Prefaces xiii
About the Companion Website xix
1 Introduction to the Study of Animal Populations 1
1.1 Population estimates 2
1.1.1 Absolute and related estimates 2
1.1.2 Relative estimates 3
1.1.3 Population indices 4
1.2 Errors and confidence 4
References 5
2 The Sampling Programme and the Measurement and Description of Dispersion 7
2.1 Preliminary sampling 7
2.1.1 Planning and fieldwork 7
2.1.2 Statistical aspects 10
2.2 The sampling programme 16
2.2.1 The number of samples per habitat unit (e.g. plant, host or puddle) 16
2.2.2 The sampling unit, its selection, size and shape 20
2.2.3 The number of samples 21
2.2.4 The pattern of sampling 24
2.2.5 The timing of sampling 26
2.3 Dispersion 27
2.3.1 Mathematical distributions that serve as models 28
2.3.2 Biological interpretation of dispersion parameters 40
2.3.3 Nearest-neighbour and related techniques: measures of population size or of the departure from randomness of the distribution 48
2.4 Sequential sampling 51
2.4.1 Sampling numbers 51
2.5 Presence or absence sampling 55
2.6 Sampling a fauna 57
2.7 Biological and other qualitative aspects of sampling 59
2.8 Jack knife and Bootstrap techniques 60
References 62
3 Absolute Population Estimates Using Capture–Recapture Experiments 77
3.1 Capture–recapture methods 78
3.1.1 Assumptions common to most methods 79
3.1.2 Estimating closed populations 86
3.1.3 Estimations for open populations 93
3.2 Methods of marking animals 103
3.2.1 Handling techniques 105
3.2.2 Release 107
3.2.3 Surface marks using paints and solutions of dyes 108
3.2.4 Dyes and fluorescent substances in powder form 112
3.2.5 Pollen 114
3.2.6 Marking formed by feeding on or absorption of dyes 114
3.2.7 Marking by injection, panjet or tattooing 116
3.2.8 External tags 116
3.2.9 Branding 117
3.2.10 Mutilation 118
3.2.11 Natural marks, parasites and genes 118
3.2.12 Rare elements 119
3.2.13 Protein marking 120
3.2.14 Radioactive isotopes 120
3.2.15 Radio and sonic tags 120
References 121
4 Absolute Population Estimates by Sampling a Unit of Habitat – Air, Plants, Plant Products and Vertebrate Hosts 139
4.1 Sampling from the air 139
4.2 Sampling apparatus 140
4.2.1 Exposed cone (Johnson–Taylor) suction trap 140
4.2.2 Enclosed cone types of suction trap including the Rothamsted 12 m trap 141
4.2.3 Rotary and other traps 143
4.3 Comparison and efficiencies of the different types of suction traps 144
4.3.1 Conversion of catch to aerial density 145
4.3.2 Conversion of density to total aerial population 146
4.4 Sampling from plants 146
4.4.1 Assessing the plant 147
4.4.2 Determining the numbers of invertebrates 147
4.4.3 The extraction of animals from herbage and debris 155
4.4.4 Methods for animals in plant tissues 163
4.4.5 Special sampling problems with animals in plant material 165
4.5 Sampling from vertebrate hosts 166
4.5.1 Sampling from living hosts 166
4.5.2 Sampling from dead hosts 169
4.5.3 Sampling from vertebrate ‘homes’ 170
References 171
5 Absolute Population Estimates by Sampling a Unit of Aquatic Habitat 183
5.1 Open water 183
5.1.1 Nets 183
5.1.2 Pumps 187
5.1.3 Water-sampling bottles 187
5.1.4 The Patalas–Schindler volume sampler 187
5.1.5 Particular methods for insects 188
5.2 Vegetation 190
5.2.1 Floating vegetation 191
5.2.2 Emergent vegetation 192
5.2.3 Submerged vegetation 194
5.3 Bottom fauna 195
5.3.1 Hand net sampling of forest litter 196
5.3.2 Sampling from under stones 197
5.3.3 The planting of removable portions of the substrate 199
5.3.4 Cylinders and boxes for delimiting an area 200
5.3.5 Trawls, bottom sledges and dredges 201
5.3.6 Grabs 205
5.3.7 Dendy inverting sampler 208
5.3.8 Box samplers and corers 209
5.3.9 Air-lift and suction devices 211
5.4 Poisons and anaesthetics used for sampling fish in rock pools and small ponds 211
References 213
6 Absolute Population Estimates by Sampling a Unit of Soil or Litter Habitat: Extraction Techniques 221
6.1 Sampling 221
6.2 Bulk staining 224
6.3 Mechanical methods of extraction 224
6.3.1 Dry sieving 224
6.3.2 Wet sieving 225
6.3.3 Soil washing and flotation 226
6.3.4 Flotation separation of plankton, meiofauna and other small animals 229
6.3.5 Separation of plant and insects by differential wetting 231
6.3.6 Centrifugation 233
6.3.7 Sedimentation 233
6.3.8 Elutriation 234
6.3.9 Sectioning 235
6.3.10 Aeration 236
6.4 Behavioural or dynamic methods 236
6.4.1 Dry extractors 237
6.4.2 Wet extractors 243
6.5 Summary of the applicability of the methods 248
References 250
7 Relative Methods of Population Measurement and the Derivation of Absolute Estimates 259
7.1 Factors affecting the size of relative estimates 259
7.1.1 The ‘phase’ of the animal 260
7.1.2 The activity of the animal 261
7.1.3 Differences in the response between species, sexes and individuals 263
7.1.4 The efficiency of the trap or searching method 264
7.2 The uses of relative methods 266
7.2.1 Measures of the availability 266
7.2.2 Indices of absolute population 266
7.2.3 Estimates of absolute population 267
7.2.4 Removal trapping or collecting 268
7.2.5 Collecting 272
7.3 Relative methods: catch per unit effort 272
7.3.1 Observation by radar 272
7.3.2 Hydroacoustic methods 273
7.3.3 Fish counters 274
7.3.4 Electric fishing 274
7.3.5 Aural detection 275
7.3.6 Exposure by plough 276
7.3.7 Collecting with a net or similar device 276
7.3.8 Visual searching and pooting 279
7.4 Relative methods: trapping 280
7.4.1 Interception traps 281
7.4.2 Flight traps combining interception and attraction 290
7.4.3 Light and other visual traps 294
7.5 Traps that attract animals by some natural stimulus or a substitute 304
7.5.1 Shelter traps 304
7.5.2 Trap host plants 305
7.5.3 Baited traps 305
7.5.4 The use of vertebrate hosts or substitutes as bait for insects 308
7.6 Using Sound 314
References 314
8 Estimates of Species Richness and Population Size Based on Signs, Products and Effects 337
8.1 Arthropod products 337
8.1.1 Exuviae 337
8.1.2 Frass 338
8.2 Vertebrate products and effects 341
8.3 Effects due to an individual insect 342
8.4 General effects: plant damage 343
8.4.1 Criteria 344
8.5 Determining the relationship between damage and insect populations 347
References 348
9 Wildlife Population Estimates by Census and Distance Measuring Techniques 355
9.1 Census methods 356
9.2 Point and line survey methods 357
9.2.1 Indices of abundance using transects 357
9.2.2 Methods based on flushing 357
9.2.3 Line transect methods: the Fourier series estimator 360
9.2.4 Point transects 365
9.3 Distance sampling software in R 365
9.4 Spatial distribution and plotless density estimators 367
9.4.1 Closest individual or distance method 367
9.4.2 Nearest-neighbour methods 368
References 369
10 Observational and Experimental Methods for the Estimation of Natality, Mortality and Dispersal 373
10.1 Natality 373
10.1.1 Fertility 373
10.1.2 Numbers entering a stage 375
10.1.3 The birth-rate from mark and recapture data 382
10.2 Mortality 382
10.2.1 Total 382
10.2.2 The death-rate from mark and recapture data 383
10.2.3 Climatic factors 383
10.2.4 Biotic factors 383
10.2.5 Experimental assessment of natural enemies 390
10.3 Dispersal 396
10.3.1 Detecting and quantifying jump dispersal 397
10.3.2 Quantifying neighbourhood dispersal 397
10.4 The measurement and description of home range and territory 410
10.4.1 The minimum convex polygon area method for estimating home range 411
10.4.2 The kernel estimation method for home range 412
10.5 The rate of colonisation of a new habitat and artificial substrates 413
10.6 The direction of migration 413
References 413
11 The Construction, Description and Analysis of Age-specific Life-tables 429
11.1 Types of life-table and the budget 429
11.2 The construction of a budget 430
11.3 Analysis of stage-frequency data 431
11.3.1 Southwood’s graphical method 432
11.3.2 Richards & Waloff’s first method 432
11.3.3 Manly’s method 434
11.3.4 Ruesink’s method 435
11.3.5 Dempster’s method 435
11.3.6 Richards & Waloff’s Second Method 436
11.3.7 Kiritani, Nakasuji & Manly’s method 437
11.3.8 Kempton’s method 438
11.3.9 The Bellows and Birley Method 439
11.4 The description of budgets and life-tables 440
11.4.1 Survivorship curves 440
11.4.2 Stock–recruitment (Moran–Ricker) curves 440
11.4.3 The life-table and life expectancy 443
11.4.4 Life and fertility tables and the net reproductive rate 444
11.4.5 Population growth rates 446
11.4.6 The calculation of r 448
11.5 The analysis of life-table data 449
11.5.1 The comparison of mortality factors within a generation 449
11.5.2 Survival and life budget analysis 451
11.5.3 Sibley’s λ contribution analysis 458
References 459
12 Age-grouping, Time-specific Life-tables and Predictive Population Models 465
12.1 Age-grouping 465
12.2 Aging young by developmental stage 466
12.3 Aging by using structures 467
12.3.1 Annelids 467
12.3.2 Crustaceans 467
12.3.3 Insects 467
12.3.4 Molluscs 472
12.3.5 Fish 473
12.3.6 Lampreys 474
12.3.7 Reptiles and amphibians 474
12.3.8 Birds 475
12.3.9 Mammals 475
12.4 Time-specific life-tables and survival rates 476
12.4.1 Physiological time 478
12.4.2 Life-table parameters 479
12.4.3 Recruitment in the field 479
12.4.4 Empirical models 479
12.4.5 Intrinsic rate models and variable life-tables 480
12.4.6 Lewis–Leslie matrices and R packages 481
References 484
13 Species Richness, Diversity and Packing 495
13.1 Diversity 496
13.1.1 Description of α- and γ-diversity 497
13.1.2 Species richness 498
13.1.3 Models for the S:N relationship 505
13.1.4 Non-parametric indices of diversity 509
13.1.5 Which model or index? 512
13.1.6 Comparing communities – diversity ordering 513
13.1.7 Procedure to determine α-diversity 515
13.1.8 Determining β-diversity 517
13.2 Similarity and the comparison and classification of samples 520
13.2.1 Measures of complementarity 521
13.2.2 Similarity indices 521
13.2.3 Multivariate analysis 525
13.3 Species packing 530
13.3.1 Measurement of interspecific association 530
13.3.2 Measurement of resource utilisation 534
13.3.3 Niche size and competition coefficients 540
References 542
14 The Estimation of Productivity and the Construction of Energy Budgets 551
14.1 Estimation of standing crop 553
14.1.1 Measurement of biomass 553
14.2 Determination of energy density 554
14.3 Estimation of energy flow 555
14.4 The measurement of production 557
14.5 The measurement of feeding and assimilation 560
14.5.1 The quality of the food eaten 560
14.6 Feeding and assimilation rates 561
14.6.1 Radiotracer techniques 561
14.6.2 Gravimetric techniques 563
14.6.3 Indicator methods 564
14.6.4 Measurement of faecal output 565
14.7 The measurement of the energy loss due to respiration and metabolic process 565
14.7.1 Calorimetric 565
14.7.2 The exchange of respiratory gases 565
14.7.3 The respiratory rate 567
14.8 The energy budget, efficiencies and transfer coefficients 573
14.8.1 The energy budget of a population (or trophic level) 573
14.8.2 Energy transfer across trophic links 574
14.9 Identification of ecological pathways using stable isotopes 576
14.10 Assessment of energy and time costs of strategies 577
References 578
15 Studies at Large Spatial, Temporal and Numerical Scales and the Classification of Habitats 587
15.1 Remote sensing data from satellites 589
15.2 Remote sensing using piloted and unmanned aircraft 591
15.3 Long-term studies 592
15.3.1 Planning spatial and temporal sampling 593
15.3.2 The classification of time series 593
15.3.3 Detecting synchrony 603
15.3.4 Measuring temporal variability 603
15.3.5 Detecting break-points 604
15.4 Geographical information systems 607
15.5 Detection of density dependence in time series 608
15.5.1 Bulmer’s (1975) test 608
15.5.2 Pollard et al.’s (1987) randomisation test 609
15.5.3 Dennis and Taper’s (1994) bootstrap approach 611
15.5.4 Using a battery of approaches to detect density dependence 611
15.6 Citizen science projects 613
15.7 Ecosystem services 613
15.8 Habitat classification 614
15.8.1 Qualitative 614
15.8.2 Quantitative 616
References 617
Index 623
