: 10.1038/ismej.2015.178

126.  Kather, R. & Martin, S.J. (2015) Evolution of Cuticular Hydrocarbons in the Hymenoptera: A Meta-analysis. Journal Chemical Ecology 41,871-883

125.  Pradella, D., Martin, S.J., Dani, F.R.(2015) Using errors by guard honeybees (Apis mellifera) to gain new insights into nest-mate recognition signals. Chemical Senses 40:649-653.

124.  Martin, S.J. (2015) Does a remote Brazilian Island hold the key to Varroa tolerance? American Bee         Journal (Nov 1217-1218)

123.  Kather, R., Drijfhout, F. P., Shemilt, S. and Martin, S.J. (2015) Evidence for colony-specific differences in chemical mimicry in the parasitic mite Varroa destructor. Chemecology 25; 215-222 DOI: 10.1007/s00049-015-0191-8

122. Kather, R., Drijfhout, F. P., Shemilt, S. and Martin, S.J. (2015) Evidence for Passive Chemical Camouflage in the Parasitic Mite Varroa destructorJournal of Chemical Ecology 41. 178-186. DOI: 10.1007/s10886-015-0548-z

121. Martin SJ & Bayfield J (2014) Is the bee louse Braula coeca (Diptera) using chemical camouflage to survive within honeybee colonies? Chemoecology 24:165-169. DOI: 10.1007/s00049-014-0158-1

120. Helanterä, H, Martin, SJ, Ratnieks FLW, (2014) Recognition of nestmate eggs in the ant Formica fusca is based on queen derived cues J. Current Zoology 60, 131-136

119.  Guillem, RM, Drijfhout, F, Martin SJ (2014) Chemical deception among ant social parasites. J. Current Zoology 60, 62-75.

118. Martin SJ, Shemilt S, Trontti K. (2014) Nest-mate recognition cues are not used during or influenced by mating in the ant  Formica exsectaEthology Ecology and Evolution 26, 40-48.

117.  Martin SJ, Vitikainen E, Shemilt S, Drijfhout FP, Sundstrom L (2013) Sources of variation in cuticular hydrocarbons in the ant Formica exsecta? Journal Chemical Ecology 39, 1415-1423.

116. Martin SJ, Carreck NL, Ball BV. (2013) The role of Deformed wing virus in the collapse of honeybee colonies. Journal of Apicultural Research 52, 251-258.

115.  Sutton P, Wilde MJ, Martin SJ, Cvacka J., Vrkoslav, V., Rowland, SJ (2013) Studies of long chain lipids in insects by (HT) GC and HTGC-MS.  J. Chrom A 1297, 236-240

114.  Badouin, H., Belkhir, K., Gregson, E.,  Galindo, J., Sundström L., Martin SJ, Butlin, RK, Smadja          CM (2013) Transcriptome characterisation of the ant Formica exsecta with new insights into the evolution of desaturase genes in social Hymenoptera. PLosOne doi:10.1371/journal.pone.0068200

113.  Martin SJ., Hardy J, Villalobos E, Martin-Hernandez R, Nikaido S, Higes M. (2013) Do the honeybee pathogens Nosema ceranae and deformed wing virus act synergistically? Environmental Microbiology Reports 5, 506-510 doi:10.1111/1758-2229.12052

112.   Martin SJ (2013) Double trouble in paradise: Small Hive Beetle joins Varroa in Hawaii. American Bee Journal. 153, 529-532.

111.   Dietemann V, Nazzi, F, Martin SJ, Anderson D, Locke B, Delaplane KS, Wauquiez Q, Tannah ll C, Frey C, Ziegelmann B, Rosenkranz P, Ellis JD (2013) Standard methods for varroa research. In V Dietemann; J D Ellis; P Neumann (Eds) The COLOSS BEEBOOK, Volume II, standard methods for Apis mellifera  pest and pathogen research. Journal of Apicultural Research 51(5):

110.* Schroeder DC, Martin SJ (2012) Deformed Wing Virus, the main suspect in unexplained honeybee deaths worldwide. Virulence 3, 1-3.

109.**  Martin SJ, et al.  (2012) Global honeybee viral landscape altered by a parasitic mite. Science 336, 1304-1306.

108. * Martin SJ, Trontti K, Shemilt S,  Drijfhout FP, Butlin, R, Jackson, D. (2012) Weak patriline effects are present in the cuticular hydrocarbon profiles of isolated Formica exsecta ants but they disappear in the colony environment. Ecology and Evolution 2:2333-2346.

107.  Choi M-B, Martin SJ, Lee J-W. (2012) Distribution, spread and impact of the invasive hornet Vespa velutina in South Korea. Journal of Asia-Pacific Entomology 15, 473-477.  

106.  Martin SJ, Shemilt S, Drijfhout FP (2012) Effect of time on colony odour stability in the ant Formica exsecta.  Naturwissenschaften 99, 327-331.

105.  Kather R, Martin SJ (2012) Cuticular hydrocarbon profiles as taxonomic tool, advantages, limitations and technical aspects. Physiological Entomology 37, 25-32.

104.* Guillem RM, Drijfhout F, Martin SJ (2012) Using chemo-taxonomy of host ants to help conserve the Large Blue butterfly. Biological Conservation 148.39-43.

103.* Martin SJ, Vitikainen E, Drijfhout FP, Jackson D. (2012) Conspecific ant aggression is correlated with chemical distance, but not with genetic or spatial distance.  Behav. Genetics. 42, 323-331.

102. Helanterä H, Lee YR, Drijfhout FP, Martin SJ (2011) Genetic diversity, colony chemical phenotype and nestmate recognition in the ant Formica fusca. Behavioural Ecol. 22, 710-716.

101. Seppa P, ... Martin SJ,... Sundstrom L (2011) The many ways to delimit species, hairs, genes and surface chemistry. Myrmecological News 15, 31-41.

100.  Kather R, Drijfhout FP, Martin SJ (2011) Task group differences in cuticular lipids in the honey bee Apis melliferaJ. Chem. Ecol. 37, 205-212.

99. * Martin SJ, Helanterä H, Drijfhout FP (2011) Is Parasite pressure a driver of chemical cue diversity in ants? Proceedings of the Royal Society B 278, 496-503.

98.    Jackson DE, Martin SJ (2010) Sex Allocation, Size matters for red spider Mites. Current Biology 20, 24 R1080

 97. Martin SJ, Carruthers, JM, Williams PH, Drijfhout, FP (2010) Host specific social parasites (Psithyrus) indicate chemical recognition system in Bumblebees. J. Chem. Ecol. 36, 855-863.

96.    Takahashi J, Yamasaki K, Mitsuhata M, Martin SJ, Ono M, Tsubaki Y (2010) Bumblebee fauna of the Nemuro peninsula, Hokkaido, Japan, with special reference to the effect of the invasive Bombus terrestris on the rare B. florilegus. Japanese Conservation Ecology 15, 101-110.

95.    Martin SJ (2010) Trouble in paradise, Varroa in Hawaii. American Bee Journal 381-384.

94.    Couvillon M, Hughes W, Perez-Sato J, Martin SJ, Roy G, Ratnieks FLW (2010) Sexual selection in honeybees, colony variation and the importance of size in male mating success. Behavioral Ecology 21, 520-525 (doi,10.1093/beheco/arq016).

93. *  Martin SJ, Ball BV, Carreck NL (2010) Prevalence and persistence of  deformed wing virus (DWV) in untreated or acaricide-treated Varroa destructor infested honey bee (Apis mellifera L.) coloniesJournal of Apicultural Research 49, 72-79.

92.    Carreck NL, Ball BV, Martin SJ (2010) The epidemiology of cloudy wing virus infections in honey bee colonies in the UK. Journal of Apicultural Research 49, 66-71.

91.  * Carreck NL, Ball BV, Martin SJ (2010) Honey bee colony collapse and changes in viral prevalence associated with Varroa destructor. Journal of Apicultural Research 49, 93-94.

90.   *Highfield AC, Nagar AEI, Mackinder LCM, NoëlJM-L, Hall MJ, Martin SJ, Schroeder DC (2009). Deformed wing virus implicated in over-wintering honeybee colony losses. Appl. Environ. Microbiol. 75, 7212-7220.

89.   *Martin SJ, Drijfhout FP (2009) A review of ant cuticular hydrocarbons J. Chem. Ecol. 35, 1151-1161. DOI 10.1007/s10886-009-9695-4.

88.    Martin SJ, et al (2009) Polygyny reduces rather than increases nestmate discrimination cue diversity in Formica exsecta ants. Insectes Sociaux 56, 375-383. DOI 10.1007/s00040-009-0035-z

87.   Chauzat M-P, Martin SJ (2009) A foreigner in France, Biological information on the Asian hornet Vespa velutina, a recently introduced species. Biologist 56, 86-91

86.   Takahashi J, Martin SJ, Ono M, Shimizu I (2010) Male production by non-natal workers in the bumblebee, Bombus deuteronymus (Hymenoptera, Apidae). J. Ethol. 28, 61-66. 

85.   Koyama S, Takagi T, Martin SJ, Yoshida T, Takahashi J (2009) Absence of reproductive             conflict during queen rearing in Apis cerana. Insectes Sociaux 56,171-175.

84.   Martin SJ, Drijfhout FP (2009) Nestmate and task cues are influenced and encoded differently within ant cuticular hydrocarbon profiles. J. Chem. Ecol. 35, 368-374.

83.  *Martin SJ, Drijfhout FP (2009) How reliable is the analysis of complex cuticular hydrocarbon profiles by multi-variate statistical methods? J. Chem. Ecol. 35, 375-382.

82.   Perez-Sato JA, Chaline N, Martin SJ, Hughes WOH, Ratnieks FLW (2009) Multi-level selection for hygienic behaviour in honeybees. Heredity DOI 10.1038/hdy.2009.20

81.   Martin SJ, Weihao Z, Drijfhout FP (2009) Long-term stability of cuticular hydrocarbons facilitates chemotaxonomy. Biological Journal of the Linnean Society 96, 732-737.

80.    Waghchoure-Camphor ES, Martin SJ (2009) Population changes of Tropilaelaps clareae mites in Apis mellifera colonies in Pakistan. J. Apic. Res. 48, 46-49 DOI 10.3896/IBRA.

79.    Martin SJ, Takahashi J, Katada S (2009) Queen condition, mating frequency, queen loss and worker reproduction in the hornets Vespa affinis and V. simillimaEcol. Entomol. 34, 43-49.

78.    Waghchoure-Camphor ES, Martin SJ (2008) Beekeeping in Pakistan, a bright future in a troubled land. American Bee Journal 148, 726-728.

77.* Martin SJ, Helanterä H,  Drijfhout FP (2008) Colony-specific hydrocarbons identify nest mates in two species of Formica ant. J. Chem Ecol. 34, 1072-1080. Doi 10.1007/s10886-008-9482-7.

76.   * Martin SJ, Vitikainen E, Helanterä H, Drijfhout FP (2008) Chemical basis of nestmate recognition in the ant Formica exsectaProceedings of the Royal Society B 275, 1271-1278.

75.    Martin SJ, Takahashi J, Ono M, Drijfhout F P (2008) Is the social parasite Vespa dybowskii using chemical transparency to get her eggs accepted? J. Ins. Phys. 54, 700-707.

74.    *Martin SJ, Helanterä H, Drijfhout FP (2008) Evolution of Species-Specific Cuticular Hydrocarbon Patterns in Formica Ants. Biological Journal of the Linnean Society 95, 131-140.

73.    *Martin SJ, Jenner EA, Drijfhout FP (2007) Chemical deterrent enables a socially parasitic ant to invade multiple hosts. Proceedings of the Royal Society B 274, 2717-2721.

72. Helantera H, Martin SJ, Ratnieks FLW (2007) Prior experience with eggs laid by non-Nestmate queens induces egg acceptance errors in ant workers. Behavioral Ecol. Sociobiol. 62, 223-228.

71.    Beekman, M., Martin SJ, Drijfhout F, Oldroyd B (2007) Higher removal rate of eggs laid by anarchistic queens-a cost of anarchy? Behavioral Ecology & Sociobiology 61, 1847-1853.

70.    Takahashi J, Inomata K, Martin SJ (2007) Mating structure and male production in Vespaanalis and V. simillima (Hymenoptera, Vespidae). Entomological Science 10, 223-229.

69.    Jackson D, Martin SJ, Ratnieks FLW, Holcombe M. (2007) Spatial and temporal variation in pheromone composition of ant foraging trails. Behavioral Ecology 18, 444-450.

68.    Martin SJ, Takahashi, Drijfhout FP (2007) Setosa membrane structure and occurrence of eicosenol in honeybees (Apis). Apidologie 38, 104-109.

67.    Martin SJ (2006) Early collapse of Vespa simillima (Hymenoptera, Vespidae) colonies in central Japan.  Entomological Science 9, 373-376.

66.    Mondragon M, Martin SJ, Vandame R (2006) Mortality of mite offspring, a major component of Varroa  destructor resistance in Africanized honeybees.  Apidologie 37, 67-74.

65.    Jensen AB, Palmer KA, Chaline N, Raine NE, Martin SJ, Tofilski A, Pedersen BV, Boomsma JJ, Ratnieks FLW (2005) 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. Conservation genetics 6, 527-537.

64.    Martin SJ, Châline N, Drijfthout F, Jones GR (2005) Role of esters in egg removal in honeybee colonies. Behavioral Ecology & Sociobiology 59, 24-29

63.    Jackson DE, Martin SJ, Holcombe M, Ratnieks FLW (2006) Longevity and detection of persistent foraging trails in Pharaoh's ants, Monomorium pharaonisAnim. Beh 71, 351-359.

62.    *Chaline N, Sandoz JC, Martin SJ, Ratnieks FLW, Jones GR (2005) Learning and discrimination of individual cuticular hydrocarbons by honey bees (Apis mellifera). Chemical Senses 30, 327-333.

61.    Martin SJ, Dils V, Billen J (2005) Morphology of the Dufour's gland within the honey bee sting gland complex. Apidologie 36, 543-436.

60.    Chaline N, Martin SJ, Ratnieks FLW (2004) Absence of nepotism towards imprisoned young queens during swarming in the honey bee. Behav. Ecol. 16, 403-409.

59.    Martin SJ, Chaline N, Jones G, Ratnieks FLW (2004) Searching for the egg-marking signal in honeybees. Journal of Negative Results 2, 1-10.

58.    Martin SJ (2004) Acaricide (pyrethroid) resistance in Varroa destructorBee World 85, 67-69.

57.    Martin SJ, Chaline N, Jones G, Ratnieks FLW (2004) Role of hydrocarbons in egg recognition in the honeybee.  Physiological Entomology 29, 395-399.

56.    Martin SJ (2004) Biological control of social wasps (Vespinae) using mermitid nematodes New Zealand Journal of Zoology 31, 241-248.

55.    Martin SJ, Chaline N, Jones G, Oldroyd B, Ratnieks FLW (2004) Egg marking pheromones of anarchistic worker honeybees (Apis mellifera). Behav. Ecol. 15, 839-844.

54.    Chaline N, Martin SJ, Ratnieks FLW (2004) Worker policing persists in a                hopelessly queenless honey bee colony (Apis mellifera). Insectes Sociaux 51, 113-116.

53.    Martin SJ, Jones G (2004) Conservation of bio-synthetic pheromone pathways in honeybees Apis.  Naturwissenschaften 91, 232-237.

52.    *Martin SJ, Medina LM (2004) Africanized honeybees possess unique tolerance to Varroa mitesTrends in Parasitology 20, 112-114.

51.    Takahashi J, Akimoto S, Martin SJ, Tamukae M, Hasegawa E (2004) Mating structure and male production in the giant hornet Vespa mandariniaApplied Entomology & Zoology 39, 343-349.

50.    *Sumpter D, Martin SJ (2004) The dynamics of virus epidemics in Varroa-infested honeybee colonysJournal of Animal Ecology 73, 51-63.

49.    Correa-Marques MH, Medina LM, Martin SJ, DeJong D (2003) Comparing data on the reproduction of Varroa destructor.  Genetics & Mol. Res. 2, 1-6.

48.    Martin SJ, Kryger P (2002) Impact of Varroa destructor on honeybees in South Africa. Apiacta 37, 60-65.

47.    Martin SJ, Elzen PJ, Rubink WR (2002) Effect of acaricide resistance of reproductive ability of the honey bee mite Varroa destructor. Exp. Appl. Acarol. 27, 195-207    

46.    Medina ML, Martin SJ, Montaño L, Ratnieks FLW (2002) Reproduction of Varroa destructor in worker brood of Africanized honey bee (Apis mellifera). Exp. Appl. Acarol. 27, 79-88.

45.   **Martin SJ, Beekman M, Wossler TC, Ratnieks FLW (2002)  Parasitic Cape honeybee workers,         Apis mellifera capensis, evade policing.  Nature 415, 163-165. 

44.    Martin SJ, Jones GJ, Ratnieks FLW, Middleton H, Châline N (2002) Reassessing the role of the honeybee (Apis mellifera) Dufour’s gland in egg  marking. Naturwissenschaften 89, 528-532.

43.    Beekman M, Wossler TC, Martin SJ, Ratnieks FLW (2002) Parasitic Cape honey bee workers are not given differential treatment by African guards. Insectes Sociaux 49, 216-220.

42.    Martin SJ, Kryger P, Wossler TC (2002) Usurpation of African Apis mellifera scutellata colonies by  A. m. capensis workers.  Apidologie 33, 215-232.

41.    Martin SJ, Kryger P (2002) Reproduction of Varroa destructor in South African honey bees, does cell space influence Varroa male survivorship?  Apidologie 33, 51-61.

40.  ** Martin SJ (2001)  The role of Varroa and viral pathogens in the collapse of

honey bee colonies, a modelling approach. Journal of Applied Ecology 38, 1082-1093.

39.    Martin SJ (2001)  Varroa destructor reproduction during the winter in Apis mellifera colonies in UK. Exp. Appl. Acarol. 25, 321-325.

38.    Martin SJ (2001)  Saving Honey bees in the Kruger National Park, American Bee J. 141, 343-346.

37.    Medina ML, Martin SJ (1999)  A comparative study of Varroa jacobsoni reproduction in worker cells of honeybees in England and Africanized bees in  Mexico. Exp. Appl. Acarol. 23, 659-667.

36.    Martin SJ  (1998)  A population dynamic model of the mite Varroa jacobsoni.  Ecological Modelling 109, 267-281.

35.   Martin SJ, Hogarth A, van Breda J, Perrett J (1998)  A scientific note on Varroa jacobsoni and the collapse of Apis mellifera colonies in the United Kingdom.  Apidologie 29, 369-370.

34.  ** Bowen-Walker PL, Martin SJ, Gunn A (1998)  The transmission of deformed wing virus between honey bees (Apis mellifera L.) by the ecto-parasitic mite Varroa jacobsoni Oud.  Journal of Invertebrate Pathology 73, 101-106.

33     Sampson C, Martin SJ (1998)  Varroa control on Jersey  Bee Biz 10, 19-21.

32.    Martin SJ (1999) Population modelling and the production of a monitoring tool for Varroa jacobsoni an ectoparasitic mite of honey bees.  Aspects of Applied Biology 53, 105-112.

31.    Bowen-Walker PL, Martin SJ, Gunn A (1997)  Preferential distribution of the parasitic mite, Varroa jacobsoni Oud. on overwintering honeybee (Apis mellifera L.) workers and changes in the level of parasitism.  Parasitology 114, 151-157.

30.   * Martin SJ, Holland K, Murray M (1997)  Non-reproduction in the honeybee mite Varroa jacobsoni.   Exp. Appl. Acarol. 21, 539-549.

29.    Martin SJ (1997)  Varroa jacobsoni population biology research in the UK.  American Bee Journal 137, 382-385.

28.    Lobb N, Martin SJ (1997)  Mortality of the mite Varroa jacobsoni during or soon after the emergence of worker or drone honeybees.  Apidologie 28, 367-374.

27.    Martin SJ, Kemp D (1997)  Average number of reproductive cycles performed by the parasitic mite Varroa jacobsoni in Apis mellifera colonies. J. Apicultural Research 36, 113-123.

26.    Martin SJ, Cook C (1996)  Effect of host brood type on number of offspring produced by the honeybee parasite Varroa jacobsoniExp. Appl. Acarol., 20, 387-390.

25.    Martin SJ (1995)  Comparison of tropical and sub-tropical colonies of the hornet Vespa affinis.  Asian Journal of Tropical Biology 2, 13-19.

24.    Martin SJ (1995)  Hornets of Malaysia. Malayan Nature Journal, 49, 71-82.

23.    Martin SJ (1995)  Colony development of the hornet Vespa affinis.Jap. J. Entomol. 63, 861-876.

22.    Martin SJ (1995)  Reproduction of Varroa jacobsoni in cells of Apis mellifera containing one or more mother mites and the distribution of these cells.  J. Apicultural Research 34, 187-196.

21.    *Martin SJ (1994)  Ontogenesis of the mite Varroa jacobsoni Oud.  in worker brood of the honeybee Apis mellifera L. under natural conditions.  Exp. Appl.   Acarol. 18, 87-100.

20.   * Martin SJ (1995)  Ontogenesis of the mite Varroa jacobsoni Oud.  in the drone brood of the honeybee Apis mellifera L. under natural conditions.  Exp. Appl. Acarol. 19, 199-210.

19.    Martin SJ (1993)  Weight changes in adult hornets, Vespa affinis.  Insectes Sociaux 40, 363-368.

18.    Martin SJ (1993)  Two nests of the hornet Vespa wilemani.  Japanese J. Entomology  61, 679-682.

17.    Martin SJ (1993)  Porbeagle sharks in Falkland waters.  Elasmo News 1,  6-7.     

16.    Martin SJ (1992)  Development of the embryo nest of Vespa affinis (Hymenoptera, Vespidae) in southern Japan.  Insectes Sociaux 39, 45-57.

15.    Martin SJ (1992)  Colony defence against ants in VespaInsectes Sociaux 39, 99-113.

14.    Martin SJ (1992)  Seasonal and altitudinal distribution of Ground Beetles (Coleoptera) in the Southern Alps of Japan.  Japanese Journal of Entomology 60, 26-38.

13.    Martin SJ (1992) Nest thermoregulation in Vespa affinisJapanese J. Entomology 60, 483-486.

12.    Martin SJ (1992)  Occurrence of the Pyralid Moth Hypsopygia mauritialis (Lepidoptera, Pyralidae) in the nests of Vespa affinis.  Japanese J. Entomology 60, 267-270.

11.    Martin SJ (1992)  Colony failure in Vespa affinisJapanese .J Entomology 60, 671-681.

10.    Martin SJ (1991)  A simulation model for colony development of the hornet Vespa simillima (Hymenoptera, Vespidae).  Japanese Journal of Entomology 59, 105-124.

9.      Martin SJ (1991)  Notes on the over-wintering of the Carpenter Bee Xylocopa albinotum (Hymenoptera, Anthophoridae).  Japanese Journal of Entomology 59, 735-741.

8.      Martin SJ (1990)  Nest thermoregulation in Vespa simillima, V. tropica and V. analis.  Ecological Entomology 15, 301-310.

7.      Martin SJ (1990)  Usurpation by conspecific queens of Vespa simillima xanthoptera Cameron (Hymenoptera, Vespidae).  Ento. Mon. Mag. 126, 71-73.

6.      Martin SJ (1990)  Queenless nests of Vespa simillima  Jap. J. Entomol. 58, 347-354.

5.      Martin SJ (1990)  Thermoregulation in the Japanese hornet, Vespa mandarinia Smith (Hymenoptera, Vespidae).  Ento. Mon. Mag. 126, 103-109.

4.      Martin SJ (1989)  Altitudinal distribution of Burying Beetles (Coleoptera, Silphidae) in the Southern Alps of Japan.  Japanese Journal of Entomology 57, 876-879.

3.      Martin SJ (1988)  Occurrence of Anthrax distigma (Diptera, Bombyliidae) in the nest of Vespa simillima xanthoptera (Hymenoptera, Vespidae).  Kontyu, 56, 461-462.

2.      Martin SJ (1988)  Thermoregulation in Vespa simillima xanthopteraKontyu, 56, 674-677.

1.      Martin SJ (1984)  Size difference between the sexes of Aphodius prodromus Ento. Mon. Mag., 120, 243-244.


Due to the limitation of space only a small selection of my10 chapters in books or booklets, 29  Refereed Conference Papers and36 general articles are presented.  

Martin, S. J. (2015) Chemical communication in Formica wood ants...........

Martin, S. J. (2011) Varroa Biology. In: Varroa - Still a problem in the 21st Century? Ed. N. Carrack & S Jones. IBRA, pages 5-10.

Drijfhout, F. P., Kather, R., Martin, S. J. (2009) The role of cuticular hydrocarbons in insects.

In: Behavioural and Chemcial Ecology. Eds Wen Zhang and Hong Liu. Nova Science Publishers, Inc. ISBN: 978-1-60741-099-7. pages 1-24.

Martin, S. J.  (2001)  Biology & Life History of Varroa mites. in Mites of the     HoneyBee, eds. T. C. Webster & K. S. Delaplane, Dadant publication pp. 131-148.

Martin, S. J.  (1998) Varroa jacobsoni : monitoring and forecasting mite populations within honey bee colonies in Britain.  MAFF publications.  12 pp. (20,000 copies in English and Welsh produced)

Martin, S. J. (1998) The Varroa Calculator, MAFF publications. (20,000 copies)

Martin, S. J. (1997) Life and death of Varroa.  In Varroa! Fight the mite.  P. Munn  & R. Jones (eds.), pp 3-10.  International Bee Research Association.  Cardiff.

Shemilt S., Martin, S. J. & Drijfhout F. (2010) Source & maintenance of cuticular hydrocarbons in the ant Formica exsecta.. XVI IUSSI Congress Denmark.

Martin, S. J. & Drijfhout F. (2006) Breaking into the fortress, the egg recognition barrier. XV IUSSI Congress Washington.

 Martin, S. J. (2004) Honey bee diseases, are they a threat to Mexican beekeeping? XVIII seminario americano de apicultura, Villahermosa, Tabasco, Mexico.

Martin, S. J., Ball, B. V., Carreck, N. (2003) The role of Deformed wing virus in the mortality of Varroa infested honeybee colonies. XXXVIIIth International Apicultural Congress, Slovenia.

Martin, S. J. & Medina, L. M. (2003) Varroa tolerance in Africanisied honey bees explained. XXXVIII th International Apicultural Congress, Ljubljana, Slovenia.

Martin, S. J. & Medina, L. M. (2003)  Africanisied bee tolerance to Varroa. 10th International Congress of Actualisation in Beekeeping, Tlaxcala, Mexico.

Martin, S. J., (2003) Interaction between honeybees, bee viruses and Varroa haplotypes. 10th International Congress of Actualisation in Beekeeping, Tlaxcala, Mexico.

Martin, S. J., Ratnieks F. W. Jones. G. (2002) How do honeybee workers discriminate between worker-laid and queen-laid eggs? XIV International Congress of the International Union for the study of social Insects, Hokkaido Univ. Sapporo, Japan, p.  124.

Martin, S.J. (2001) A graphic computer model which generates realistic brood patterns found in social hornet (Vespa) colonies despite using simple rules.  From Individual to collective behaviour in biological systems, Newton Institute, Cambridge, UK.

Martin, S. J. & Kryger, P. (2001) Impact of Varroa destructor on the honey bees of South Africa. XXXVI th International Apicultural Congress, Durban, South Africa.

Martin, SJ. (1999) The virus, mite and model: a useful approach towards Varroa control. XXXVI International Apicultural Congress, Vancouver, Canada: p 141-142.

Martin, SJ. & Ball, B. V. (1998) Variations in the virulence of Varroa infestations. XIII International Congress of the International Union for the study of social Insects, Australia.

Martin, S.J. (2012) Hawaii provides a key piece of the Varroa Virus puzzle. BBKA News, 208, 17-19.

Martin, S.J. (2012) The Butterfly, Owl or Lizard in memory of Dr. D. Stradling. Antenna 36, 219-220.

Martin, S.J. (2010) Hawaii, ants, viruses, and honeybees (Honeybee research at Sheffield). 100th special issue of Beekeepers Quarterly Magazine.

Martin, S. J. (2003) Is the capensis clone a threat to beekeeping? BBKA news April.

Martin, S. J. (1997) The Varroa model.  Bee Craft, 79: 360.

Martin, S. J. (1993-2000) seven key government reports.