Snails and their trails: The multiple functions of trail-following in gastropods

Biological Reviews

Volumn 88, Issue 3, 2013, Pages 683-700

  Ng, Terence P T ^a   Saltin, Sara H ^b   Davies, Mark S ^c   Johannesson, Kerstin ^b   Stafford, Richard ^d   Williams, Gray A ^a  

^a UNIVERSITY OF HONG KONG   (Hong Kong)

^b UNIVERSITY OF GOTHENBURG   (Sweden)

^c UNIVERSITY OF SUNDERLAND   (United Kingdom)

^d UNIVERSITY OF BEDFORDSHIRE   (United Kingdom)

Author keywords

Communication; Gastropod; Homing; Mate search; Mucus; Pheromone; Self organisation

Indexed keywords

PHEROMONE;

ANIMAL; ANIMAL BEHAVIOR; ARTICLE; GASTROPOD; INTERPERSONAL COMMUNICATION; MATE SEARCH; MOTOR ACTIVITY; MUCUS; PHYSIOLOGY; SELF-ORGANISATION;

COMMUNICATION; GASTROPOD; HOMING; MATE SEARCH; MUCUS; PHEROMONE; SELF-ORGANISATION;

ANIMALS; BEHAVIOR, ANIMAL; GASTROPODA; MOTOR ACTIVITY; MUCUS;

ANIMALIA; GASTROPODA; METAZOA;

EID: 84880037098     PISSN: 14647931     EISSN: 1469185X     Source Type: Journal    
DOI: 10.1111/brv.12023     Document Type: Article

References (217)

1. Akino, T. (2002). Chemical camouflage by myrmecophilous beetles Zyras comes (Coleoptera: Staphylinidae) and Diaritiger fossulatus (Coleoptera: Pselaphidae) to be integrated into the nest of Lasius fuliginosus (Hymenoptera: Formicidae). Chemoecology 12, 83-89.
2. Alfaro, A. C. (2007). Migration and trail affinity of snails, Littoraria scabra, on mangrove trees of Nananu-i-ra, Fiji islands. Marine and Freshwater Behaviour and Physiology 40, 247-255.
3. Amerongen, H. W. & Chia, F. S. (1982). Behavioural evidence for a chemoreceptive function of the cerebral organs in Paranemertes peregrine Coe (Hoplonemertea: Monostilifera). Journal of Experimental Marine Biology and Ecology 64, 11-16.
4. Bailey, S. E. R. (1989). Daily cycles of feeding and locomotion in Helix aspersa. Haliotis 19, 23-31.
5. Baur, B. (1992). Random mating by size in the simultaneously hermaphroditic land snail Arianta arbustorum: experiments and an explanation. Animal Behaviour 43, 511-518.
6. Beckett, T. W. (1968). Limpet movements. Tane 14, 43-65.
7. Blair, G. M. & Seapy, R. R. (1972). Selective predation and prey location in the sea slug Navanax inermis. Veliger 15, 119-124.
8. Bonabeau, E., Dorigo, M. & Theraulaz, G. (1999). Swarm Intelligence: From Natural to Artificial Systems. Oxford University Press, New York.
9. Bonabeau, E., Theraulaz, G., Deneubourg, J. L. & Camazine, S. (1997). Self-organisation in social insects. Trends in Ecology & Evolution 12, 188-193.
10. Bordereau, C., Lacey, M. J., Sémon, E., Braekman, J. C., Ghostin, J., Robert, A., Sherman, J. S. & Sillam-Dussès, D. (2010). Sex pheromones and trail-following pheromone in the basal termites Zootermopsis nevadensis (Hagen) and Z. angusticollis (Hagen) (Isoptera: Termopsidae: Termopsinae). Biological Journal of the Linnean Society 100, 519-530.
11. Bousfield, J. D., Tait, A. I., Thomas, J. D. & Towner-Jones, D. (1981). Behavioural studies on the nature of stimuli responsible for triggering mucus trail tracking by Biomphalaria glabrata. Malacologicol Review 14, 49-64.
12. Branch, G. M. (1975). Mechanisms reducing intraspecific competition in Patella Spp.: migration, differentiation and territorial behaviour. Journal of Animal Ecology 44, 575-600.
13. Branch, G. M. (1981). The biology of limpets: physical factors, energy flow, and ecological interactions. Oceanography and Marine Biology: An Annual Review 19, 235-380.
14. Bretz, D. D. & Dimock, R. V. (1983). Behaviorally important characteristics of the mucous trail of the marine gastropod Ilyanassa obsoleta (Say). Journal of Experimental Marine Biology and Ecology 71, 181-191.
15. Burchard, R. P. (1982). Trail following by gliding bacteria. Journal of Bacteriology 152, 495-501.
16. Calow, P. (1974). Some observations on locomotory strategies and their metabolic effects in two species of freshwater gastropods, Ancylus fluviatilis Müll. and Planorbis contortus Linn. Oecologia 16, 149-161.
17. Cammaerts, R., Detrain, C. & Cammaerts, M. C. (1990). Host trail following by the myrmecophilous beetle Edaphopaussus favieri (Fairmaire) (Carabidae: Paussinae). Insectes Sociaux 37, 200-211.
18. Capinera, J. L. (1980). A trail pheromone from silk produced by larvae of the range caterpillar Hemileuca oliviae (Lepidoptera: Saturniidae) and observations on aggregation behaviour. Journal of Chemical Ecology 6, 655-664.
19. Cate, J. M. (1968). Mating behaviour in Mitra idae Melvill, 1893. Veliger 10, 247-252.
20. Cavalier-Smith, T. (1998). A revised six-kingdom system of life. Biological Reviews 73, 203-266.
21. Chapman, M. G. (1998). Variability in trail-following and aggregation in Nodilittorina unifasciata Gray. Journal of Experimental Marine Biology and Ecology 224, 49-71.
22. Chapman, M. G. & Underwood, A. J. (1996). Influences of tidal conditions, temperature and desiccation on patterns of aggregation of the high-shore periwinkle, Littorina unifasciata, in New South Wales, Australia. Journal of Experimental Marine Biology and Ecology 196, 213-237.
23. Chase, R. (1986). Lessons from snail tentacles. Chemical Senses 11, 411-420.
24. Chase, R. & Croll, R. P. (1981). Tentacular function in snail olfactory orientation. Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology 143, 357-362.
25. Chase, R., Pryer, K., Baker, R. & Madison, D. (1978). Responses to conspecific chemical stimuli in the terrestrial snail Achatina fulica (Pulmonata: Sigmurethra). Behavioral Biology 22, 302-315.
26. Chelazzi, G., Della Santina, P. & Parpagnoli, D. (1990). The role of trail following in the homing of intertidal chitons: a comparison between three Acanthopleura spp. Marine Biology 105, 445-450.
27. Chelazzi, G., Della Santina, P. & Vannini, M. (1985). Long-lasting substrate marking in the collective homing of the gastropod Nerita textilis. The Biological Bulletin 168, 214-221.
28. Chelazzi, G., Focardi, S. & Deneubourg, J.-L. (1989). Analysis of movement patterns and orientation mechanisms in intertidal chitons and gastropods. In Behavioural Adaptation to Intertidal Life (eds G. Chelazzi and M. Vannini ), pp. 173-184. Plenum Press, London.
29. Chelazzi, G., Innocenti, R. & Della Santina, P. (1983). Zonal migration and trail-following of an intertidal gastropod analyzed by LED tracking in the field. Marine Behavior & Physiology 10, 121-136.
30. Chelazzi, G., Le Voci, G. & Parpagnoli, D. (1988). Relative importance of airborne odours and trails in group homing of Limaeus flavus (Linnaeus) (Gastropoda, Pulmonata). Journal of Molluscan Studies 54, 173-180.
31. Clifford, K. T., Gross, L., Johnson, K., Martin, K. J., Shaheen, N. & Harrington, M. A. (2003). Slime-trail tracking in the predatory snail, Euglandina rosea. Behavioural Neuroscience 117, 1086-1095.
32. Coffin, M. R. S., Barbeau, M. A., Hamilton, D. J. & Drolet, D. (2012). Effect of the mud snail Ilyanassa obsoleta on vital rates of the intertidal amphipod Corophium volutator. Journal of Experimental Marine Biology and Ecology 418-419, 12-23.
33. Coleman, R. A., Goss-Custard, J. D., Dit Durell, S. E. A. L. V. & Hawkins, S. J. (1999). Limpet Patella spp. consumption by oystercatchers Haematopus ostralegus: a preference for solitary prey items. Marine Ecology Progress Series 183, 253-261.
34. Connor, V. M. (1986). The use of mucous trails by intertidal limpets to enhance food resources. The Biological Bulletin 171, 548-564.
35. Connor, V. M. & Quinn, J. F. (1984). Stimulation of food species growth by limpet mucus. Science 255, 843-844.
36. Cook, S. B. (1969). Experiments on homing in the limpet Siphonaria normalis. Animal Behaviour 17, 679-682.
37. Cook, S. B. (1971). A study of homing behaviour in the limpet Siphonaria alternata. The Biological Bulletin 141, 449-457.
38. Cook, A. (1977). Mucus trail following by the slug Limax grossui Lupu. Animal Behaviour 25, 774-781.
39. Cook, A. (1979). Homing by the slug Limax pseudoflavus. Animal Behaviour 27, 545-552.
40. Cook, A. (1980). Field studies of homing in the pulmonate slug Limax pseudoflavus Evans. Journal of Molluscan Studies 46, 100-105.
41. Cook, A. (1985a). Functional aspects of trail following in the carnivorous snail Euglandina rosea Ferussac. Malacologia 26, 173-181.
42. Cook, A. (1985b). Tentacular function in trail following by the pulmonate slug Limax pseudoflavus Evans. Journal of Molluscan Studies 51, 240-247.
43. Cook, A. (1989). Factors affecting prey choice and feeding technique in the carnivorous snail Euglandina rosea Ferussac. Journal of Molluscan Studies 55, 469-477.
44. Cook, A. (1992). The function of trail following in the pulmonate slug, Limax pseudoflavus. Animal Behaviour 43, 813-821.
45. Cook, A. (1994). Trail following in slugs: the stimulus, its reception and the behavioural response. Ethology Ecology and Evolution 6, 55-64.
46. Cook, A. (2001). Behavioural ecology: on doing the right thing, in the right place at the right time. In The Biology of Terrestrial Molluscs (ed. G. M. Barker ), pp. 447-487. CABI Publishing, Wallingfort.
47. Cook, A., Bamford, O. S., Freeman, J. D. B. & Teideman, D. J. (1969). A study of the homing habit of the limpet. Animal Behaviour 17, 330-339.
48. Cook, S. B. & Cook, C. B. (1975). Directionality in the trail-following response of the pulmonate limpet Siphonaria alternata. Marine Behavior & Physiology 3, 147-155.
49. Cottrell, J. M., Henderson, I. F., Pickett, J. A. & Wright, D. J. (1993). Evidence for glycosaminoglycans as a major component of trail mucus from the terrestrial slug, Arion ater L. Comparative Biochemistry and Physiology Part B: Biochemical & Molecular Biology 104, 455-468.
50. Crothers, J. H. (1985). Dog-whelks: an introduction to the biology of Nucella lapillus (L.). Field Studies 6, 291-360.
51. Cummins, S. F., Erpenbeck, D., Zou, Z., Claudianos, C., Moroz, L. L., Nagle, G. T. & Degnan, B. M. (2009). Candidate chemoreceptor subfamilies differentially expressed in the chemosensory organs of the mollusc Aplysia. BMC Biology 7, 28.
52. Curtis, L. A. (1993). Parasite transmission in the intertidal zone: vertical migrations, infective stages, and snail trails. Journal of Experimental Marine Biology and Ecology 173, 197-209.
53. Davies, M. S. & Beckwith, P. (1999). Role of mucus trails and trail-following in the behaviour and nutrition of the periwinkle Littorina littorea (L.). Marine Ecology Progress Series 179, 247-257.
54. Davies, M. S. & Blackwell, J. (2007). Energy saving through trail following in a marine snail. Proceedings of the Royal Society of London, Series B: Biological Sciences 274, 1233-1236.
55. Davies, M. S. & Cliffe, E. J. (2000). Adsorption of heavy metals in seawater to limpet pedal mucus. Bulletin of Environmental Contamination and Toxicology 64, 228-234.
56. Davies, M. S., Edwards, M. & Williams, G. A. (2006). Movement patterns of the limpet Cellana grata (Gould) observed over a continuous period through a changing tidal regime. Marine Biology 149, 775-787.
57. Davies, M. S. & Hawkins, S. J. (1998). Mucus from marine molluscs. Advances in Marine Biology 34, 1-71.
58. Davies, M. S., Hawkins, S. J. & Jones, H. D. (1990a). Mucus production and physiological energetics in Patella vulgata L. Journal of Molluscan Studies 56, 499-503.
59. Davies, M. S., Jones, H. D. & Hawkins, S. J. (1990b). Seasonal variation in the composition of pedal mucus from Patella vulgata L. Journal of Experimental Marine Biology and Ecology 144, 101-112.
60. Davies, M. S., Hawkins, S. J. & Jones, H. D. (1992a). Pedal mucus and its influence on the microbial food supply of two intertidal gastropods, Patella vulgata L. and Littorina littorea (L.). Journal of Experimental Marine Biology and Ecology 161, 57-77.
61. Davies, M. S., Jones, H. D. & Hawkins, S. J. (1992b). Pedal mucus production in Littorina littorea (L.). In Proceedings of the Third International Symposium on Littorinid Biology (eds J. Grahame, P. J. Mill and D. G. Reid ), pp. 227-233. The Malacological Society of London, London.
62. Davies, M. S., Jones, H. D. & Hawkins, S. J. (1992c). Physical factors affecting the fate of pedal mucus produced by the common limpet Patella vulgata L. Journal of the Marine Biological Association of the United Kingdom 72, 633-643.
63. Davies, M. S. & Hutchinson, S. J. (1995). Crystalline calcium in littorinid mucus trails. Hydrobiologia 309, 117-121.
64. Davies, M. S. & Knowles, A. J. (2001). Effects of trematode parasitism on the behaviour and ecology of a common marine snail (Littorina littorea (L.)). Journal of Experimental Marine Biology and Ecology 260, 155-167.
65. Davies, M. S. & Williams, G. A. (1995). Pedal mucus of a tropical limpet, Cellana grata: energetics, production and fate. Journal of Experimental Marine Biology and Ecology 186, 77-87.
66. Davies, M. S. & Williams, G. A. (1997). Mucus production by Siphonaria species in Hong Kong. In Proceedings of the Forth International Workshop on the Marine Flora and Fauna of Hong Kong and Southern China (ed. B. S. Morton ), pp. 303-313. Hong Kong University Press, Hong Kong.
67. Davis, E. C. (2007). Investigation in the laboratory of mucous trail detection in the terrestrial pulmonate snail Mesodon thyroidus (Say, 1817) (Mollusca: Gastropoda: Polygyridae). American Malacological Bulletin 22, 157-164.
68. Davis-Berg, E. C. (2012). The predatory snail Euglandina rosea successfully follows mucous trails of both native and non-native prey snails. Invertebrate Biology 131, 1-10.
69. Dehnhardt, G., Mauck, B., Hanke, W. & Bleckmann, H. (2001). Hydrodynamic trail following in harbor seals (Phocavitulina). Science 293, 102-104.
70. Denny, M. W. (1980a). Locomotion: the cost of gastropod crawling. Science 208, 1288-1290.
71. Denny, M. W. (1980b). The role of gastropod pedal mucus in locomotion. Nature 285, 160-161.
72. Denny, M. W. & GOSLINE, J. M. (1980). The physical properties of the pedal mucus of the terrestrial slug, Ariolimax columbianus. Journal of Experimental Biology 88, 375-393.
73. Donovan, D. & Carefoot, T. (1997). Locomotion in the abalone Haliotis kamtschatkana: pedal morphology and cost of transport. Journal of Experimental Biology 200, 1145-1153.
74. Dunn, D. C. (1982). An examination of cues involved in mucous trail following by the mud snail Ilyanassa obsoleta. MA Thesis: Wake Forest University.
75. Edwards, D. C. (1968). Reproduction in Oliviella biplicata. Veliger 10, 297-304.
76. Edwards, M. & Davies, M. S. (2002). Functional and ecological aspects of the mucus trails of the intertidal prosobranch gastropod Littorina littorea (L.). Marine Ecology Progress Series 239, 129-137.
77. Edwards, S. F. & Welsh, B. L. (1982). Trophic dynamics of a mud snail (Ilyanassa obtsoleta (Say)) population on an intertidal mudflat. Estuarine, Coastal and Shelf Science 14, 663-686.
78. Erlandsson, J. (2002). Do reproductive strategy and breeding season influence the presence of mate recognition in the intertidal snail Littorina? Invertebrate Reproduction and Development 41, 53-60.
79. Erlandsson, J. & Kostylev, V. (1995). Trail following, speed and fractal dimension of movement in a marine prosobranch, Littorina littorea, during a mating and a non-mating season. Marine Biology 122, 87-94.
80. Erlandsson, J. & Johannesson, K. (1994). Sexual selection on female size in a marine snail, Littorina littorea (L.). Journal of Experimental Marine Biology and Ecology 181, 145-157.
81. Evison, S. E. F., Hart, A. G. & Jackson, D. E. (2008). Minor workers have a major role in the maintenance of leafcutter ant pheromone trails. Animal Behaviour 75, 963-969.
82. Farkas, S. R. & Shorey, H. H. (1972). Chemical trail-following by flying insects: a mechanism for orientation to a distant odor source. Science 178, 67-68.
83. Farkas, S. R., Shorey, H. H. & Gaston, L. K. (1974). Sex pheromones of Lepidoptera. Influence of pheromone concentration and visual cues on aerial odor-trail following by males of Pectinophora gossypiella. Annals of the Entomological Society of America 67, 633-638.
84. Fitzgerald, T. D. (1993). Trail and arena marking by caterpillars of Archips cerasivoranus (Lepidoptera: Tortricidae). Journal of Chemical Ecology 19, 1479-1489.
85. Funke, W. (1968). Heimfindevermogen und Ortstreue bei Patella L. (Gastropoda: Prosobranchia). Oecologia 2, 139-142.
86. Furry, K., Swain, T. & Chiszar, D. (1991). Strike-induced chemosensory searching and trail followed by prairie rattlesnakes (Crotalus viridis) preying upon deer mice (Peromyscus maniculatus): chemical discrimination among individual mice. Herpetologica 47, 69-78.
87. Galbraith, R. T. (1965). Homing behaviour in the limpets Acmaea digitalis and Lottia gigantea. American Midland Naturalist 74, 245-246.
88. Garrett, C. M., Boyer, D., Card, W. C., Roberts, D. T., Murphy, J. B. & Chiszar, D. (1996). Comparison of chemosensory behavior and prey trail-following behavior in the varanoid lizards Varanus gouldii and Heloderma suspectum. Zoo Biology 15, 255-265.
89. Garrity, S. D. (1984). Some adaptations of gastropods to physical stress on a tropical rocky shore. Ecology 65, 559-574.
90. Gehlbach, F. R., Watkins, J. F. & Kroll, J. C. (1971). Pheromone trail-following studies of typhlopid, leptotyphlopid, and colubrid snakes. Behaviour 40, 282-294.
91. Gerlach, J. (1999). The ecology of western Indian Ocean carnivorous land snails. Phelsuma 7, 14-24.
92. Gilly, W. F. & Swenson, R. P. (1978). Trail following by Littorina: washout of polarized information and the point of paradox test. The Biological Bulletin 155, 439.
93. Gläser, N., Wieskotten, S., Otter, C., Dehnhardt, G. & Hanke, W. (2011). Hydrodynamic trail following in a California sea lion (Zalophus californianus). Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology 197, 141-151.
94. Gonor, J. J. (1965). Predator-prey between two marine prosobranch gastropods. Veliger 7, 228-232.
95. Gould, S. J. & Vrba, E. S. (1982). Exaptation - a missing term in the science of form. Paleobiology 8, 4-15.
96. Guo, F., Huang, Z.-B., Huang, M.-Q., Zhao, J. & Ke, C.-H. (2009). Effects of small abalone, Haliotis diversicolor, pedal mucus on bacterial growth, attachment, biofilm formation and community structure. Aquaculture 293, 35-41.
97. Hadfield, M. G. & Switzer-Dunlap, M. (1984). Opisthobranchs. In The Mollusca, Volume 7: Reproduction (eds A. Tompa, N. H. Verdonk and J. A. Biggelaar ), pp. 209-350. Academic Press, New York.
98. Hall, J. R. (1972). Intraspecific trail-following in the marsh periwinkle Littorina irrorata (Say). Veliger 16, 72-75.
99. Hawkins, S. J. & Hartnoll, R. G. (1983). Grazing of intertidal algae by marine invertebrates. Oceanography and Marine Biology: An Annual Review 21, 195-282.
100. Heller, J. (1993). Hermaphroditism in molluscs. Biological Journal of the Linnean Society 48, 19-42.
101. Heller, S. B. & Halpern, M. (1981). Laboratory observations on conspecific and congeneric scent trailing in garter snakes (Thamnophis). Behavioural and Neural Biology 33, 372-377.
102. Herndl, G. J. & Peduzzi, P. (1989). Potential microbial utilisation rates of sublittoral gastropod mucus trails. Limnology and Oceanography 34, 780-784.
103. Hewatt, W. G. (1940). Observations on the homing limpet Acmaea scabra Gould. American Naturalist 24, 205-208.
104. Hirano, Y. & Inaba, A. (1980). Siphonaria (Pulmonate limpet) survey of Japan. I. Observations on the behaviour of Siphonaria japonica during breeding season. Publications of the Seto Marine Biological Laboratory 25, 323-334.
105. Holland, B. S., Chock, T., Lee, A. & Sugiura, S. (2012). Tracking behavior in the snail Euglandina rosea: first evidence of preference for endemic vs. biocontrol target pest species in Hawaii. American Malacological Bulletin 30, 153-157.
106. Holmes, S. P. (2002). The effect of pedal mucus on barnacle cyprid settlement: a source for indirect interactions in the rocky intertidal? Journal of the Marine Biological Association of the UK 82, 117-129.
107. Hughes, R. N. & Answer, P. (1982). Growth, spawning and trematode infection of Littorina littorea (L.) from an exposed shore in North Wales. Journal of Molluscan Studies 48, 321-330.
108. Hutchinson, N., Davies, M. S., Ng, J. S. S. & Williams, G. A. (2007). Trail following behaviour in relation to pedal mucus production in the intertidal gastropod Monodonta labio (Linnaeus). Journal of Experimental Marine Biology and Ecology 349, 313-322.
109. Iwai, N., Sugiura, S. & Chiba, S. (2010). Prey-tracking behavior in the invasive terrestrial planarian Platydemus manokwari (Platyhelminthes, Tricladida). Naturwissenschaften 97, 997-1002.
110. Iwasaki, K. (1992). Factors affecting individual variation in resting site fidelity in the patellid limpet Cellana toreuma (Reeve). Ecological Research 7, 305-331.
111. Iwasaki, K. (1998). Interindividual trail following by the intertidal patellid limpet Cellana toreuma. Journal of the Marine Biological Association of the United Kingdom 78, 1019-1022.
112. Jamon, M. (1994). An analysis of trail-following behaviour in the wood mouse. Animal Behaviour 47, 1127-1134.
113. Janson, K. & Sundberg, P. (1983). Multivariate morphometric analysis of two varieties of Littorina saxatilis from the Swedish west coast. Marine Biology 74, 49-53.
114. Jarau, S., Dambacher, J., Twele, R., Aguilar, I., Francke, W. & Ayasse, M. (2010). The trail pheromone of a stingless bee, Trigona corvina (Hymenoptera, Apidae, Meliponini), varies between populations. Chemical Senses 35, 593-601.
115. Jessee, W. F. (1968). Studies of homing behaviour in the limpet Acmaea scabra. Veliger 11(Suppl), 52-55.
116. Johannesson, K., Havenhand, J. N., Jonsson, P. R., Lindegarth, M., Sundin, A. & Hollander, J. (2008). Male discrimination of female mucus trails permits assortative mating in a marine snail species. Evolution 62, 3178-3184.
117. Johannesson, K., Rolán-Alvarez, E. & Ekendahl, A. (1995). Incipient reproductive isolation between two sympatric morphs of the intertidal snail Littorina saxatilis. Evolution 49, 1180-1190.
118. Johannesson, K., Saltin, S. H., Duranovic, I., Havenhand, J. N. & Jonsson, P. R. (2010). Indiscriminate males: mating behaviour of a marine snail compromised by a sexual conflict? PLoS ONE 5, e12005.
119. Johnson, M. P., Hughes, R. N., Burrows, M. T. & Hawkins, S. J. (1998). Beyond the predation halo: small scale gradients in barnacle populations affected by the relative refuge value of crevices. Journal of Experimental Marine Biology and Ecology 231, 163-170.
120. Kauffman, S. A. (1993). The Origins of Order: Self-organization and Selection in Evolution. Oxford University Press, New York.
121. Kemppainen, P., Panova, M., Hollander, J. & Johannesson, K. (2009). Complete lack of mitochondrial divergence between two species of NE Atlantic marine intertidal gastropods. Journal of Evolutionary Biology 22, 2000-2011.
122. Kideys, A. E. & Hartnoll, R. G. (1991). Energetics of mucus production in the common whelk Buccinum undatum L. Journal of Experimental Marine Biology and Ecology 150, 91-105.
123. Kofoed, L. H. (1975). The feeding biology of Hydrobia ventrosa (Montagu). II. Allocation of the components of the carbon-budget and the significance of the secretion of dissolved organic material. Journal of Experimental Marine Biology and Ecology 19, 243-256.
124. Krause, J. & Ruxton, G. D. (2002). Living in Groups. Oxford University Press, Oxford.
125. Kuanpradit, C., Stewart, M. J., York, P. S., Degnan, B. M., Sobhon, P., Hanna, P. J., Chavadej, J. & Cummins, S. F. (2012). Characterization of mucus-associated proteins from abalone (Haliotis) - candidates for chemical signaling. FEBS Journal 279, 437-450.
126. Kutschera, U., Pfeiffer, I. & Ebermann, E. (2007). The European land leech: biology and DNA-based taxonomy of a rare species that is threatened by climate warming. Naturwissenschaften 94, 967-974.
127. Lai, Y.-T., Chen, J.-H. & Lee, L.-L. (2011). The chemosensory ability of the predatory leech Whitmania laevis (Arhynchobdellida: Haemopidae) for prey searching. Chemoecology 21, 67-74.
128. Laimek, P., Clart, S., Stewart, M., Pfeffer, F., Wanichanon, C., Hanna, P. & Sobhon, P. (2008). The presence of GABA in gastropod mucus and its role in inducing larval settlement. Journal of Experimental Marine Biology and Ecology 354, 182-191.
129. Lassance, J.-M., Groot, A. T., Lienard, M. A., Antony, B., Borgwardt, C., Andersson, F., Hedenstrom, E., Heckel, D. G. & Lofstedt, C. (2010). Allelic variation in a fatty-acyl reductase gene causes divergence in moth sex pheromones. Nature 466, 486-491.
130. Lavenex, P. & Schenk, F. (1998). Olfactory traces and spatial learning in rats. Behavioural Processes 56, 1129-1136.
131. Lee, O. H. K. & Davies, M. S. (2000). Mucous production and morphometrics in the mangrove littorinids, Littoraria melanostoma and L. ardouiniana. In The Marine Flora and Fauna of Hong Kong and Southern China V (ed. B. Morton ), pp. 241-253. Hong Kong University Press, Hong Kong.
132. Leonard, J. L. & Lukowiak, K. (1985). Courtship, copulation, and sperm trading in the sea slug, Navanax inermis (Opisthobranchia: Cephalaspidea). Canadian Journal of Zoology 63, 2719-2729.
133. Li, D. & Graham, L. D. (2007). Epiphragmin, the major protein of epiphragm mucus from the vineyard snail, Cernuella virgata. Comparative Biochemistry and Physiology, Part B: Biochemistry & Molecular Biology 148, 192-200.
134. Lim, S. S. L. & Tan, T. L. (2008). The use of infrared spectroscopy as a test for species-specific pedal mucus in gastropod molluscs - a comparative study in Moreton Bay and Singapore. In Proceedings of the Thirteenth International Marine Biological Workshop, the Marine Fauna and Flora of Moreton Bay, Queensland: Memoirs of the Queensland Museum: Nature (Volume 54, eds P. J. F. Davie and J. A. Phillips ), pp. 349-354. Queensland Museum, Queensland.
135. Lincoln, B. J., Simpson, T. R. E. & Keddie, J. L. (2004). Water vapour sorption by the pedal mucus trail of a land snail. Colloids and Surfaces. B, Biointerfaces 33, 251-258.
136. Little, C., Williams, G. A., Morritt, D., Perrins, J. M. & Stirling, P. (1988). Foraging behaviour of Patella vulgata L. in an Irish sea-lough. Journal of Experimental Marine Biology and Ecology 120, 1-21.
137. Lorenzi, A. T. & Martins, M. F. (2008). Análise colorimétrica e espectroscópica do muco de caracóis terrestres Achatina sp alimentados com ração diferenciada. Revista Brasileira de Zootecnia 37, 572-579.
138. Lowe, E. F. & Turner, R. L. (1976). Aggregation and trail following in juvenile Bursatella leachii pleii. Veliger 19, 153-155.
139. Marin, A., Alvarez, L. A., Cimino, G. & Spinella, A. (1999). Chemical defence in cephalaspidean gastropods: origin, anatomical location and ecological roles. Journal of Molluscan Studies 65, 121-131.
140. McDonnell, R. J., Paine, T. D. & Gormally, M. J. (2007). Trail-following behaviour in the malacophagous larvae of the aquatic sciomyzid flies Sepedon spinipes spinipes and Dictya montana. Journal of Insect Behaviour 20, 367-376.
141. McFarlane, I. D. (1980). Trail-following and trail-searching behaviour in homing of the intertidal gastropod mollusc, Onchidium verruculatum. Marine Behavior & Physiology 7, 95-108.
142. McFarlane, I. D. (1981). In the intertidal homing gastropod Onchidium verruculatum (Cuv.) the outward and homeward trails have a different information content. Journal of Experimental Marine Biology and Ecology 51, 207-218.
143. Mohammed, S. Z. (1999). Aspects on clustering and movements of the intertidal gastropod, Planaxis sulcatus (Gastropoda/Planaxidae) in the Suez Canal. Indian Journal of Marine Sciences 28, 320-324.
144. Moomjian, L., Nystrom, S. & Rittschof, D. (2003). Behavioral responses of sexually active mud snails: kairomones and pheromones. Journal of Chemical Ecology 29, 497-501.
145. Muñoz, J. L. P., Camus, P. A., Labra, F. A., Finke, G. R. & Bozinovic, F. (2008). Thermal constraints on daily patterns of aggregation and density along an intertidal gradient in the periwinkle Echinolittorina peruviana. Journal of Thermal Biology 33, 149-156.
146. Nakashima, Y. (1995). Mucous trail following in 2 intertidal nudibranchs. Journal of Ethology 13, 125-128.
147. Navarro, J. M. & Torrijos, R. A. (1995). Fisiología energética de Concholepas concholepas (Bruguière, 1789) (Gastropoda: Muriciae) en la bahía de Yaldad, sur de Chile. Revista Chilena de Historia Natural 68, 61-77.
148. Ng, T. P. T., Davies, M. S., Stafford, R. & Williams, G. A. (2011). Mucus trail following as a mate-searching strategy in mangrove littorinid snails. Animal Behaviour 82, 459-465.
149. Niu, C. J., Nakao, S. & Goshima, S. (1998). Energetics of the limpet Lottia kogamogai (Gastropoda: Acmaeidae) in an intertidal rocky shore in southern Hokkaido, Japan. Journal of Experimental Marine Biology and Ecology 224, 167-181.
150. Norton, C. G. & Bronson, J. M. (2006). The relationship of body size and growth to egg production in the hermaphroditic freshwater snail, Helisoma trivolvis. Journal of Molluscan Studies 72, 143-147.
151. Nowak, M. A. (2006). Five rules for the evolution of cooperation. Science 314, 1560-1563.
152. Nuutinen, V. & Butt, K. R. (1997). The mating behaviour of the earthworm Lumbricus terrestris L. (Oligochaeta: Lumbricidae). Journal of Zoology 242, 783-798.
153. Ohgushi, R. (1954). Ethological studies on the intertidal limpets. 1. On the tidal rhythmic activities of two species of limpets. Japanese Journal of Ecology 4, 120.
154. Paine, R. T. (1963). Food recognition and predation on opisthobranchs by Navanax inermis (Gastropoda: Opisthobranchia). Veliger 6, 1-9.
155. Painter, S. D., Clough, B., Garden, R. W., Sweedler, J. V. & Nagle, G. T. (1998). Characterization of Aplysia attractin, the first water-borne peptide pheromone in invertebrates. The Biological Bulletin 194, 120-131.
156. Panova, M., Boström, J., Hofving, T., Areskoug, T., Eriksson, A., Mehlig, B., Mäkinen, T., André, C. & Johannesson, K. (2010). Extreme female promiscuity in a non-social invertebrate species. PLoS ONE 5, e9640.
157. Panova, M., Hollander, J. & Johannesson, K. (2006). Site-specific genetic divergence in parallel hybrid zones suggests non-allopatric evolution of reproductive barriers. Molecular Ecology 15, 4021-4031.
158. Pearce, T. A. & Gaertner, A. (1996). Optimal foraging and mucus trail following in the carnivorous land snail Haplotrema concavum. Malacological Review 29, 85-99.
159. Peck, L. S., Culley, M. B. & Helm, M. M. (1987). A laboratory energy budget for the ormer Haliotis tuberculata L. Journal of Experimental Marine Biology and Ecology 106, 103-123.
160. Peck, L. S., Prothero-Thomas, E. & Hough, N. (1993). Pedal mucus production by the antarctic limpet Nacella concinna (Strebel, 1908). Journal of Experimental Marine Biology and Ecology 174, 177-192.
161. Peduzzi, P. & Herndl, G. J. (1991). Mucus trails in the rocky intertidal: a highly active microenvironment. Marine Ecology Progress Series 75, 267-274.
162. Pescador-Rubio, A., Stanford-Camargo, S. G., Páez-Gerardo, L. E., Ramĺrez-Reyes, A. J., Ibarra-Jiménez, R. A. & Fitzgerald, T. D. (2011). Trail marking by caterpillars of the silverspot butterfly Dione juno huascuma. Journal of Insect Science 11, 55.
163. Peters, R. S. (1964). Function of the cephalic tentacles in Littorina planaxis Philippi (Gastropoda: Prosobranchiata). Veliger 7, 143-148.
164. Quick, H. E. (1960). British slugs (Pulmonata; Testacellidae, Arionidae, Limacidae). Bulletin of the British Museum (Natural History), Zoology Series 6, 105-226.
165. Quinet, Y. & Pasteels, J. M. (1995). Trail following and stowaway behaviour of the myrmecophilous staphylinid beetle, Homoeusa acuminata, during foraging trips of its host Lasius fuliginosus (Hymenoptera: Formicidae). Insectes Sociaux 42, 31-44.
166. Raftery, R. E. (1983). Littorina trail following: sexual preference, loss of polarized information, and trail alterations. Veliger 25, 378-382.
167. Reid, D. G., Dyal, P. & Williams, S. T. (2012). A global molecular phylogeny of 147 periwinkle species (Gastropoda, Littorininae). Zoologica Scripta 41, 125-136.
168. Reise, H. (2007). A review of mating behaviour in slugs of the genus Deroceras (Pulmonata: Agriolimacidae). American Malacological Bulletin 23, 137-156.
169. Richardson, A. M. M. (1975). Energy flux in a natural population of the land snail Cepaea nemoralis L. Oecologia 19, 141-164.
170. Robins, R. G. & Hamilton, P. V. (1996). Role of the cephalic tentacles in trail following by Littoraria irrorata (Say). Journal of Molluscan Studies 62, 537-539.
171. Ruf, C., Costa, J. T. & Fiedler, K. (2001). Trail-based communication in social caterpillars of Eriogaster lanestris. Journal of Insect Behaviour 14, 231-245.
172. Saltin, S. H. (2010). Complex male mate choice in marine snails Littorina. MSc Thesis: University of Gothenburg.
173. Saltin, S. H., Schade, H. & Johannesson, K. (in Press). Preference of males for large females causes a partial mating barrier between a large and a small ecotype of Littorina fabalis (W. Turton, 1825). Journal of Molluscan Studies.
174. Santelices, B. & Bobadilla, M. (1996). Gastropod pedal mucus retains seaweed propagules. Journal of Experimental Marine Biology and Ecology 197, 251-261.
175. Saur, M. (1990). Mate discrimination in Littorina littorea (L.) and L. saxatilis (Olivi) (Mollusca: Prosobranchia). Hydrobiologia 193, 261-270.
176. Schüpbach, H. U. & Baur, B. (2008). Experimental evidence for a new transmission route in a parasitic mite and its mucus-dependent orientation towards the host snail. Parasitology 135, 1679-1684.
177. Shaheen, N., Patel, K., Patel, P., Moore, M. & Harrington, M. A. (2005). A predatory snail distinguishes between conspecific and heterospecific snails and trails based on chemical cues in slime. Animal Behaviour 70, 1067-1077.
178. Skingsley, D. R., White, A. J. & Weston, A. (2000). Analysis of pulmonate mucus by infrared spectroscopy. Journal of Molluscan Studies 66, 363-371.
179. Skov, M. W., Volkelt-Igoe, M., Hawkins, S. J., Jesus, B., Thompson, R. C. & Doncaster, C. P. (2010). Past and present grazing boosts the photo-autotrophic biomass of biofilms. Marine Ecology Progress Series 401, 101-111.
180. Sleeper, H. L. & Fenical, W. (1977). Navenones A-C: trail-breaking alarm pheromones from the marine opisthobranch Navanax inermis. Journal of the American Chemical Society 99, 2367-2368.
181. Smith, A. M. (2002). The structure and function of adhesive gels from invertebrates. Integrative and Comparative Biology 42, 1164-1171.
182. Smith, A. M. (2006). Biological Adhesives. Springer-Verlag, Berlin, Heidelberg.
183. Smith, A. M. & Morin, M. C. (2002). Biochemical differences between trail mucus and adhesive mucus from marsh periwinkle snails. The Biological Bulletin 203, 338-346.
184. Solé, R. V., Bonabeau, E., Delgado, J., Fernández, P. & Marín, J. (2000). Pattern formation and optimization in army ant raids. Artificial Life 6, 219-226.
185. Stafford, R. (2002). The role of environmental stress and physical and biological interactions on the ecology of high shore Littorinids in a temperate and a tropical region. PhD Thesis: University of Sunderland.
186. Stafford, R. (2010). Crossing fitness valleys during the evolution of limpet homing behaviour. Central European Journal of Biology 5, 274-282.
187. Stafford, R. & Davies, M. S. (2005). Spatial patchiness of epilithic biofilm caused by refuge-inhabiting high shore gastropods. Hydrobiologia 545, 279-287.
188. Stafford, R., Davies, M. S. & Williams, G. A. (2007). Computer simulations of high shore littorinids predict small-scale spatial and temporal distribution patterns on real rocky shores. Marine Ecology Progress Series 342, 151-161.
189. Stafford, R., Davies, M. S. & Williams, G. A. (2008). Self-organization of intertidal snails facilitates evolution of aggregation behavior. Artificial Life 14, 409-423.
190. Stafford, R., Davies, M. S. & Williams, G. A. (2012a). Cheats in a cooperative behaviour? Behavioural differences and breakdown of cooperative behaviour in aggregating littorinids. Marine Ecology 33, 66-74.
191. Stafford, R., Davies, M. S. & Williams, G. A. (2012b). Misinterpreting the potential benefits of aggregation for reducing desiccation in the intertidal: a simple analogy. Marine Ecology 33, 512-515.
192. Stafford, R., Williams, G. A. & Davies, M. S. (2011). Robustness of self-organised systems to changes in behaviour: an example from real and simulated self-organised snail aggregations. PLoS ONE 6, e22743.
193. Stephenson, T. A. (1936). The marine ecology of the South African coast, with special reference to the habits of limpets. Proceedings of the Linnean Society of London 148, 74-79.
194. Stevens, A. (1995). Trail following and aggregation of myxobacteria. Journal of Biological Systems 3, 1059-1068.
195. Stirling, D. & Hamilton, P. V. (1986). Observations on the mechanism of detecting mucous trail polarity in the snail Littorina irrorata. Veliger 29, 31-37.
196. Switzer-Dunlap, M. & Hadfield, G. M. (1979). Reproductive patterns of Hawaiian aplysiid gastropods. In Reproductive Ecology of Marine Invertebrates (ed. S. E. Srancyk ), pp. 197-210. University of South Carolina Press, Columbia.
197. Takeichi, M., Hirai, Y. & Yusa, Y. (2007). A water-borne sex pheromone and trail following in the apple snail, Pomacea canaliculata. Journal of Molluscan Studies 73, 275-278.
198. Tankersley, R. A. (1989). The effect of trail-following on the locomotion of the marsh periwinkle Littorina irrorata (Mesogastropoda: Littorinidae). Marine Behaviour and Physiology 15, 89-100.
199. Thomas, R. F. (1973). Homing behaviour and movement rhythms in the pulmonate limpet, Siphonaria pectinata Linnaeus. Proceedings of the Malacological Society of London 40, 303-311.
200. Todd, C. D. (1977). The ecology of intertidal nudibranchs, with special reference to distribution, reproduction and population dynamics. PhD Thesis: University of Leeds.
201. Todd, C. D. (1979). The population ecology of Onchidoris bilamellata (Gastropoda: Nudibranchia). Journal of Experimental Marine Biology and Ecology 41, 213-255.
202. Tong, L. K. Y. (1988). The reproductive biology of Thais clavigera and Morula musiva (Gastropoda: Muricidae) in Hong Kong. Asian Marine Biology 5, 65-75.
203. Townsend, C. R. (1974). Mucus trail following by the snail Biomphalaria gIabrata (Say). Animal Behaviour 22, 170-177.
204. Trott, T. J. (1978). Trail following by the mud snail Ilyanassa obsoleta. MSc Thesis: Wake Forest University.
205. Trott, T. J. & Dimock, R. V. (1978). Intraspecific trail following by the mud snail Ilyanassa obsoleta. Marine Behavior & Physiology 5, 91-101.
206. Underwood, A. J. (1979). The ecology of intertidal gastropods. Advances in Marine Biology 16, 111-210.
207. Ushadevi, S. V. & Krishnamoorthy, R. V. (1980). Do slugs have silver track pheromone? Indian Journal of Experimental Biology 18, 1502-1504.
208. Wareing, D. R. (1986). Directional trail following in Deroceras reticulatum (Muller). Journal of Molluscan Studies 52, 256-258.
209. Webb, J. K. & Shine, R. (1992). To find an ant: trail- following in Australian blindsnakes (Typhlopidae). Animal Behaviour 43, 941-948.
210. Weissburg, M. J., Doall, M. H. & Yen, J. (1998). Following the invisible trail: kinematic analysis of mate-tracking in the copepod Temora longicornis. Philosophical Transactions of the Royal Society Series B: Biological Sciences 353, 701-712.
211. Wells, M. J. & Buckley, S. K. L. (1972). Snails and trails. Animal Behaviour 20, 345-355.
212. Williams, G. A. & Morritt, D. (1991). Patterns of foraging in Patella vulgata (L.). In The Ecology of Lough Hyne (eds A. Myers, C. Little, M. Costello and J. Partridge ), pp. 60-91. Royal Irish Academy.
213. Wilmer, P. (1990). Invertebrate Relationships: Patterns in Animal Evolution. Cambridge University Press, Cambridge.
214. Wilson, D. M. & Hoy, R. R. (1968). Optomotor reaction, locomotory bias, and reactive inhibition in the milkweed bug Ooncopeltus and the beetle Zophobas. Journal of Comparative Physiology, A: Neuroethology, Sensory, Neural, and Behavioral Physiology 58, 136-152.
215. Wright, J. R. (1977). The construction of energy budgets for three intertidal rocky shore gastropods Patella vulgata, Littorina littorea and Nucella lapillus. PhD Thesis: University of Liverpool.
216. Yen, J., Sehn, J. K., Catton, K., Kramer, A. & Sarnelle, O. (2011). Pheromone trail following in three dimensions by the freshwater copepod Hesperodiaptomus shoshone. Journal of Plankton Research 33, 907-916.
217. Zahradnik, T. D., Lemay, M. A. & Boulding, E. G. (2008). Choosy males in a littorinid gastropod: male Littorina subrotundata prefer large and virgin females. Journal of Molluscan Studies 74, 245-251.