Index [assets.cambridge.org]€¦ · Aeolidia papillosa (nudibranch sea-slug) predator marked by prey pheromone, 148 Aeshena umbrosa (dragonfly nymph) predator, 157 African mole rats,

Index

Aardwolf, see Proteles cristatusAbsolute configuration, 306–307Acanthomyops (ant), 5Accessory olfactory bulb (AOB),

165, 183, 184, 201(5R,6S)-6-Acetoxy-5-hexadecanolide, 75Acomys cahirinus (spiny mouse)

kin recognition cues, 110Acroneuria carolinensis (stonefly), 236Across-fibre patterning, 171Actias luna (silk moth)

antenna as sieve or paddle, 167Active space, 209

diffusion in air and water, 210–212multi-component pheromones,

212, 214trail, 215

Acyrthosiphon pisum (pea aphid)suicide hypothesis, 152

Adrenal glandsfemale mice, Lee–Boot effect, 195

Aeolidia papillosa (nudibranch sea-slug)predator marked by prey

pheromone, 148Aeshena umbrosa (dragonfly nymph)

predator, 157African mole rats, see Heterocephalus

glaber (naked mole rat)Aggregation pheromones, 74

aposematic insects, 75commercial exploitation

managing bark beetles, 266stopping marine fouling, 266

dilution of risk, 75ecophysiological benefits, 79

assembly pheromone and ticks, 80

overwintering garter snakes, 80intraspecific eavesdropping, 236

males only signal until femalescome, 82

Schistocerca gregaria(desert locust), 76

in space

oviposition pheromones, 75, 76, 77

Panulirus interruptus (lobster), 75settlement of marine

invertebrates, 26, 77–79in time (synchronisation)

coordinating external fertilisa-tion, 9, 91, 41–43

larval release, 77Agrotis ipsilon (black cutworm moth)

male response activated by juvenilehormone, 189

Alarm pheromonescommercial exploitation

for aphid control, 265hatchery trout taught to

recognise predator, 254costs to responding

aphid drop, 152costs to sender

eavesdropping by predators, 229, 234

evolution from pre-existing chemical cues, 9

evolution in related individuals,147–157

family groups, 147, 148, 149kin selection, 148–157

evolution in unrelated individuals,157–162

Diadema antillarum(sea urchin), 158

fish, 158–162predator labelling with host

pheromoneAeolidia papillosa (sea-slug), 148predatory fish, 161

propaganda by plants to repel aphids, 244, 245prey to escape ants, 244robber bees, 243slave-making ants, 241, 244

Alcelaphus buselaphus cokei (Coke’s hartebeest)

self marking and presentation, 92Aleochara curtula (rove beetle), 61Alkylmethoxypyrazine, 75Allee effects, 75

feeding, 79Dendraster excentricus

(sand dollar), 80mating, 79

external fertilisation, 79internal fertilisation, 79

overcoming tree defencesbark beetles, 81, 81

for wildlife conservation, 254Allelochemicals, 2Allomarking, 111Allomones, 2

deceit, propaganda, 230Alloparental care, 123

egg dumping, 230Alpheus heterochaelis (snapping shrimp)

modulation of visual threat signalsby pheromones, 18

Alternative mating strategiessatellite males

eavesdropping parasitoid pressure, 233

Nauphoeta cinerea (African cockroach), 60

she-malesAleochara curtula (rove beetle), 61Thamnophis sirtalis parietalis

(garter snake), 60Alzheimer’s disease, diagnosis, 297Amblyomma hebraeum (tick)

pest management, 262Amniotic fluid, 199Amphibians

Bufo borealis (western toad)tadpoles, alarm pheromone,

escape from predator, 157Litoria splendida (magnificent

tree frog)peptide sex pheromone, 21

tadpoles, kin, and alarm

371

Italic numbers refer to figures, boxes and tables

Organisms are nearly all indexed under taxonomic names in main entries; where commonnames are used a cross-reference is provided to the taxonomic name (except for groups).Taxonomic and/or common names are used in subentries.

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372 INDEX

Amphibians (cont.)pheromones, 157see also Salamanders and newts

Amphid sensilla, 203Amphiprion (anemone fish), 238, 239Amygdala

interaction between main olfactoryand vomeronasal organ (VNO)outputs, hamster, 185

Anal glandCastor canadensis (beaver)

variation between families, 96Meles meles (European badger), 111Mustela erminea (stoat)

repel prey, 265Anas platyrhynchos (mallard duck), 21Andrena nigroaenea (solitary bee)

duped by orchid, 240, 2423�-Androstenol, 253, 288, 289

steroid sex pheromones in pig, 20in truffles, use of sows to find, 253

5�-Androstenone, 191, 288, 289Boar Mate™, 252, 253, 254

Anguilla rostrata (American eel)zigzag behaviour, 227

Animal welfareunderstanding olfactory world, 254

Anomala osakana (Osaka beetle), 17Anonymous signals

social insects, 102Anosmias

induced changes on exposure, 191specific, 291

Antagonists, 66, 68, 187, 227, 260Antelopes

Kobus kob (Uganda kob)lekking, 58

Kobus leche (Kafue lechwe)lekking, 58

Litocranius walleri (gerenuk), 89Oreotragus oreotragus

(klipspringer antelope)scent marks eavedropped by

ticks, 233Ourebia ourebi (oribi), 87

border maintenance by multimale groups, 96

territories, 96Raphicerus melanotis (grysbok), 13

Antennal lobe, 165macroglomerular complex

(MGC), 165main, 165

Anthonomus grandis (boll weevil), 20pest management, 263

Anthopleura elegantissima (sea anemone)alarm pheromone (anthopleurine),

148, 149Anthopleurine, 148

soluble polar pheromone, 15Anti-aggregation pheromones, 235Antilocapra americana (pronghorn

antelope)alert signals, 148

Antorbital glandsterritorial and self marking

Alcelaphus buselaphus cokei(Coke’s hartebeest), 92

territorial marking by antelope, 89, 233

Aphaenogaster, see NovomessorAphrodisiac pheromones, see Male

pheromonesAphrodisin, 182Apis cerana japonica (Japanese

honeybee), 154–155, 156–157Apis mellifera, see HoneybeesAplysia (sea-slug)

peptide sex pheromone, 15Apocephalus paraponerae (phorid fly)

eavesdrops host pheromone, 233Apolipoprotein D (apoD), 290Aposematism

insects, aggregation, 75toad tadpoles, 157

Applications of pheromonesanimal husbandry

primer pheromones, 252signalling pheromones, 253

aquaculturehatchery trout taught to

recognise predator odour, 254induce sperm production in

carp, 254beneficial insects

honeybees, 255captive breeding rare species, 254

kin recognition, mate choice, 254commercialisation, 267–268

economics, 268public policy, 268

pest managementalarm pheromones, aphids, 265deterrent odours, 264–265lure and kill or mass trap,

260, 261–264

marine fouling organisms, 266mating disruption, 257–260,

261, 267monitoring, 256, 256, 257pest resistance to pheromones, 267primer pheromones, 265–266push-pull or stimulo-deterrent

diversionary strategies, 265, 266

self-protecting plants, 266pets

cats, spray marking, 254minipigs, 254

Tuber melanosporum (truffles), finding, 253

Aquatic pheromonesalarm pheromones

Diadema antillarum (sea urchin), 158

fish, 158toad tadpoles, 157

coordination of external fertilisationCarassius auratus (goldfish), 9, 19Nereis succinea (polychaete worm), 42

Dendraster excentricus (sand dollar)larval settlement, 80

Diadema antillarum (sea urchin)alarm responses, 158

eicosanoids (PUFAs)barnacle hatching pheromone, 77

Homarus americanus (lobster)precopulatory mate guarding,

61, 62peptides

Aplysia (sea-slug), 15Nereis succinea (polychaete

worm), 42sodefrin (newt), 15, 16

polar pheromoneanthopleurine, sea anemone,

15, 148, 149prostaglandins

Carassius auratus (goldfish), 19,27–28

Temora longicornis (copepod), 213see also Pheromones, aquatic habitats;

Signals, design, aquatic habitatsArachnids

mitesVarroa, 116, 262

phalangids (harvestmen)use ant alarm pheromone,

defence, 244






INDEX 373

spidersbolas spiders, 240, 243Habronestes bradleyi, attracted to

fighting ants, 229Linyphia litogiosa, 59

ticksguanine, assembly pheromone, 80Ixodes neitzi, 233

Argyortaenia velutinana (moth)male pheromone response wider

than range females produce, 69Argyropelecus hemigymnus

(deep-sea hatchet fish), 207Arms race

bark beetles and their predators, 234

Arthropodin, see Settlement-inducingprotein complex (SIPC)

Artificial insemination (AI), 253Assembly pheromone

ticks, 80Assortative mating, 65Asymmetric tracking, 68Atemeles pubicollis (beetle social para-

site of Myrmica ants), 246Atta (leaf-cutter ants), 114, 263

Atta texanasensitivity of workers to trail

pheromone, 136

‘Badges’ of statusNauphoeta cinerea (African

cockroach), 44Bark beetles

push-pull or stimulo-deterrent diversionary strategies, 266

see also Aggregation pheromones;Allee effects; Arms race; Dendroctonus; Exploitation ofpheromones by organisms; Ips;Mice; Pheromones

Barnaclesdisrupting settlement, 266egg hatching pheromone, 77larval settlement, 26, 78

Beetle aggregation pheromones, 81Behavioural manipulation by semen

componentsDrosophila melanogaster, 63humans? 64Thamnophis sirtalis parietalis

(garter snake), 63Benzaldehyde, 5, 243

Bioassays, see Pheromone identifica-tion, bioassays

BirdsAnas platyrhynchos (mallard duck)

possible sex pheromone, 21eavesdropping vole scent marks by

ultraviolet cuesButeo lagopus (rough-legged

buzzard), 232Falco tinnunculus (kestrel), 232

glandsanal, 21sebaceous, 21uropygidial, 21

nest recognition, 21pheromones, 21procellariiform seabirds

olfaction, 21zigzag flight upwind, 225

cis-(Z)-�-Bisbolene epoxide, 233Boar Mate™, 252, 253, 254Boiga irregularis (brown tree snake)

pest management, 262Bombykol

identification of, 25Bombus, see BumblebeesBombyx mori (silk moth), 23

antennae, design, 166Bos taurus (bulls and cows), 253Boundary layer

effects on olfactory sensors, 174turbulence, 219

Brain, see Olfactionendo-Brevicomin, 235exo-Brevicomin, 5Bruce effect

memory and pregnancy block inmice, 201

pregnancy block by malepheromones, 195

Bufo borealis (western toad)tadpoles, alarm pheromone, 157

Bumblebees (Bombus)marking visited flowers, 85

Butenandt, Adolf, 23Buteo lagopus (rough-legged

buzzard), 2322-sec-Butyl-dihydrothiazole

(thiazole or SBT), 193, 194

C20 polyunsaturated fatty acids (PUFAs) (eicosanoids), 77

Caenorhabditis elegans (nematode), 203

Callinectes sapidus, (blue crab)rheotaxis, 224

Callithrix jacchus (common marmoset)subordinate female ovulation

suppressed, 197Camponotus socius (ant)

group recruitment, 134Canis

Canis latrans (coyote)urine repells rodents, 265

Canis lupus (grey wolf)border marks in territory, 97suppression of subordinate

reproduction, 124Canis lupus familiaris, see DogsCanis sinensis (Ethiopian wolf)

border marks in territory, 97Carassius auratus (goldfish)

primer and releaser pheromones, 19

sex pheromones evolved from hormones, 9, 27–28

Castesdifferences

pheromone secretions, ants, 114responses to pheromones, social

aphids, 153social parasites copy odour badges

of termite host castes, 248see also Honeybees

Castor canadensis (beaver)anal glands, 96avoid marks at low population

densities, 91castoreum, 96kin recognition, 111pest management, 265suppression of subordinate

reproduction, 124Castoreum, 96Central-place foragers

naked mole rat 133social insects, 133

Ceratitis capitata (Mediterranean fruit fly)

leks, 58, 59eavesdropped by predator, 232

Ceratovacuna lanigera (social aphid)soldiers respond, non-soldiers

flee, 153soldiers attack predator, 153

Cerocebus albigena (mangabey,primate), 271






374 INDEX

c-fos, 182Chemical camouflage, 247–248Chemical mimicry, 248–249Chemical structure, 302Chemoreception, 164Chirality, 3055�-Cholestane-3,24-dione

trail pheromone, Malacosoma, 130Chrysopa slossonae (lacewing larvae)

escapes detection by chemical camouflage, 247

cis (in chemical name), 305Citral, 243Clean-up enzymes, 177Clethrionomys glareolus (bank vole)

females prefer odours of dominantmale, 46

Cloacal gland, 16Coccinella septempunctata (ladybird

beetle)aposomatic pheromones, 75

Colony defencecoordinated response to

danger, 155Colony-level selection

honeybees, 107Colophina monstrifica (social

aphid), 153Combinatorial coding

glomeruli, 170Communal nesting

enabled by synchronous breeding, 196

mouse preference for kin, 54, 55Communication, 2, 3

privacy in, 16sex communication channel, 66see also Composite signals;

Pheromones; SignalsComposite (combined) signals

chirp and odours, naked mole ratforagers, 136

parallel sensory channels, 17pheromone and stridulation

recruitmentAphaenogaster (Novomessor)

(ant), 140Conflict between the sexes

Linyphia litogiosa (spider), 59Nicrophorus (burying beetle), 59semen components

Drosophila melanogaster, 63humans, 64

Thamnophis sirtalis parietalis(garter snake), 63–64

Conscious and unconscious responsesto odours, 185

human responses, 185electroencephalogram (EEG), 185functional magnetic resonance

imaging (fMRI), 185Contact chemoreception

ants detecting colony identity, 4Contests, 44–46Coolidge effect, 57, 202

Gryllus bimaculatus (cricket), 57male rodents, 57Vipera berus (adder), 57

Cooperative breeders, 123–127plural breeders, 192

subordinate female ovulationsuppressed, 124, 126, 127,197–198

singular breeders, 124Cooperative signal, 122Coptotermes (termite), 130Coremata, 10, 11

honest signal, 3sexual selection, 47, 49

Cornicle secretionsaphids, 150

Corpora allata, 190termites, 119

Corynebacteriumbacteria in human armpits, 291

Costs (of signalling)eavesdropping by parasites

Ixodes neitzi (ticks), 233parasitoid insects, 232

eavesdropping by predators, 231ant-decapitating flies, 234lekking Ceratitis capitata (Mediter-

ranean fruit fly), 232spiders attracted to fighting

ants, 229low metabolic cost of pheromones

Anthonomus grandis (boll weevil), 20

Litoria splendida (magnificent treefrog), 21

long-distance communication,207

riskinvestigating marks, beavers, 89

timemarking, antelope, 87

Countermarking, see Over-markingCourtship

Cerocebus albigena (mangabey,primate), 271

Creatonotus gangis (arctiid moth), 10Drosophila melanogaster (fruit fly),

18, 70Grapholitha molesta (oriental fruit

moth), 7Homarus americanus (lobster), 61, 62humans, 277Macaca mulatta (rhesus monkey), 281newts, 16salamanders , pheromone delivery

patterns, 4species recognition, 64, 70, 72

Crataegus mollis (hawthorn), 85Creatonotus gangis (arctiid moth)

coremata, 3, 10Cremogaster dohrni artifex (attendant

ant of butterfly), 238Crocuta crocuta (hyena)

expose scent glands on approach, 91Cross-fostering experiments

Acomys cahirinus (spiny mouse)diet cues, 110

Odocoileus hemionus hemionus (black-tailed deer), 199

honeybees, 106MHC preferences of mice, 55Spermophilus beldingi (Belding’s

ground squirrel), 110Crotalaria

plant source of pyrrolizidine alka-loids (PAs), 47

CrustaceansAlpheus heterochaelis (snapping

shrimp)modulation of visual threat by

pheromones, 18barnacles

larval settlement, 26Semibalanus balanoides (barnacle),

77settlement behaviour, 78

copepodsLepeophtheirus salmonis (salmon ec-

toparasite), control, 262crabs

Callinectes sapidus (blue crab), 224Rhitropanopeus harrisii (mud crab),

77Homarus americanus (lobster)






INDEX 375

contest, 44precopulatory mate guarding,

61, 62tropotaxis, 222

Panulirus interruptus (Californiaspiny lobster)

aggregation for defence, 75plankton

Temora longicornis (copepod), 213Culex (mosquito), 263

oviposition pheromone, 75Cuticular hydrocarbons

ant trail pheromones, 140colony recognion

honeybees, 106ants, 108deception, 247–249Polistes fuscatus (paper wasp), 107

deception, 61dipteran sex pheromones, 70dipteran speciation, 71Gryllus bimaculatus (cricket)

mate choice, 51, 57solid phase microextraction (SPME)

from living ants, 33Cydia pomonella (codling moth)

pest management, 258Cynictis penicillata (yellow mongoose)

scent marking, 90Cynops pyrrhogaster (red-bellied newt)

courtship, 16sodefrin, 15, 16

Cypris larva (barnacle), 78L-Cysteine-glutathione disulfide, 42

Danaidone, 4Danaus gilippus (queen butterfly)

transfer pheromone direct to female, 4

Darwin, Charlessexual selection on pheromones, 37

Dauer larvaCaenorhabditis elegans

(nematode), 203Dear-enemy phenomenon (territorial

defence), 99, 112Deer

Odocoileus hemionus columbianus(black-tailed deer)

alert signals, 28, 147–148composite (combined) alarm

signal, 17scent marking, 91

2,3-Dehydro-exo-brevicomin (brevicomin or DHB), 5, 193, 194

2,5-Demethylpyrazine, 194Dendraster excentricus (sand dollar), 80Dendroctonus (bark beetles), 81

Dendroctonus brevicomiseavesdropped by predator, 234

Dendroctonus frontalis, 82male inhibition by male

pheromones, 235Dendroctonus micans, 81, 81Dendroctonus montanus, 81Dendroctonus pseudotsugae, 82Dendroctonus valens, 81

Dermacentor variabilis (tick)pest management, 262

Desmognathus ochrophaeus (salamander)male injects pheromone into

female, 4Development

agehamster, 189honeybee worker responses, 190honeybee, queen mandibular

pheromone (QMP), 121, 122Agrotis ipsilon (black cutworm

moth), 189caste differences in social insects

pheromone production andperception, 190

learningmale hormone surge to vaginal

odours (hamster), 182maternal behaviour, 191, 200Rhagolitis pomonella (apple maggot

fly), 190–191social insects, 107, 248

Schistocerca gregaria (desert locust), 76

sensitive periodewe learning odour of her lamb,

199, 200kin recognition, 110newly adult ant, 248

sex differences in brainmammals, 190moths, 190

switchdauer pheromone, 203social insects, 119, 204, 204


Diastereoisomers, 307Diet

effect on odourAcomys cahirinus (spiny

mouse), 110humans, 274

quality and mate choiceMicrotus pennsylvanicus

(meadow vole), 46Utetheisa ornatrix (tiger moth)

hydroxydanaidal (HD), 472,5-Dimethylpyrazine

female mice, Lee-Boot effect, 195N,N-Dimethyluracil, 130Dinoponera quadriceps (ant)

cuticular hydrocarbons, non-destructive study by solidphase microextraction (SPME),33–34

Diomedea (albatross)navigation, food location, 21, 225

Dipsosaurus dorsalis (iguana)composite (combined) signal with

ultraviolet cue, 14Disassortative mating

MHC genotype, 55(Z,Z,E)-Dodecatrienol, 130, 140(Z)-7-Dodecen-1-yl acetate, 1Dogs (Canis lupus familiaris)

ability to track odour trail, anddetect its direction, 218

as detector of cow oestrus, 253distinguishing humans by

smell, 274forensic use of smell, 296olfactory sensory sensitivity, 167scent marking landmarks, 99

Dolichotis patagonum (mara)males directly mark females, 100

DominanceHomarus americanus (lobster), 44, 61mice, 39, 93, 193, 194Microcebus murinus (lesser mouse

lemur), 196Microtus pennsylvanicus (meadow

vole), 46Nauphoeta cinerea (African

cockroach), 40, 44, 45queen, social insects, 118in queenless ants, 34–34social mammals, 124

Drakaea glyptodon (orchid) dupeswasp, 241






376 INDEX

Drosophila, 36, 71, 188, 209Drosophila adiastola subgroup, 72Drosophila mauritiana, 72Drosophila melanogaster, 63, 72

conflict between the sexes, 63courtship, pheromones, and

speciation, 18, 70, 71genetic engineering, 35–36, 70

Drosophila sechellia, 72Drosophila simulans, 71

Dufour’s gland, 11, 130ant trail, 139Lasioglossum zephyrum (sweatbee), 108parasitoid wasps, 84, 84queen bee egg mark, 122slave-making ants, 241, 244

Eavesdropping, see Exploitation ofpheromones by organisms

EchinodermsDendraster excentricus (sand dollar)

larval settlement, feedingbenefit, 80


Eciton (army ants), foraging, 141–143Egg dumping, 230

fish, 231insects, 230

Egg hatching pheromoneSemibalanus balanoides (barnacle), 77

Eicosanoids (PUFAs) (prostaglandinsin vertebrates)

barnacle hatching pheromone, 77Electroantennogram (EAG), 29, 30Electroencephalogram (EEG)

response to odours, 185Electroolfactogram (EOG), 189Electrovomerogram (EVG), 194Elephants

African (Loxodonta africana), 47mate guarding, 62

Asian (Elephas maximus), 1, 5, 47musth, testosterone elevation, 48pheromone shared with moths, 2

musthfemale response, 47male aggressiveness, 47temporal gland secretion, 47, 48testosterone elevation, 47, 48urine, 47

young males smell different fromadults, 47

Elephas maximus, see Elephants, AsianEnantiomers, 17, 305Enemy specification, 155Enoclerus lecontei

eavesdropping, 234, 236Entgegen (E), opposite, 308Environmental odour cues and

recognition, 103, 110Epiphyas postvittana (light brown apple

moth)pest management, 257, 258, 259

EphestiaEphestia elutella (tobacco moth), 47Ephestia kuehniella (flour moth)

pest management, 263Equus (horse), flehmen to oestrous

urine, 179Esox lucius (pike), 254

labelled with prey alarmpheromone, 159–161

naïve prey learn odour, 159predator attraction, interference

hypothesis, 162, 162Estrus, see Hormones; OestrusEstrogen, see Hormones; Oestrogen4-Ethyl-2-methoxyphenol, 44Ethyl-trans-cinnamate, 7Euglossine bees

courtship needs plant perfumeoils, 12

Eupoecilia ambiguella (grape moth)pest management, 258

Eusociality, 116conflict over reproduction in

societies, 118continuum (reproductive skew),

117, 117social insects and social mammals,

parallels, 127see also Social insects; Social

mammalsEvolution, see Pheromones,

evolutionExperience, see Development;

LearningExperimental methods, see

Pheromone identificationExploitation of pheromones by

organismsarms race, 230code breaking, 229deception

bolas spiders, 240, 243

cost of excluding deceivers, 249pollination by sexual deception,

240, 241, 242predators, guests and parasites of

social insects, 244–249propaganda, 241–244, 245

deception by mimic chemicalsenabled by shared biochemistry

in all life, 230eavesdropping, 229

aggregation pheromones, 81, 234,235, 236

alarm pheromones, 229, 234egg dumping, 230–232enabled by characteristics of

olfaction, 230intraspecific, sex pheromones as

aggregation pheromones, 236prey responses to predator

pheromones, 154–155, 156–157,236, 243, 264–265

sex pheromones, 231–232territorial marking

pheromones, 233intraspecific

hormones evolving intopheromones, 8

mutualism, 237‘communication’ between leaf-

cutter ants and fungus, 238ants and lycaenid butterfly

caterpillars, 237aphids tended by ants, 237sea anemones and anemone fish,

238–240External fertilisation

coordination by pheromones, 41Carassius auratus (goldfish), 9, 19Nereis succinea (polychaete

worm), 42

Fabré, Jean-Henri, 23Faeces, territorial marking

antelopes (oribi), 96Meles meles (European badger), 98salamanders, 95

Falco tinnunculus (kestrel), 232Farnesene, 130, 193, 194

produced by wild potato, 244, 245Feliway™ (synthetic cheek gland

pheromone), 254Female choice, see Mate choice; Sexual

selection






INDEX 377

Female pheromoneseffects on females

biases sex ratio in pregnancy, 197delayed puberty, 194, 195oestrus synchronisation, 196, 282social insects, 115–116, 119, 121, 123subordinate female ovulation

suppressed, 197suppressed oestrus cycles, 194, 195

effects on malesMicrocebus murinus (lesser mouse

lemur), 196Femoral gland

Iguana iguana (lizard)testosterone influence, 46

Fishalarm pheromones, 158–162Amphiprion (anemone fish

symbiotic with sea anemones),238, 239

Anguilla rostrata (American eel)zigzag behaviour, 227

Argyropelecus hemigymnus (deep-seahatchet fish)

biggest nasal organ for body size,vertebrate, 207

Carassius auratus (goldfish)pheromones from hormones, 9,

19, 27–28, 43, 189Esox lucius (pike)

labelled with prey alarmpheromone, 159–161

naïve prey learn odour, 159odour to teach hatchery rainbow

trout, 254predator attraction, interference

hypothesis, 162, 162Gadus morhua (cod)

zigzag orientation in plume, 225Gasterosteus aculeatus (stickleback)

MHC and mate choice, 55Lepomis cyanellus (green sunfish)

alloparental carer, 230–231Notropis umbratili (redfin shiner)

interspecific egg dumper, 231Oncorhynchus mykiss (rainbow trout)

taught to recognise predatorodour, 254

Petromyzon marinus (marine lamprey)control, 262

Phoxinus phoxinus (Europeanminnow)

alarm pheromone, 158

Plotsus lineatus (catfish)scanning electron microscopy of

olfactory epithelium, 169Premnas (anemone fish symbiotic

with sea anemones), 238Primiphales promelas (fathead min-

now), 158Puntius (androgen effects on elec-

troolfactogram (EOG)), 189salmon

zigzag orientation in plume, 225Flame ionisation detector (FID), 30Flehmen

Bos taurus (bulls and cows), 253Elephas maximus (Asian elephant), 48Equus (horse), to oestrous urine, 179

Flicks, antennalbehaviour of smelling, 174Homarus americanus (lobster), 174

Floaters, 99Fluctuating asymmetry, 49

humans, 280Panorpa japonica (scorpion fly), 49

Formica subintegra (slave-making ant),241, 244

Formicoxenus (shampoo ant)social parasite of ants (Myrmica),

247Frequency dependant responses

parasitoid wasp superparasitism,188

Fright response, 158–159Functional groups, 302Functional magnetic resonance

imaging (fMRI)humans, 174, 175, 185

Gadus morhua (cod)zigzag orientation in plume, 225

Galago (bushbaby)nocturnal, use of pheromones, 13

Galaxolide, 185Gargaphia solani (aubergine lace bug)

intraspecific egg dumping, 230subsocial care, alarm pheromone,

147Garter snake, see Thamnophis sirtalis

parietalisGas liquid chromatography (GC), see

Pheromone identification, separating chemicals

Gasterosteus aculeatus (stickleback), 55Genetics

Caenorhabditis elegans (nematode)using mutants, 203

Drosophila, 35–36, 70humans, perception of odours,

specific anosmias, 291–292moths, pheromone signal, 69olfactory receptor proteins (OR), 169

Geometrical isomersEntgegen (E), Zusammen (Z), 308

Geraniolalarm pheromone

Gargaphia solani (aubergine lacebug), 147

Gestalt modelcolony odour, 108

Glands, 9, 10, 11adrenal

female mice, Lee–Boot effect, 195anal

birds, 21Castor canadensis (beaver), 96Meles meles (European badger),

111Mustela erminea (stoat), 265

antorbitalterritorial marking by antelope

(gerenuk) 89, 233cloacal, 16closable, due to eavesdropping

predators, 233Dufour’s

ant trail, 139Lasioglossum zephyrum (sweatbee),

108parasitoid wasps, 84, 84queen bee egg mark, 122slave-making ants, 241

femoralIguana iguana (lizard), 46

labialtermites, 138

mandibularTrigona (stingless bee), 136

musk, 148Nasonov

honeybees, 131poison

ant, 131postcloacal

salamander, 95postpharyngeal

ants, 108, 248preorbital, see antorbital






378 INDEX

Glands, (cont.)preputial

mice, 193pygidial

ant, 134sebaceous

birds, 21humans, 284signal life, 14

some vary more within aspecies, 113

sternalant, 130Malacosoma americanum (tent

caterpillar), 130Polistes (wasp), 107termite, 130, 137

subcaudalMeles meles (European badger), 111

sweatapocrine, 284eccrine, 284

temporalelephants, 46

uropygidialbirds, 21

Glomerulicombinatorial coding, 170–174functional units of processing, 172

L-Glutamic acid, 42L-Glutamine, 42Gossyplure, 303G-proteins, 166Grandlure, 263Grapholitha molesta (oriental fruit moth)

courtship display of hair pencils, 7pest management, 258response to pheromone blend, 67

‘Green beard’ phenomenon, 104, 105Gryllus bimaculatus (cricket)

Coolidge effect, 57mate choice for optimal

outbreeding, 51Guanine, 80

Habronestes bradleyi (spider), attractedto fighting ants, 229

Hair pencilsmoth, 7

Hamilton’s ‘selfish-herd’ theoryfish shoals and alarm

pheromone, 159Hamilton’s rule, kin selection, 116

Hawaiian Drosophila fruit flies, 72Drosophila adiastola subgroup, 72speciation, 72

Helicoverpa zea (corn ear wormmoth), 187

Helogale undulata (dwarf mongoose)individual recognition, 112

(Z)-7-Heneicosene, 61Heneicosane, 842-Heptanone, 194Herpestes auropunctatus (Indian

mongoose)individual recognition, 112

Heterocephalus glaber (naked molerat), 125

colony odour, parallels with socialinsects, 112

queen suppression of worker reproduction, 124

recruitment of foragers, 136Heterodera glycines (soybean cyst

nematode)pest management, 258, 260

E-10,Z-12-Hexadecandien-1-ol, 25Z-9-Hexadecenal, 130Hexanoic acid, 130Hinterland marking, 97–98Homarus americanus (lobster), 44

antennal flicks, 174precopulatory mate guarding, 61, 62tropotaxis, 222

Homo sapiens, see HumansHonest signal, 2, 44, 45

queen pheromonehoneybees, 121, 121

reflecting biological state ofmarker

territorial scent marks, 89Utetheisa ornatrix (tiger moth), 47,

49, 49, 50Honeybees (Apis mellifera)

Africanised bees, monitoringspread of, 256, 256

cell capping, odour cues, 116colony odour sources, 106colony-level signature, 106comb wax odour cues, 106manipulating behaviour for bee-

keepingdelaying swarming, 255directing pollination, 255luring swarms, 255

marking visited flowers, 85

multiple paternity in nestblending of template colony

odour, 105Nasonov pheromone

marking new nesting cavity, 131swarming, 131

pheromones, summary, 115queen mandibular pheromone

(QMP), 121as honest signal, 121as sex pheromone, 19rapid colony response to loss, 121transmission, 121

queen-rearing nepotism, 107queen retinue, 116sting pheromone attracts other

bees to sting, 154swarming, 121Varroa mite, 116, 262

Hormonescortisol and recognition of human

babies, 276humans

birth, 201cortisol and recognition of

human babies, 276juvenile hormone ( JH)

Agrotis ipsilon (black cutwormmoth), 189–190

social insects, 204, 204oestrogen, 198oxytocin in ewes, 199, 200, 201pheromone influence on hormones

female stimulates testosterone secretion in lesser mouselemur, 196

male hormone surge to vaginalodours in hamster, 182

puberty acceleration in rodents,ungulates and non-human primates, 193

responses induced via olfactorysystem, 185

sexual activity in Petaurus breviceps (sugar glider), 196

testosteroneandrogen effects on electroolfac-

togram, fish, 189femoral gland, Iguana iguana

(lizard), 46graded response of scent glands

in Microtus pennsylvanicus(meadow vole), 46






INDEX 379

hamster male brain needs to respond, 189

male sexual response in mammals, 183

musth and temporal gland secretion in elephants, 47, 48

see also Primer pheromones; Female pheromones; Malepheromones

Host-discrimination, 83Host-marking pheromones (HMP), 74,

83–85evolution of, 83host-discrimination, 83individual recognition, 85

Human leucocyte antigen (HLA), seeHumans, mate choice; Majorhistocompatibility complex;Mammals, mate choice

Humansadvertisement of oestrus, 281

‘copulins’, 281unresolved in humans, 282

age, changes withperception, 271secretions, 275sensitivity, 297

anosmiasinduced changes on exposure,

191specific, 291

antiperspirants and deodorants,288

axilla (armpit) hair and odour creating bacteria, 288

behavioural manipulation by semen hormones? 64

candidate compounds, 285axillae (armpits), 287–291glands, 284sites over body, 286, 287

cleanliness not necessarily avirtue, 276

cultural and social aspects ofodours, 273

osmologies, 273Napoleon, 273sexual attraction, 273

importance of odours in humanbehaviour and biology, 274

mate choice, 2775�-androstenone, 277body symmetry and, 280

kibbutz or Westermarck effect,280

human leucocyte antigen (HLA),52, 278

major histocompatibilitycomplex (MHC), 52–56,278–280

sexually selected traits, 277mothers and newborn babies, 275

correlation of mother cortisoland baby recognition, 276

importance of natural odours,275

speed of learning, 275odour preferences

learning, 271neonates different from older

children and adults, 271odours and memory

experimental tests, 276Proust, 276

olfactionmapping brain activity, 174, 175,

185olfactory receptors

genes and pseudogenes, 170perception of odours, 291

changes during menstrual cycle,292, 293

changes with puberty, 292specific anosmias, 291, 292variation between individuals,

291perfumes, 273, 279phenotype matching, 276primer pheromones

age of female puberty, 285menstrual synchrony, 282–285steroids and mood, 295

recognition, 274children, sibs, parents, 275security blankets, 276self, partners, 275T-shirt sniff tests, 274

semen components, 64truffles, 253, 288using human odours, 295

‘human pheromones’, 298forensics, 296medical diagnosis, 296–297mood changers, 298

vomeronasal organ (VNO),putative, 294

genetic evidence, 295location, 294neuroanatomical and histologi-

cal evidence, 295see also Primates

HutteritesMHC and partner choice, 280

Hydrobates pelagicus (storm petrel)nest recognition, 21

Hydrocarbons, see Cuticularhydrocarbons

Hydroides (polychaete worm)plankton settlement, 79

3-Hydroxy-2-butanone, 446-Hydroxy-6-methyl-3-heptanone,

193, 194Hydroxydanaidal (HD), 50

derived from plant pyrrololizidinealkaloids in diet, 47

Hypothalamus, 184Microtus ochrogaster (prairie vole), 198oxytocin

maternal behaviour, 199, 200Hypoxanthine 3-N-oxide, 158

Iguana iguana (lizard)forest habitat, volatile pheromone,

14–15femoral gland

testosterone influence, 46Imprinting

mate recognition, 198odours, recognising kin and colony

members, 199Individual recognition

generalisation across glandodours, 113

Helogale undulata (dwarf mongoose),112

Herpestes auropunctatus (Indianmongoose), 112

Homarus americanus (lobster), 46humans, 274–275mechanisms, within a species some

glands vary more than others,113

Meles meles (European badgers), 112mice, 112pair bonding

Niveoscincus microlepidotus(lizard), 113

Proteles cristatus (aardwolf)‘dear-enemy’ recognition, 113






380 INDEX

Information centreMalacosoma americanum (tent

caterpillar), 133Inosine, 42Integrated pest management

(IPM), 255pheromones in control of tomato

pests, 259Ips (bark beetles)

Ips paraconfusus, 235discovery of synergy, 35

Ips pinibioassays, 24

Ips typographuspest management, 264

(R)-(�)-ipsdienol, 234(S)-(�)-ipsdienol, 234Iridomyrmex (ants), 130Isomers, 304

constitutionalfunctional group, 304positional, 304

naming of, 305stereoisomers

chirality and enantiomers (optical isomers), 305–307

Ixodes neitzi (tick)responds to scent marks of host

antelope, 233

(R)- and (S)-Japonilure enantiomersuse by sympatric beetles, 17

Juvenile hormone (JH)Agrotis ipsilon (black cutworm moth)

male response activated by, 189social insects, 204

ants, 204termites, 119

Kairomones, 2eavesdropping, 230

Kangaroosmate guarding by male, 62

Keifera lycopersicella (tomato pin worm)pest management, 258, 259, 261

Kibbutz or Westermarck effecthuman mate choice, 280

Kin recognition 103–105mate choice

conservation projects, 254mechanisms, 103

low metabolic costs, 103Kin selection, Hamilton’s rule, 116

KobusKobus kob (Uganda kob), 58Kobus leche(Kafue lechwe), 58

Labelled lines, 187Labial gland

termites, 138Lasioderma serricorne (cigarette beetle)

aggregation pheromone, 83, 84Lasioglossum zephyrum (sweatbee)

guard bees discriminate relatedness, 108

Lasius (ants)trail pheromone, 130Lasius alienus

alarm pheromone, causes panicand escape, 155

Lasius fuliginosustrail following, 216

Lasius nigerforaging effort, food value, 139

Lateral olfactory tract, 183Learning

fish, conditioned alarm response topredator odours, 159, 191, 254

mammalsinduced peripheral changes in

response to odours after contact, 191

maternal behaviour, 191needed to distinguish oestrous

from dioestrous female odours,191

maternalewes bond quicker on subsequent

births, 200odour stimulation and

noradrenaline release inolfactory bulbs, 199

Rhagolitis pomonella (apple maggotfly), 190–191

see also DevelopmentLee–Boot effect, 194, 195Leks, 57

Ceratitis capitata (Mediterraneanfruit fly), 58

cost, eavesdropped by predators,231–232

Kobus kob (Uganda kob), 58Kobus leche (Kafue lechwe), 58Lutzomyia longipalpis (sandfly), 58Xylocopa fimbriata (carpenter bee),

57

Lemur catta (ring-tailed lemur)group territorial ‘stink fights’, 90

Leontopithecus rosalia (lion tamarin)suppression of subordinate

reproduction, 127Lepeophtheirus salmonis (salmon

ectoparasite)pest management, 262

Lepomis cyanellus (green sunfish)alloparental carer, 230–231

Leptothorax acervorum (ant)tandem running recruitment, 134

Leptotyphlops dulcis (blind snake), 247Lestrimelitta limao (robber bee), 243Linoleic acid, 84Linyphia litogiosa (spider)

conflict between the sexes, 59Liriomyza sativae (leafminer fly), 259Litocranius walleri (gerenuk), 89Litoria splendida (magnificent tree

frog)peptide sex pheromone, 21

Lobesia botrana (grape vine moth)pest management, 258

Lordosis, 20, 184, 253Loxodonta africana, see Elephants,

AfricanLuteinising hormone (LH), 183

Microtus ochrogaster (prairie vole),198

Luteinising hormone-releasing hormone (LHRH), 198

Lutzomyia longipalpis (sandfly), 263lekking, 58

Macaca mulatta (rhesus monkey), 281Macroglomerular complex (MGC), 165Macrotermes (termite)

nest building, 143–144Maculinea rebeli (lycaenid butterfly)

manipulates ant host, 249Main olfactory bulb (MOB), 165Main olfactory epithelium (MOE), 165,

178, 179, 182, 183Main olfactory system

detection of pheromones, 181Oryctolagus cuniculus (European

rabbit), 181Sus scrofa (pig), 184

sheep maternal behaviour, 201see also Olfaction

Major histocompatibility complex(MHC)






INDEX 381

bacterial flora and odours, 53Bruce effect, 195communal nesting, mouse

preference for kin, 54, 55familial imprinting

odour preferences for differencein mates, 55

human leucocyte antigen (HLA),52, 278

kin recognition in other vertebrates(fish, birds, reptiles), 56

mate choice, 52MHC diversity, Gasterosteus

aculeatus (stickleback), 55mice and humans, 54mutual distinguishing MHC of

odours inrats, mice and humans, 52

not universal, 56odours as marker for degree of

kinship, 52semi-natural experiments, 52

preferenceslearning, 55women taking oral

contraceptives, 54selection pressures maintaining

diversity of MHC alleles, 55, 56disassortative mating, 55higher implantation rates if dis-

similar, 55reduced chance of inbreeding if

dissimilar, 55source of the odours, 52–54

Major urinary proteins (MUPs),194, 195

longevity of signal, slow release,193

elephants, 195mice, 195

Malacosoma (tent caterpillars)trail pheromone, 130Malacosoma americanum, 133

laying trail, 135Male pheromones

diverse release structures, 7, 10, 70effects on females

leks, 57–59mice, Bruce effect (pregnancy

block), 195oestrus induction and puberty

acceleration (Whitten and Vandenbergh effects), 192

oestrus synchronisation, mice,194

effects on males, 196sexual activity of subordinates

suppressed, 196selection by female to avoid hybrid

matings, 70Mammals

group, kin, family, and individualrecognition, 109

individual recognition, 112Helogale undulata (dwarf

mongoose), 112Herpestes auropunctatus (Indian

mongoose), 112Meles meles (European badger), 112mice, 112

kin recognitionAcomys cahirinus (spiny mouse),

110mate choice

major histocompatibility complex (MHC), 52–56

mother–infant recognition, 109scent marking, 87scent sources, summary, 10secretory glands, 11see also Humans; Primates; Social

mammalsMan, see HumansMandibular gland

Trigona (stingless bee), 136Manduca sexta (tobacco horn moth)

zigzag orientation upwind, 225Marine invertebrates, alarm

responses, 158Marking enemies for further attack,

155Marsupials

kangaroosmate guarding by male, 62

Monodelphis (opossum)vomeronasal organ (VNO) needed

for oestrus induction, 181Petaurus breviceps (sugar glider)

scent marks affect sexual activity, 196

MastophoraMastophora bisaccata (bolas spider),

243Mastophora hutchinsoni (bolas

spider), 240Mate choice

avoiding lethal alleles, ‘t-complex’in mice, 56

diet qualityMicrotus pennsylvanicus (meadow

vole), 46fluctuating asymmetry

humans, 280Panorpa japonica (scorpion fly), 49

MHC diversity, see Major histocompatibility complex,mate choice

optimal outbreeding, 51Gryllus bimaculatus (cricket), 51Lasioglossum zephyrum (sweat-

bees), 51parasites, 51

mice, preference for uninfectedmates, 51

paternal investment and femalechoice, 47

Utetheisa ornatrix (tiger moth), 47pheromones

Ephestia elutella (tobacco moth), 47Mate guarding, see Sperm competition

and mate guardingMaternal behaviour and recognition

humans, 275sheep, 199

Mating disruption, as pest control,258

Medial amygdaloid nucleus, 183Medial preoptic area (MPOA), 182, 183Megaponera (ant), 130Meles meles (European badger)

anal glands, 111border and hinterland latrines,

98, 98individual recognition, 112subcaudal glands, 111

Memory and pregnancy block (Bruceeffect) in mice, 201

accessory olfactory bulb (AOB), 201Mesocricetus auratus (hamster) , 183

aphrodisin, 182over-marking, 100self-matching, 105

Messor barbarus (harvester ant)matching trail patterns to food

distribution, 1394-Methyl-3-heptanone, 2346-Methyl-5-hepten-2-one, 243(E)-3-Methyl-2-hexenoic acid ((E)-3M2H

or TMHA), 289






382 INDEX

Methyl-4-methylpyrrole-2-carboxylate,130, 136

2-Methyliazolidine, 44MHC, see Major histocompatibility

complexMice (Mus musculus domesticus)

Bruce effectmemory and pregnancy block,

201individual recognition, 112Lee–Boot effect, 195male pheromones

oestrus induction and puberty acceleration in females (Whitten and Vandenbergh effects), 192

marking rates, dominants and subordinates, 94, 94

mate choicepreference for uninfected

mates, 51mate choice to avoid lethal alleles,

‘t-complex’, 56pheromone marks

importance for animal wel-fare, 254

scent marking by dominantmale, 93

signal sites, urine posts, 94similar pheromone to bark beetle, 5‘t-complex’, 56Vandenbergh effect, 192Whitten effect, 192

Microcebus murinus (lesser mouselemur)

female urine stimulates spermatogenesis and testosterone secretion, 196

pheromones and sex ratios, 197sexual activity of subordinates

suppressed, 196Microdon mutabilis (syrphid fly)

matches its ant host, 248Microtus

Microtus ochrogaster (prairie vole),198

unfamiliar male odours, stimulation of oestrus, 198

Microtus pennsylvanicus (meadowvole)

suppression of subordinate reproduction, 124, 126,197–198

graded response of scent glandsto testosterone implants, 46

Microtus pinetorum (pine vole)suppression of subordinate

reproduction, 127Miriamide, 84Mitral/tufted cells, 171, 172

Bruce effect (memory of mate), 201memory for lamb, 200

MolluscsAplysia (sea-slug), peptide sex

pheromone, 15snails, direction of a trail, 217

Monodelphis (opossum)vomeronasal organ (VNO) needed

for induction of oestrus, 181Monogamous partner recognition,

202Monozygotic twins

human, dogs have difficulty distinguishing, 274

sheep, lamb recognition experiment, 109

Mother–infant recognitionhumans, 275sheep, 109

Mouse, see MiceMulti-component pheromones, 187

alarm pheromones in socialinsects, 154

ant, alarm pheromone signal,212, 214

asymmetric tracking, 68major and minor components, 66male response wider than range

females produce, 69response is to whole blend, moths,

67specificity, 17, 66synergy

insects, 35mammals, 195

Musca domestica (house fly)pest management, 263

Mus musculus domesticus, see MiceMushroom body, 165Musk, 185Musk gland, 148Mustela

Mustela erminea (stoat), anal glands,265

Mustela putorius furo (ferret), 190Musth, see Elephants

Myrmecaphodius excavaticollis (beetle)parasite of Solenopsis (fire ant), 247

Myrmecophilous invertebrates, 245Myrmica (ants), 247

Myrmica sabuletimatching foraging effort to food

value, 139Myzus persicae (aphid)

repelled by alarm pheromone fromplant, 244

Naming pheromonesfunctional groups

prefixes and suffixes, 303Napoleon, 273Nasonov pheromone

honeybees, 131, 255Nasutitermes (termite), 120, 155

marking enemies for further attack, 155

recruitment pheromones, 140Nauphoeta cinerea (African cockroach)

‘badges’ of status, 44honest signal, 44, 45male contest and female choice,

40, 44satellite males, 60

NematodesCaenorhabditis elegans (model

system), 203Heterodera glycines (soybean cyst

nematode)mating disruption with sex

pheromone, 258, 260number of olfactory receptor

types, 172Nemeritis (parasitoid wasp)

host marking pheromone, 84(E,E,E)-Neocembrene-A

(E-6-cembrene A), 140Nereis succinea (polychaete worm)

pheromones coordinate externalfertilisation, 42

Newts, see Salamanders and newtsNezara viridula (southern green stink

bug)sex pheromone blend change due

to parasitoid, 233Nicrophorus (burying beetle), 59Niveoscincus microlepidotus (lizard)

individual recognitionpair bonding, 113

Noradrenaline, 198






INDEX 383

role in Bruce effect, 201Norepinephrine (NE), see

NoradrenalineNotropis umbratili (redfin shiner fish)

interspecific egg dumper, 231Novomessor (ant)

recruitment, 140signals, 18

Nuptial dancepolychaete worms, 42

Odocoileus hemionus columbianus (black-tailed deer), see Deer

Odorant-binding proteins (OBP), 169,177, 178

Odour sieve, 167Oecophylla

Oecophylla longinoda (African weaverant)

alarm pheromone signal, 212, 214laying trail, 135territories, 88–89

Oecophylla smaragdinaterritories, 89

Oestrogen, see Hormones, oestrogenOestrus

Microtus ochrogaster (prairie vole), 198rats, 181, 191, 193, 196synchronisation

humans, 282–283, 285mice, 194rats, 196

Olfactionacross-fibre patterning, 171amygdala

interaction between main olfac-tory and vomeronasal organ(VNO) outputs, hamster, 184

behaviour of smelling (sniffs andflicks), 174

brainmacroglomerular complex (MGC)

not needed for response to allpheromones, 188

mapping activity, 172, 173,175–176, 185

moths and sex pheromones, 165,186–188

outputs modulated, 178, 182similarities in vertebrates and

insects, 165combinatorial coding, 170glomeruli, 170–174

integration of olfactory and visualinputs, 178

main olfactory systemfunctional overlap with

vomeronasal olfactory system(VNO), 180–86

mapping brain activity, fluorescing dyes, 172, 173functional magnetic resonance

imaging (fMRI)humans, 174, 175, 185

memories for odoursrenewal of olfactory sensory

neurons (OSNs), 173odotypes

hypothesis for coding features ofodour molecules, 170, 171

olfactory receptorstypes (and organisation)nematodes, insects, and

mammals, 172sensitivity

predator olfactory sensory neurons (OSNs) as sensitive asprey’s, 231

smell (olfaction) vs. taste (gustation), 164

specificityclean-up enzymes, 177odorant-binding proteins (OBPs),

177synaptic organization, 172

insect antennal lobe, 172mammalian olfactory bulb, 172

temporal coding in brains, 174fast oscillations of neural

activity, 174humans, 175fast oscillations, synchronisation,

174Olfactory bulb

ewe learns individual odour oflamb, 199, 200

Olfactory cuesrecognition

mates, species, kin, 198Olfactory organs

amphid sensillae (nematode),203–204


biggest nasal organ for body sizein vertebrates, 207

functional design, 166–168, 174lobsters

antennule sensory hairs, 174, 221sensitivity

dogs, 167moth antennae, 166

Olfactory receptor proteins (OR), 166evolution from G-protein-coupled

receptors, 176genes, 169–170

humans, 170Olfactory sensory neurons (OSNs),

165, 166renewal, 173similarity across animal

kingdom, 169stem cells, 169

Olfactory space, 169Oncorhynchus mykiss (rainbow trout)

taught to recognise predatorodour, 254

Ondata zibethicus (muskrats)monitoring, 257

Ophrys sphegodes (orchid)dupes solitary bee, 240, 242

Optical isomers, 305Optimal outbreeding

mate choice, 51Oreotragus oreotragus (klipspringer

antelope)scent marks eavesdropped by

ticks, 233Orientation behaviour, 206

active space, 209diffusion, 210–212


mate location, 207arresting, 207chemical plumes, 206combining information from

different senses, 223directly guided, 207idiothetic, 207, 208indirectly guided, 207, 208kinesis, 207, 208

klinokinesis, 208orthokinesis, 208

odour ‘landscape’, 206odour concentration gradient, 207odour stimulus

plumes, 210–227short range diffusion, 210–213, 214






384 INDEX

Orientation behaviour, (cont.)trails, 210, 213,214–218

precise stimuli, 207ranging, 209

strategies, 209sampling

sequential, 209simultaneous, 209

scalecommunication strategies,

212, 221self-steered, 207

counter-turning, 225taxis, 207, 208

anemotaxis, 222klinotaxis, 208rheotaxis, 222teleotaxis, 208tropotaxis, 208, 213, 222

teleology, 207threshold (K), 210turbulence

transmission in currents, 158virtual-reality chemical goggles, 223

Oryctolagus cuniculus (European rabbit)nipple pheromone, 181, 183

Ostariophysi, 158Ostrinia nubilalis (European corn borer

moth)pheromone signal genetics, 69

Ourebia ourebi (oribi), territories, 96Over-marking, 99–100

Castor canadensis (beaver), 96message centre, 99scent blending, 99scent masking, 99used in mate choice in house

mice? 94Oviposition-marking pheromones, see

Host-marking pheromonesOviposition pheromones

Simulidae (blackfly), 75Culex (mosquito), 75, 77Schistocerca gregaria (desert locust), 76

Ovis aries, see SheepOxytocin

peptide hormone, 199, 200

Pachycondyla (ants)Pachycondyla laevigata

mass recruitment, 134Pachycondyla obscuricornis

individual hunter, 134

Panorpa japonica (scorpion fly)symmetrical males more

attractive, 49Panulirus interruptus (California spiny

lobster)aggregation for defence, 75

Paraleptophlebia adoptiva (mayfly), 236Paraponera clavata (ant)

eavesdropped by parasitoid fly, 233Parasitoid insects

fliesApocephalus paraponerae

(phorid), 233Trichopoda pennipes (tachinid), 233

host suicide hypothesis, 152wasps

host marking, 84Telenomus euproctidis, 232Trichogramma pretiosum, 85

Parkinson’s disease, 297Paternal investment

Utetheisa ornatrix (tiger moth),47, 49, 50

Pectinophora gossypiella (pink bollwormmoth)

males, wider pheromone responsethan range females produce, 69

pest management, 258, 267Peptide pheromones, 15

aphrodisin, 182Litoria splendida (magnificent tree

frog), 21sodefrin, 15, 16specificity, 16

Perfumes, 273, 279Periglomerular cells, 172Periplaneta americana (American

cockroach)sex pheromone, 16

Periplanone-Bsex pheromone, Periplaneta

americana (Americancockroach), 16

Petaurus breviceps (sugar glider)scent marks affect sexual

activity, 196Petromyzon marinus (marine lamprey)

pest management, 262Pharmacophagy, 12Pheidole (ants)

Pheidole dentataenemy specification, 155

Pheidole pallidula

matching foraging effort to foodvalue, 139

Pheromone binding proteins(PBPs), 177

Pheromone identification, 25bioassays, 25–29

appropriate concentrations,28, 188

examples, 24, 25–29subtractive approach, 35Y-maze, 133

by comparative approachants, 32mice, 32

challenges, 23, 36collection of chemical signals, 29

aeration and cold trap, 24antelope scent marks, 29entrainment, Poropak-QTM,

Tenax™, cold trap, 31solid phase microextraction

(SPME), 31, 32, 33–34solvent washes or whole glands,

29, 31composite (combined) signals, 29electrophysiology

electroantennogram (EAG), 29, 31electroolfactogram (EOG), 29, 189electrovomerogram (EVG), 194single cell recording (SCR), 29

humansfunctional magnetic resonance

imaging (fMRI),174, 185organoleptic tests, 289‘scratch & sniff’ odorants, 297T-shirt sniff tests, 274twins studies, 292

ionised airtracking plumes, 221

model systems, 36, 203neurotransmitter dopamine as

plume tracer, 221new techniques

brain imaging, 36genetic analysis, 36genetic engineering, 35–36, 70genomics, 35

separating chemicalsfractionation, 25, 31gas liquid chromatography (GC or

GLC), 30, 32gas liquid chromatography–mass

spectroscopy (GCMS), 32






INDEX 385

high performance liquid chromatography (HPLC), 26

signal redundancydeer alarm signal, 28

Pheromones, 1advertising

Homarus americanus (lobster), 61aquatic habitats, 15assembly

guanine, tick, 80calling pattern, by mating system

and habitatstored-product insects, 82

caste differencesmandibular gland secretions,

114, 115changes with age

elephant males, 47humans, 275queen mandibular pheromone

(QPM) blend, 122convergent, 4–6cooperative breeding, 113–127

inhibition or suppression of subordinate reproduction, 126

coordinating externalfertilisation, 41

Nereis succinea (polychaeteworm), 42

Carassius auratus (goldfish), 9, 19costs of signalling, see Costscuticular hydrocarbons

dipteran sex pheromones, 70social insects, colony odours,

106, 108direct methods of transfer, 4

into bloodstream, 4to female, 4

dominance status, 44eavesdropped by visual cues, 232egg hatching pheromone (barnacle)

eicosanoids (PUFAs), 77evolution

alarm pheromones, 9clues from geographical

variation, 69, 232–234, 248eavesdropping, 8, 234effect of habitat, 82, 97–98enabled by olfactory receptor

proteins and combinatorialbrain circuits, 176

from existing chemical cues, 9,19, 42, 43, 189

honeybees, sociality, 114sensory drive, 6, 7

glandsmammals, summary, 10wasps, bees, ants, and termites,

summary, 11glycoproteins

barnacle settlement, 78salamander, 4

graded responsevole scent glands to

testosterone, 46hormone-based, 9host marking, 83–85insect

molecular weight related to function, 14

larval aggregation pheromonesbark beetles, 81

long distanceproduced by females, 41produced by males, 41

longevitymajor urinary proteins (MUPs), 193

major urinary proteins (MUPs)elephants, 195mice, 195

mammalsaphrodisin, 182detection by main olfactory

system, 181molecular weight related to

habitat, 13, 15marking of patches

bees, 85ladybirds, 85parasitoid wasps, 85

metabolic cost, lowAnthonomus grandis (boll weevil), 20Litoria splendida (magnificent tree

frog), 21molecular weight related to habitat

aquatic, 15terrestrial, 13, 15

multi-componentalarm pheromones in social

insects, 154moths, 17, 66provide specificity in insects and

vertebrates, 17recruitment, 141

multiple messages in odoursmammals, 102

social insects, 102nipple

Oryctolagus cuniculus (Europeanrabbit), 181, 183

oviposition, 75, 76, 77peptides

aphrodisin, 182larval settlement pheromone,

barnacle, 78Litoria splendida (magnificent tree

frog), 21marine invertebrates, 15pumping pheromone, crab, 77sodefrin, 15, 16

pheromone binding proteins (PBPs)polar molecules

anthopleurine, 15prostaglandins, 9pumping, 77queen egg marking, honeybee, 122queen mandibular pheromone

(QMP), 121, 204queen pheromones in social

insects, 118–123release as pulses rare, 221reproduction in social groups, 113sex ratio effects

Microcebus murinus (lesser mouselemur), 197

sourcesplants, 11, 12, 234symbiotic bacteria, 11, 111waste products, 80

stereoisomersimportance in invertebrates and

vertebrates, 17steroids, 9synergy in multi-component

pheromonesinsects, 35mammals, 35, 195

terrestrial habitats, 13–15transmission in currents

fish alarm pheromone, 158see also Aggregation pheromones;

Alarm pheromones; Antagonists; Glands; Honeybees; Host-markingpheromones; Pheromone identi-fication; Primer pheromones;Recruitment pheromones;Releaser pheromones; Sexpheromones; Signals; Territories






386 INDEX

Philopatry, 125Phoxinus phoxinus (European minnow)

alarm pheromone, 158Pieris (cabbage butterfly)

host-marking pheromone, 84Pigs

minipig pets, 254see also Sus scrofa

Piriform cortex, 165Plankton,

commercial applications, disrupt-ing settlement, 266

settlementbarnacle, 26, 78Dendraster excentricus (sand

dollar), 80Hydroides (polychaete worm), 79

3D trailsTemora longicornis (copepod), 213

Plantsdeception by wild potatoes, 244bee pheromones from orchids, 12pyrrolizidine alkaloids (PAs), 12, 47tree defences against bark

beetles, 81see also Pheromones, sources, plants

Platysoma cylindrica, eavesdropping,234, 236

PlethodonPlethodon cinereus (red-backed

salamander)contest, 46feeding and breeding

territories, 46scent matching hypothesis and

territory, 95Plethodon jordani (salamander)

pheromone transfer by male, 4postcloacal gland, 95

Plotsus lineatus (catfish), 169Plumes

turbulence, visualisation, 219tracking plumes, 221

ionised air, 221neurotransmitter dopamine, 221

Podisus maculiventris (spined soldierbug)

eavesdropped by parasitoid, 233Pogonomyrmex badius (harvester ant)

alarm pheromone signal, 212Poison gland, ant, 130Polistes fuscatus (paper wasp)

kin recognition mechanisms, 107

Popilla japonica ( Japanese beetle), 17Postcloacal gland

salamander, 95Postpharyngeal gland

ants, 108host source of hydrocarbons for

parasite, 248Precopulatory mate guarding, see

Sperm competition and mateguarding

Predators and parasitoids, 2324-Pregnen-17�-,20�-diol-3-one

(17,20�-P), 9Pregnancy block, male pheromones

(Bruce effect), 195Premnas (anemone fish), 238Preorbital gland, see Antorbital glandPreputial gland

mice, 193Primates

Callithrix jacchus (common mar-moset)

subordinate female ovulationsuppressed, 197

Cerocebus albigena (mangabey), 271Leontopithecus rosalia (lion tamarin)

cooperative breeding not odourcontrolled, 127

Microcebus murinus (lesser mouselemur)

bias of offfspring sex ratios, 197female urine stimulates sper-

matogenesis and testosteronesecretion, 196

sexual activity of subordinatessuppressed, 196

micro-osmic, 271olfactory receptors, 169–170

genes, 170Saguinus fuscicollis (saddle-back

tamarin)multiple messages in odours, 102subordinate female ovulation

suppressed, 197scent glands and scent

marking, 272see also Humans

Primer pheromones, 18can be fast acting, 20Carassius auratus (goldfish), 19gregarising factor in egg foam

Schistocerca gregaria (desert locust), 76

juvenile hormone ( JH)social insects, 204ants, 204

maturation synchronySchistocerca gregaria (desert

locust), 76pest management, 266reproduction, 192

daylength and pheromones insheep, 192

similiarity of action in mammalsand insects, 20

social insectsqueen effects in ants, 204

termitescaste control, 119

Primiphales promelas (fathead minnow)alarm pheromone, 158

3-Propyl-1,2-dithiolane, 265Prostaglandins, 9

15-keto-prostaglandin-F-2�

(15-keto-PGF2�), 9fish peripheral response, 189

prostaglandin F-2� (PGF2�), 9Prostephanus truncatus (larger grain

borer beetle), 256males only signal until females

come, 82monitoring, 256

Proteles cristatus (aardwolf)‘dear-enemy’ individual

recognition, 113Protocerebrum, 165Proust, memory, 276Pseudaletia (moth), 5Pseudogates

termites, 119Pseudogenes, 170Puberty acceleration, see HormonesPUFAs (C20 polyunsaturated fatty

acids), 77Puntius (cyprinid fish), 189Pygidial gland, ant, 134Pyrrolizidine alkaloids (PAs), 12, 47

Queen mandibular pheromone (QMP),see Pheromones

Queen pheromones in social insects,see Pheromones

Racemate or racemic mixture, 306Ranging behaviour, 209Raphicerus melanotis (grysbok), 13






INDEX 387

Rattus norvegicus (Norway rat)follows trails from good food

sources, 137Recognition mechanisms, 103

direct familiarisation, 104indirect familiarisation (phenotypic

matching), 104Microtus ochrogaster (prairie vole),

197–198phenotypic matching hypothesis

field tests, 111recognition allele (‘green beard’

phenomenon), 104, 105fire ants, 105

sensitive period for learning, 105social insects

learning needed before discrimination, 103–109

usually by learning, 103–104Recruitment pheromones

alarmApis cerana japonica (Japanese

honeybee), 154, 156–157marking enemies for further

attack, 153, 154, 155ants

competition strategy, 140group recruitment, 134laying trail, 135mass communication, 129, 134scout ants, 134tandem running, 134trail specificity, 134worker sensitivity to trail

pheromone, 136aphids, social

soldiers attack enemy, 153, 153competition strategy, 140

ants, 140Trigona (stingless bee), 140

convergence in markingbehaviour, 135

coordinated attackVespa mandarinia japonica (giant

hornet), 154–155, 157expand colony diet, 136food distribution, recruitment

type, 136longevity related to food

supply, 137Malacosoma americanum (tent

caterpillar)laying trail, 135

matching foraging effort to foodvalue, 138

Rattus norvegicus (Norway rat)follow trails from good food

sources, 137scout ants, 134termites, 137

laying trail, 135trail specificity, 135

trail specificity, 140see also Alarm pheromones; Self-

organising systemsReinforcement, 65Releaser pheromones, 18

Carassius auratus (goldfish), 19mammal, 20nippleOryctolagus cuniculus (European rab-

bit), 181Reproductive character

displacement, 65Reproductive skew, 116, 117Response to pheromones, factors

affecting, 188, 191context

aphids tended by ants, 237honeybees, 188parasitoid wasps, 188

hormonescentral nervous system

effects, 189peripheral effects, 189

receiver’s characteristics, 188see also Learning; Development

Reticulitermies f lavipes (termite)recruitment pheromones, 140

Reynolds numbers (Re), 167copepod pheromone trails, 213

RhagolitisRhagolitis cerasi (cherry fruit fly)

pest management, 264Rhagolitis pomonella (apple

maggot fly)Crataegus mollis (hawthorn),

original host, 85evolution of host-marking

behaviour, 85experience of marking

pheromone needed forresponse, 190–191

host marking pheromone, 84Rhitropanopeus harrisii (mud crab)

pumping pheromone, 77

Ritualised signals, 3Ropalidia marginata (paper wasp)

non-kin may join colony, 108

Saguinus fuscicollis (saddle-backtamarin)

multiple messages in odours, 102subordinate female ovulation

suppressed, 197Salamanders and newts

courtship pheromone delivery patterns, 4

Cynops pyrrhogaster (red-belliednewt)

courtship, 16sodefrin, 16

Desmognathus ochrophaeusmale injects pheromone into

female, 4Plethodon cinereus (red-backed

salamander)postcloacal gland, 95scent matching, 95

Sanguinis fuscicollis (saddle-backtamarin)

multiple messages in odours, 102subordinate female ovulation

suppressed, 197Satellite males, see Alternative mating

stategiesSaturnia pavonia (emperor moth)

early demonstration ofpheromone, 23

Scaleboundary layer

effects on olfactory sensors, 174chemical plumes, 206communication strategies, 212design of olfactory organs, 166–168diffusable pheromones in water, 213Reynolds numbers (Re), 167, 213

Scent markingborder maintenance hypothesis, 90,

96–97Castor canadensis (beaver), 96ecological factors, patterns between

and within related species, 91males directly mark females

Dolichotis patagonum (mara), 100marking behaviour of house mice, 94non-territorial mammals, 100response by mice varies by

competitive ability, 94






388 INDEX

Scent marking (cont.)salamanders, nose-tapping, 95scent fence hypothesis, 91scent matching hypothesis, 90,

91–96mice, 93–94reducing cost of territorial

defence, 93salamanders, 95self-marking and presentation, 92testing the hypothesis, mice, 95

using vertebrate scent marks to census, 257

Schedorhinotermes (termites), 5Schedorhinotermes lamianus

foraging and soldiers/workers,138

pheromones guide food collection, 138

Schistocerca gregaria (desert locust)gregarising factor in egg foam, 76maturation synchrony, 76oviposition pheromone, 76pheromones in life cycle, 76

Scramble competition, 43–44Carassius auratus (goldfish), 43moth males responding to female

pheromones, 43selection for male moth sensitivity,

43, 166Thamnophis sirtalis parietalis

(red-sided garter snake), 43Sebaceous gland

birds, 21humans, 284signal life, 14

Sebum, 13Self-organising systems, 141

ant foraging, 141Eciton (army ants), 142matching resources to effort, 143modelling food distribution, 142stochastic effects, 143

termite nests, 143–145stigmergy, 143

‘Selfish herd’ theory, 159Semen

behavioural manipulation byDrosophila melanogaster, 63humans? 64Thamnophis sirtalis parietalis

(red-sided garter snake), 63–64Semibalanus balanoides (barnacle)

egg hatching pheromoneeicosanoids (PUFAs), 77

Semiochemicals, 1Sensitive periods, 201

ewe learning odour of her lamb,199, 200

hormonal state and learning, 199sex ratio effects

Microcebus murinus (lesser mouselemur), 197

termite caste development, 205Sensory drive, 6, 7Serricorone, 84Settlement-inducing protein complex

(SIPC), 78Settlement of marine vertebrates, 77–79Sex differences

in brainmacroglomerular complex (MGC)

in male moths, 190response to pheromones

Carassius auratus (goldfish), 189sensitivity to 3�-androstenol in

sows, 253see also Development

Sex pheromonesaphrodisin, 182Aplysia (sea-slug)

peptide sex pheromone, 15arachnids

Linyphia litogiosa (spider), 59crustaceans

Homarus americanus (lobster), 61, 62Temora longicornis (copepod), 213

exploitationcommercial, 253, 254natural, 231, 232–233

insectsCeratitis capitata (Mediterranean

fruit fly), 58, 59Lutzomyia longipalpis (sandfly), 58Nicrophorus (burying beetle), 59Periplaneta americana (American

cockroach), 16Xylocopa fimbriata (carpenter

bee), 57mammals, role of experience, 182sodefrin, Cynops pyrrhogaster (newt),

15, 16speciation, 64see also Development; Female

pheromones; Malepheromones

Sex ratioMicrocebus murinus (lesser mouse

lemur)bias of sex ratio in pregnancy,

197Sexual selection, 37

contests, same-sex, 38Homarus americanus (lobster), 44Nauphoeta cinerea (African cock-

roach), 40, 44female choice

elephants, 47Ephestia elutella (tobacco moth), 47Nauphoeta cinerea (African

cockroach), 40, 44male characters

coremata, 3, 10, 11, 47, 49mate choice, 37–38mate quality and courtship, 46mechanisms

direct benefit, 38, 39imprinting, 40indicator, 40runaway (‘Fisherian’, ‘sexy sons’),

38, 39, 59scramble competition, 43–44sexual competition, 38which sex should call? 40

‘Sexy faeces’ hypothesisPlethodon cinereus (red-backed

salamander), 46She-males, see Alternative mating

strategiesSheep (Ovis aries)

mate choice not universal, 56maternal behaviour, 199–201mother–infant recognition, 109reproduction, 192synchronised oestrus, 252

Signals, 1–2anonymous

social insects, 102butterflies

visual cues and pheromones, 14composite, 17–18

parallel sensory channels, 17with ultraviolet cues, 14

cooperative, 122design

aquatic habitats, 15contrasting modalities, 12habitat and daily activity, 13function and environment, 13






INDEX 389

lizard signals in contrasting habitats, 14

longevity, selection on, 13molecular weight and function,

13molecular weight and habitat,

13–15terrestrial habitats, 13

Dipsosaurus dorsalis (iguana)composite signal with ultraviolet

cue, 14dishonest, 121distinguishing chemical signals

from noise, 206honest, 3, 22, 44, 45

queen pheromoneshoneybees, 121, 121territorial scent marks, 89Utetheisa ornatrix (tiger moth),

47, 49, 49, 50low metabolic cost of pheromones

Anthonomus grandis (boll weevil), 20

Litoria splendida (magnificent treefrog), 21

mechanisms for specificitypeptides (amino acid sequence), 16unique molecules, 16multi-component, 17stereoisomerism, 17

modulationintensity, 17by sound, 18of visual threat by

pheromones, 18ON–OFF

clean-up enzymes, 177recruitment, 129

ants, 134redundancy, 17

deer alert signal, 28, 148ritualised, 3sexually selected, 37synergy, 35see also Alarm pheromones;

Pheromones, evolutionSimulidae (blackfly)

oviposition pheromone, 75, 263Singular breeders, 124Sitophilus (beetle), 83Smell, see OlfactionSnakes

Boiga irregularis (brown tree snake)

control, 262detecting trail direction, 217, 217Leptotyphlops dulcis (blind snake)

follows ant pheromone trails, 247tongue-flicking

trail following, 215Thamnophis sirtalis parietalis

(red-sided garter snake)overwintering aggregations, 80

Vipera berus (adder)Coolidge effect, 57

Sniffingbehaviour of smelling, 174

Social insectsalarm pheromones, 152–157ants

Atta texana, 136Oecophylla longinoda (African

weaver ant), 129, 212, 214Solenopsis invicta (fire ant), 123

aphidsalarm pheromones, 152–153Ceratovacuna lanigera, 153Colophina monstrifica, 153

caste differencespheromone secretions, 114reproductives and workers, 118response, 153

caterpillarsMalacosoma americanum (tent

caterpillar), 133colony odours

variation, 248guard bees

defense of colonies, 105discrimination of relatedness, 108recognition mechanisms, 105

Polistes fuscatus (paper wasps)kin recognition mechanisms, 107

Ropalidia marginatanon-kin may join colony, 108

parallels with social mammals, 127pheromones, 114primer pheromones

development of ant castes, 204queen mandibular pheromone

(QMP), 121action via juvenile hormone ( JH)

levels, 204queen pheromones, 118–123

control or cooperative signal, 118evolution, 122fire ants, 123

reproductive division of labour, 116termites

colony recognition, 109territories

Oecophylla longinoda (Africanweaver ant), 88–89

Oecophylla smaragdina (ant), 89Trigona (stingless bees)

odour beacons rather than trails,136

worker policing, 122see also Honeybees; Recruitment

pheromones; Self-organisingsystems

Social mammalsclan recognition, 111

Meles meles (European badger) , 111cooperative breeding, 124–127

evolution, 125inhibition or suppression of

subordinate reproduction, 126control by pheromones, 124, 126control not by pheromones, 124,

125, 127kin recognition

Castor canadensis (beaver), 111Spermophilus beldingi (Belding’s

ground squirrel), 110parallels with social insects, 112,

127primer pheromones and

reproduction, 123–124recruitment, 136

Heterocephalus glaber (naked molerats), 134

reproductive division of labour, 116shared colony odour, parallels

with social insects, 112Sodefrin, peptide pheromone, 15, 16Solanum berthaultii (wild potato),

255, 245Solenopsis (fire ants), 130, 137

Solenopsis invictaenemy specification, 155mass communication

recruitment, 134queen mutual inhibition, 123

Solid phase microextraction (SPME),31, 32

Speciation, 64–72allopatric, 64, 65Drosophila melanogaster

cuticular hydrocarbons, 70






390 INDEX

Speciation, (cont.)reproductive character

displacement, 65male pheromones, selection by

female to avoid hybrid matings, 70

signal shift, 68asymmetric tracking, 68

sympatric, 64, 65, 66Specificity of male moth responses to

pheromones, 68Specific-Mate Recognition System, 65Sperm competition and mate

guarding, 61–64cryptic female choice, 63

Tribolium castaneum (flour beetle),63

Utethiesa ornatrix (tiger moth), 63precopulatory mate guarding,

61–62consorting by male kangaroos, 62elephants, 62Homarus americanus (lobster), 61, 62

Spermophilus beldingi (Belding’sground squirrel)

kin recognition cues, 110Spindasis lohita (lycaenid butterfly),

238Squalene, 43‘Standing behaviour’, see LordosisStem cell, olfactory, 168–169Stereoisomers, 305Sternal gland

ant, 130Malacosoma americanum (tent

caterpillar), 130Polistes (paper wasp), 107termites, 130, 137

Steroids, 95�-cholestane 3,24-dione

Malacosoma trail pheromone, 130Stigmergy, see Self-organising systemsStimulo-deterrent diversionary

strategy (SDDS), 265Stochastic effects, 143Stored product insects

calling pattern, by mating systemand habitat, 82

Lasioderma serricorne (cigarette beetle), 83

Prostephanus truncatus (larger grainborer beetle) , 82, 256

Sitophilus (beetle), 83

Trogoderma granarium (Khapra beetle), 83

Stria terminalis, 183Subcaudal gland

Meles meles (European badger), 111Superparasitism, 83Sus scrofa (pig)

female response3�-androstenol, 20, 184, 2535�-androstenone, 184

lordosis by oestrous sow to malepheromones, 20, 184, 253

Swarming, see HoneybeesSweat glands

apocrine, 284eccrine, 284

Synergy, 35Synomones, 2

mutualism, 230

Tadpoles, kin, and alarmpheromones, 157

Talpa europaea (European mole), 91Taste

smell (olfaction) vs. taste(gustation), 164

taste buds, 164Taxis, 207, 208‘t-complex’ in mice, 56Telenomus euproctidis (parasitoid wasp)

eavesdrops host, 232Teleology, 207Temnochila chlorodia (beetle), 234Temora longicornis (copepod), 3D trails,

213Temporal gland

elephants, 46Termites

beetle mimics its termite host, 248caste-change pheromones, 119colony recognition odours, 109developmental pathways, 120pest management, 263primer pheromones, 119soldiers, control of numbers, 119

Termitophilous invertebrates, 245Territories, 87

costs (of signalling)Oreotragus oreotragus (klipspringer

antelope), 233dear-enemy phenomenon, 99economics of scent marking, 97

patterns, 97

variation by habitat inHyaenidae, 98

feeding and breedingPlethodon cinereus (red-backed

salamander), 46group territorial defence

Lemur catta (ring-tailed lemur)‘stink fights’, 90

‘owner advantage’Oecophylla longinoda (African

weaver ant), 88scent marking

border maintenance hypothesis,90, 96–97

composite signals withultraviolet cue, 99

lamp-post effect, 99social insects

Oecophylla longinoda (Africanweaver ant), 88–89

Testosterone, see Hormones,testosterone

Tetramorium caespitum (ant), foraging,143

Thamnophis sirtalis parietalis (gartersnake)

conflict between the sexessemen components, 63–64

detecting trail direction, 217, 217male pheromone, squalene, 43overwintering aggregations, 80she-males, 60

Thanasimus dubiuseavesdropping, 234

Thiazole, see Butyl dihydrothiazoleTicks

Amblyomma hebraeumpest management, 262

Dermacentor variabilispest management, 262

guanine, assembly pheromone, 80Ixodes neitzi responds to scent marks

of host antelope, 233Tools to study pheromones, see

Pheromone identificationTongue-flicking, snakes, 215Trail pheromones, see Recruitment

pheromonestrans (in chemical name), 305Tree defences, resins and toxic

chemicals, 81Tribolium castaneum (flour beetle), 82

cryptic female choice, 63






INDEX 391

Trichogramma pretiosum (parasitoidwasp), 85

Trichoplusia ni (cabbage looper moth)both sexes call, 41mutation in pheromone blend, 68male response requires hormone, 189

Trichopoda pennipes (tachinid)selection presure on host, 233

Trichopsenius frosti (beetle)matches its termite host, 248

(Z)-7-Tricosene, 61(E)-4-Tridecenyl acetate, 259Trigona (stingless bees), 5, 136, 140

Trigona angustularesists pheromone propaganda

by robber bee, 243Trigona subterranea

victim of pheromone propagandaby robber bee, 243

Trogoderma granarium (Khapra beetle),83

Trophallaxisoral, 119proctodeal, 119

Truffles (Tuber melanosporum)3�-androstenol, 253use of sows to find, 253human taste for, 288

Tuber melanosporum, see TrufflesTufted cells, 172

Ultraviolet (UV) cuesbirds eavesdrop vole scent marks

by UV, 232Dipsosaurus dorsalis (iguana),

composite signal with UV, 14Undercrowding

Allee effects, 75Urine

active only on contactMicrotus ochrogaster (prairie vole),

197–198dominant male effective, 192Homarus americanus (lobster)

contest, 44major urinary proteins (MUPs)

elephants, 195mice, 195

reflects hormonal and dominancestates in rodents, 46

Uropygidial gland, 21Utetheisa ornatrix (tiger moth)

female choice, 3, 63

honest signal, 47, 49, 49, 50hydroxydanaidal (HD) from diet, 47sexual selection, 39, 70

Vaginocervical stimulation duringbirth

hormonal releasers, 200Vandenbergh effect (puberty

acceleration), 192Vanillic acid, 260Varroa mite

pest management, 262response to honeybee larval

odours, 116Ventral amygdalofugal pathway, 183Verbenone, 235Veromessor (ant)

benzaldehyde as defensive compound, 5

Vertebratesandrogens

gland secretion rates, 46dual olfactory system, 178pheromones to attract mates, 41see also Amphibians; Antelopes;

Deer; Dogs; Fish; Mammals;Mice; Pheromones; Primates;Salamanders and newts;Social mammals; Wolves; othertaxonomic entries

Vespa mandarinia japonica (giant hornet), 154, 156–157

Vespula (wasps)Vespula germanica, 155

eavesdropping prey leks, 232, 231Vespula vulgaris, 155

Vipera berus (adder), Coolidge effect, 57Vomeronasal organ (VNO), 165,

178–185female VNO, urine contact, 198induction of oestrus

VNO needed, 181VNO not needed, 181

Vomeronasal amygdala, 165Vomeronasal olfactory system, 165

functional overlap with main olfactory system, 180–186

in humans? 294memory and pregnancy block

(Bruce effect) in mice, 201molecular architecture and

differences from main olfactory system (MOE),

179–180mapping to glomeruli, 180receptor proteins, 179receptor specificity, 179response to proteins, 180

some pheromones detected bymain olfactory system(MOE), 181

trail following by snakes, 215–217

Westermarck or kibbutz effecthuman mate choice, 280

Whitten effect (oestrus induction),192

WolvesCanis simensis (Ethiopian)

border marks in territory, 97Canis lupus (grey)

border marks in territory, 97control of reproduction, 124

Worker policing, 122

Xylocopa fimbriata (carpenter bee),leks, 57

Yponomeuta (small ermine moths)calling time, 66host plants, 66, 67pheromone blend, 66, 67

Zaspilothynnus trilobatus (thynninewasp)

duped by orchid, 241Zigzags

swimming or flying up plumes, 224Zootermopsis nevadensis (termite)

nymph laying trail, 135Zusammen, (Z) together, 308






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Index [assets.cambridge.org]€¦ · Aeolidia papillosa (nudibranch sea-slug) predator marked by prey pheromone, 148 Aeshena umbrosa (dragonfly nymph) predator, 157 African mole rats,