Functional and Evolutionary Ecology of Fleas: A Model for Ecological Parasitology

Cover......Page 1
Half-title......Page 3
Title......Page 5
Copyright......Page 6
Contents......Page 7
Preface......Page 11
PART I BRIEF DESCRIPTIVE ECOLOGY: WHAT DO FLEAS DO?......Page 17
1.1 Infraorders and families......Page 19
1.2 Temporal pattern of discovery of flea species......Page 20
2.1 Avian and mammalian hosts......Page 25
2.2 ‘Realized’ and available hosts......Page 26
2.3 Number of flea species among host orders......Page 31
2.4 Fleas, small mammals and biogeography......Page 32
2.5 Concluding remark......Page 33
3.1 General patterns of geographical distribution......Page 34
3.2 Fleas on islands......Page 37
3.3 Size of geographical range......Page 38
3.4 Relationship between flea and host(s) geographical ranges......Page 41
3.5 Concluding remark......Page 44
4.1 Ancestral and sister taxa......Page 45
4.2 Origin of flea parasitism......Page 48
4.3 Phylogenetic relationships within Siphonaptera......Page 49
4.4 Cophylogeny of fleas and their hosts......Page 50
4.5 Flea diversification: intrahost speciation, host-switching or climate?......Page 57
4.6 Concluding remarks......Page 60
5.1 Mating and oviposition......Page 61
5.2 Larvae......Page 64
5.3 Pupae......Page 67
5.4 Imago......Page 68
5.5 Seasonality......Page 70
5.5.1 Examples of annual cycles......Page 71
5.5.2 Reproductive diapause and overwintering......Page 74
5.5.3 Classification of annual cycles......Page 75
5.5.4 Variation in seasonality: host biology, climate and evolutionary constraintsclimate and evolutionary constraints......Page 80
5.6 Concluding remark......Page 83
6.1.1 Dermatological diseases caused by flea parasitism......Page 84
6.1.2 Fleas as vectors of infectious diseases......Page 85
6.2 Veterinary aspects......Page 88
6.3 Fleas in human habitats......Page 90
6.4 Concluding remarks......Page 92
PART II FUNCTIONAL ECOLOGY: HOW DO FLEAS DO WHAT THEY DO?......Page 93
7.1.1 Size dimorphism and Rensch’s rule......Page 95
7.1.2 Shape dimorphism......Page 97
7.2 Physiological gender differences......Page 98
7.2.1 Feeding parameters......Page 99
7.2.2 Metabolic rate......Page 101
7.2.3 Water content......Page 105
7.3 Gender differences in behaviour......Page 106
7.4 Responses to environmental factors......Page 107
7.5 Ecology of sex ratios......Page 109
7.6 Concluding remarks......Page 118
8.1 On-host locomotion......Page 119
8.2.1 Mechanics of a flea jump......Page 120
8.2.2 Jumping capacity, sexual size dimorphism and morphology......Page 121
8.2.3 Jumping performance and metabolic rate......Page 124
8.2.4 Ecological correlates of jumping performance......Page 127
8.2.5 Jumping rate and activity......Page 128
8.2.6 Walking and climbing......Page 129
8.3 Concluding remark......Page 130
9 Ecology of host selection......Page 131
9.1.1 Definitions......Page 132
9.1.2 Flea abundance and the taxonomic distance among host species......Page 133
9.1.3 Geographical change of a principal host......Page 137
9.2.1 Density dependent host selection......Page 138
9.3.1 Are fleas ‘ ideal’? Host search and location......Page 146
Signals extrinsic to the host......Page 147
Signals intrinsic to the host......Page 152
9.3.2 Are fleas ‘free’? Dispersal and host-to-host transfer......Page 156
9.4 Distribution of fleas on the body of a host......Page 157
9.4.1 Sessile and stick-tight fleas......Page 158
9.4.2 Non-sessile fleas and host grooming......Page 160
9.4.3 Microclimatic differences, interspecific competition or the effect of
the original host?......Page 161
9.5.1 ‘Body’ and ‘nest’ fleas......Page 163
9.5.2 Reliability of body infestation data......Page 165
9.6 Concluding remarks......Page 169
10.1 Mouthparts and host skin......Page 170
10.2 Measures of feeding success......Page 172
10.2.1 Blood meal size and rate of engorgement......Page 173
10.2.2 Rate of digestion......Page 175
10.2.3 Energetic cost of blood digestion......Page 176
10.2.4 Number of blood meals necessary for oviposition......Page 177
10.3.1 Host species......Page 178
10.3.2 Host gender and age......Page 186
10.3.3 Host body conditions......Page 187
10.4.1 Interspecific variation in feeding patterns......Page 189
10.4.2 Intraspecific variation in feeding patterns......Page 190
10.5 Environment-related effects......Page 194
10.6 Concluding remarks......Page 197
11.1 Measures of reproductive success......Page 198
11.2.1 Host species......Page 201
11.2.2 Host gender and age......Page 207
11.2.3 Host body conditions......Page 208
11.3.1 Flea species......Page 215
11.3.2 Flea age......Page 216
11.3.3 Flea density......Page 217
11.4.1 Air temperature and relative humidity......Page 222
11.4.2 Substrate texture......Page 230
11.4.3 Light regime, photoperiod and host odour......Page 231
11.5 Concluding remarks......Page 232
12 Ecology of flea virulence......Page 233
12.1 Host metabolic rate......Page 234
12.2 Host body mass and growth rate......Page 239
12.3 Host haematological parameters......Page 243
12.4 Host features related to sexual selection......Page 244
12.5 Host behaviour......Page 245
12.6 Host survival......Page 248
12.7 Host fitness......Page 251
12.8 Concluding remarks......Page 253
13 Ecology of host defence......Page 255
13.1.1 Avoidance in space......Page 256
13.1.2 Avoidance in time......Page 258
13.2.1 Managing nest infestation......Page 259
13.2.2 Managing body infestation......Page 261
13.2.3 Neurophysiology of anti-parasitic grooming......Page 266
13.2.4 Trade-offs of the behavioural defence against fleas......Page 268
13.2.5 Withstanding host grooming......Page 270
13.3 Third line of defence: immune response against fleas......Page 271
13.3.1 Third line of defence: immune response......Page 273
13.3.2 Acquired resistance......Page 274
13.3.3 Cross-reactivity......Page 277
13.3.4 Immunocompetence......Page 279
13.3.5 Host gender and immune response......Page 282
13.3.6 Immune responses and maternal effects......Page 284
13.3.7 Immune responses and infestation patterns......Page 286
13.3.8 Costs and trade-offs of the immune defence against fleas......Page 291
13.3.9 Coping with the host immune response......Page 293
13.4 Concluding remarks......Page 294
PART III EVOLUTIONARY ECOLOGY: WHY DO FLEAS DO WHAT THEY DO?......Page 297
14.1 Measures of host specificity......Page 299
14.2.1 Patterns of host specificity......Page 301
14.2.2 Is host specificity a flea species character?......Page 302
14.2.3 Geographical variation of host specificity......Page 305
14.2.4 Distribution of specialization and flea species richness......Page 310
14.3.1 Abundance and fitness achieved in hosts......Page 311
14.3.2 Geographical range......Page 316
14.4 Host specificity and host features......Page 319
14.4.1 Predictability: host body mass and longevity......Page 320
14.4.2 Predictability: host abundance, coloniality and spatial distribution......Page 322
14.4.3 Host defensibility......Page 323
14.5.1 Is host specificity directional and irreversible?......Page 324
14.5.2 Is host specificity evolutionarily heritable?......Page 328
14.5.3 Coevolution and host opportunism......Page 329
14.6 Applicative aspects of host specificity studies......Page 330
14.6.1 Probability of flea species discovery......Page 331
14.6.2 Transmission of the plague pathogen......Page 334
14.7 Concluding remarks......Page 336
15 Ecology of flea populations......Page 337
15.1 Measuring abundance and distribution......Page 338
15.2 Is abundance a flea species character?......Page 340
15.3.1 Are fleas aggregated among hosts?......Page 344
15.3.2 Why does flea aggregation vary interspecifically?......Page 345
15.3.3 Why does flea aggregation vary intraspecifically?......Page 349
15.3.4 Dynamics of flea aggregation: does an uninfested host stay
uninfested for ever?......Page 351
15.4.1 Host species......Page 354
15.4.2 Host gender......Page 356
15.4.3 Host age......Page 362
15.5 Relationship between flea abundance and prevalence......Page 368
15.6.1 Host density......Page 372
15.6.2 Inferring regulating mechanisms......Page 380
15.6.3 Host spatial behaviour......Page 384
15.6.5 Environmental factors......Page 387
15.7 Concluding remarks......Page 390
16 Ecology of flea communities......Page 391
16.1.1 Patterns of species co-occurrences......Page 393
16.1.2 Nested pattern......Page 398
16.2 Local versus regional processes governing flea communities......Page 402
16.3.1 Patterns of interspecific interactions inferred from patterns of abundance......Page 407
16.3.2 Patterns of interspecific interactions inferred from patterns of aggregation......Page 412
16.4 Negative interspecific interactions......Page 417
16.5 Similarity in flea communities: geographical distance or similarity in host composition?......Page 421
16.6 Concluding remarks......Page 424
17 Patterns of flea diversity......Page 426
17.1 Flea diversity and host body......Page 427
17.3 Flea diversity and host population......Page 431
17.4.1 Flea diversity among host species and the effect of host community structure......Page 433
17.4.2 Flea diversity among host communities......Page 435
17.5.1 Patterns within a host species......Page 440
17.6 Flea diversity and the off-host environment......Page 441
17.6.1 Flea diversity and productivity......Page 443
17.6.2 Flea diversity and elevation......Page 445
17.6.3 Flea diversity and latitude......Page 446
17.7 Parasites of other taxa......Page 447
17.8 Concluding remarks......Page 451
18 Fleas, hosts, habitats......Page 452
18.1.1 Description of habitats, hosts and fleas......Page 453
18.1.2 Habitat variation in flea species composition......Page 454
18.1.3 Mechanisms of distribution of the two Xenopsylla species......Page 458
18.2.1 Description of habitats, hosts and fleas......Page 459
18.2.2 Habitat variation in flea species composition......Page 461
18.3 Other examples......Page 467
18.4 Concluding remarks......Page 470
19
What further efforts are needed?......Page 471
19.1 Where are we now and what do we have?......Page 472
19.2.1 Multifaceted vision......Page 474
19.2.2 Flea—host associations as integral entities......Page 475
19.2.3 External validity......Page 476
19.2.4 Laboratory, field and comparative approaches......Page 477
19.3 Not only a pure science . . .......Page 479
References......Page 482
Index......Page 599