waders and their estuarine food supplies - Vlaams Instituut voor de ...

More documents

Recommendations

Info

$Wetenschap - \Nijverheid- Handel$

PREDICTING SEASONAL AND ANNUAL FLUCTUATIONS IN THE LOCAL EXPLOITION OF DIFFERENT PREY Fig. I. Map .'I the Study area along ihe Frisian coast. The food supply was measured in the Nes area (7.3 ha: site N I. The birds were counted in the Nes area and in twenty counting areas around the Nes area, in total MX) ha ol tidal Hals. See als,. ihe map ol Fig, I" Methods Study area The study was performed between 1977 and 1986 on intertidal Hats along die Frisian coast in ihe Dutch Wadden Sea (Fig. I). The study area was intersected by long rows of poles, these being ihe remnants of brushwood groynes made in the sixties to enhance sedimentation. The former groynes delimited the study area, in total 396 ha. and divided it into 53 subareas. The eastern subareas were situated around mean sea level and the clay fraction (< 2 pm) in the substrate was 2-51. The more western subareas were al about 25 cm above mean sea level and the substrate was soft with a clay content of 10-20%. For a more extensive description of the study area, see Zwarts et al. (1992). The Oystercatchers feeding in this area used several roosts in the immediate surroundings, but at high tide the majority were found in the Paesenserpolder or on the island Engelsmanplaat. In total, about 20 000 Oystercalchers foraged on the tidal flats between the Frisian coast and the island Engelsmanplaat. We individually marked more than 3000 birds wilh the aid of 3 colour rings per bird, of which one with bar-codes. Most were captured with mist and cannon nets on the tidal roost in the Paesenserpolder. 236 Measuring the food supply Sampling Samples of the macrozoobenthos inhabiting the mudflats were taken along 27 transects in August during four years (1977-1980). More detailed data were collected in the eastern part of the study area, where 146 plots of 0.1 ha were pegged out around two observation towers. One to four samples of the benthic fauna (179 cm-) were taken nearly every month during seven years (1980-1986. and less frequently in 1978 and 1979) in the 73 sites around one of these towers, the Nes area, in earlier papers also indicated as site N. This series of measurements is used here. A comparison between the Nes data and the samples taken along the transects covering the entire area showed thai ihe Nes samples were representative for the area as a whole, especially for the most eastern part of ihe area (Zwarts 1988). The measurements in the Nes area started in December 1977 but. to extend the series of measurements, we will use for August 1977 the samples of two of the 27 transects that crossed the Nes area. Condition, growth and mortality The laboraiotv procedures used to determine the biomass of the benthic animals have been described in Zwarts (1991). The growth of the bivalves could sometimes be ealeu-
PREDICTING SEASONAL AND ANNUAL FLUCTUATIONS IN THE LOCAL EXPLOITION OF DIFFERENT PREY kited directly from the length frequencv distribution in the monthly samples, but this was usually impossible because two or even more year classes occurred together. The year classes v. ere identified using the Bhattaeharya method for separating cohorts (MPA module of the Compleat ELEFAN software package, version 1.0: Gayanilo et nl. 1988). This technique made it possible lo estimate the monthly mortality and growth in length for ihe separate cohorts. Estimating prey production Using the relationships between size and llesh weight, determined for all sampling data, we also calculated for each cohort the fluctuation in the biomass of ihe average indiv idual and for the cohort population as a whole. This allowed us to calculate the production per species, separately for the age classes. The production can be estimated by adding either the growth increments or the weight losses caused by size-dependent mortality (Crisp 1984). We estimated the production by calculation of the monthly weight loss since this made it possible to indicate to what degree the elimination of biomass due to prey moriality had been determined by the predation pressure of the Oystercatchers. The elimination of the prey biomass is determined by the product of two terms: the mean weight of the pre) averaged over two consecutive sampling dale's and the rfayrr-flSf in the pre" denshj between both sample dates. Both, prey weight and the numbers disappeared, were calculated separately per cohort and recalculated per month if the intervening period was longer than a month. The eliminated biomass per cohort was summed to arrive at the total monthly elimination of biomass per prey species. Since Oystercatchers do not take Cockles < 10 mm. Macoma < 11 mm and the Soft-shell Clam Mya arenaria < 17 mm (Zwarts ei al. 1996a). the elimination of biomass belonging to the size classes ignored by Oystercatchers will be given separately, as well as for prey living out nl reach of ihe bill. i.e. Mya living more llian 6 cm below the surface (Zwarts & Wanink 1984). Burying depth The burying depth of the bivalves was measured once or twice a month during seven years (1980-1986). The methods have been described by Zwarts & Wanink (1989). The combination of biomass samples and the depth measuremenis was used to describe the annual fluctuation in the biomass acluallv View Irom the sea wall over Ihe study area. The observation tower was built in the middle of the .Nes area at the cross of two 237
Page 1 and 2:
WADERS AND THEIR ESTUARINE FOOD SUP
Page 3 and 4:
plia^ohi. WADERS AND THEIR ESTUARI
Page 5 and 6:
Waders and their estuarine food sup
Page 7 and 8:
WADERS AND THEIR ESTUARINE FOOD SUP
Page 10 and 11:
figuren: Dick Visser omslagfoto: Ja
Page 12 and 13:
15 Versatility of male curlews (Num
Page 15 and 16:
That science is making progress, ma
Page 17 and 18:
INTRODUCTION The remnants of brushw
Page 19 and 20:
When hcnlhic bivalves extend llien
Page 21 and 22:
the burrow depths and on the fracti
Page 23 and 24:
INTRODUCTION Ms,i invest 409 of (he
Page 25 and 26:
able lo sw itch 10 oiher prey which
Page 27 and 28:
Long-term, broadly-based research c
Page 29 and 30:
Chapter 1 SEASONAL VARIATION IN BOD
Page 31 and 32:
SEASONAL VARIATION IN BODY WEIGHT O
Page 33 and 34:
Tahle 1 Weight loss ('•- ± SE) m
Page 35 and 36:
Page 37 and 38:
Page 39 and 40:
i 60 50 40 30 20 10 0 • INFESTED
Page 41 and 42:
240- SEASONAL VARIATION IN BODY WEI
Page 43 and 44:
20 10 0 -10 -20h 2 3 4 5 6 7 8 9 se
Page 45 and 46:
a E 400 - S. plana 35 mm 320 240 16
Page 47 and 48:
Page 49 and 50:
Chapter 2 HOW THE FOOD SUPPLY HARVE
Page 51 and 52:
FOOD SUPPLY HARVESTABLE BY WADERS H
Page 53 and 54:
Methods The study sites were situat
Page 55 and 56:
less than that of bivalves, partly
Page 57 and 58:
I Fig. 3). Peak condition was reach
Page 59 and 60:
total biomass since most of those t
Page 61 and 62:
on the basis of the preceding and t
Page 63 and 64:
However, for obvious reasons, we ma
Page 65 and 66:
prey. A decrease in the prey densit
Page 67 and 68:
Tahle 2. The intake rate ol < lyste
Page 69 and 70:
* ' % -. . ; a X - . - * • ^ •
Page 71 and 72:
(Zwarts & Wanink 1989). In quiet we
Page 73 and 74:
general law relating handling time
Page 75 and 76:
nearly twice as much time (0.79 s).
Page 77 and 78:
4 5 6 7 8 9 size class of Corophium
Page 79 and 80:
and annually. Second, the lower siz
Page 81 and 82:
Prey switching Waders feeding on ti
Page 83 and 84:
autumn and spring, and the contrary
Page 85 and 86:
where they switch to surface-living
Page 87 and 88:
Chapter 3 BURYING DEPTH OF THE BENT
Page 89 and 90:
DEPTH AND SIPHON CROPPING IN SCROBI
Page 91 and 92:
I DEPTH AND SIPHON CROPPING IN SCRO
Page 93 and 94:
DEPTH AND SIPHON CROPPING IN SCROBI
Page 95 and 96:
Table 3. Three-Way analysis Of vari
Page 97:
Chapter 4 SIPHON SIZE AND BURYING D
Page 100 and 101:
15 20 size (mm) Fig. 3. Cerasioderm
Page 102 and 103:
10 size (mm) SIPHON SIZE AND DEPTH
Page 104 and 105:
o, 7 01 5 | * CL 5- SIPHON SIZE AND
Page 106 and 107:
4 Q) Si •a a. 16- 20- A predator:
Page 108 and 109:
prey risk for an animal al a depth
Page 110 and 111:
Table 6. Size al which there is a m
Page 112 and 113:
* ' 1 /-/ "».-^*«C
Page 114 and 115:
face and so expose themselves to a
Page 116 and 117:
surface in die containers was varie
Page 118 and 119:
50 OOF 310.00 I 5.00 ^ 1.00 3=" 0.5
Page 120 and 121:
Tahle 1. Macoma and Scrobicularia.
Page 122 and 123:
siphon would not have to reach the
Page 124 and 125:
known siphon weight and burying dep
Page 127 and 128:
ACCESSIBLE PREY ARE OFTEN IN POOR C
Page 129 and 130:
1 2 3 4 burying depth (cm) Kin. I.
Page 131 and 132:
I—1 • • : : : : ! - ! -|-r 0
Page 133 and 134:
Chapter 7 DOES AN OPTIMALLY FORAGIN
Page 135 and 136:
OPTIMAL FORAGING AND THE FUNCTIONAL
Page 137 and 138:
100 200 300 prey density (n-m-2) Fi
Page 139 and 140:
Table I. Results of five one way an
Page 141 and 142:
Table 2. Results of eight iwo-wav a
Page 143 and 144:
Page 145 and 146:
prey density II III Fig. 12. Freque
Page 147 and 148:
Page 149:
PREY SIZE SELECTION AND INTAKE RATE
Page 152 and 153:
Predicted 'passive size selection'
Page 154 and 155:
lig. 2. Larger Mussels are less ava
Page 156 and 157:
Fig. 5. Scrobicularia plana. Size c
Page 158 and 159:
and maiiv are too thick-shelled to
Page 160 and 161:
The measurements of handling time i
Page 162 and 163:
1 2 3 4 5 6 7 probing depth (cm PRE
Page 164 and 165:
valves may vary by a factor of two
Page 166 and 167:
optimal foraging model, the minimum
Page 168 and 169:
their gut is full? Do they reduce t
Page 171 and 172:
PREY PROFITABILITY AND INTAKE RATE
Page 173 and 174:
catchers to take only certain prey
Page 175 and 176:
licult error of estimate arose if p
Page 177 and 178:
Counts of feeding and non-feeding b
Page 179 and 180:
PREY PROFITABILITY AND INTAKE RATE
Page 181 and 182: 20 30 40 50 shell length (mm) PREY
Page 183 and 184: PREY PROFITABILITY AND INTAKE RATE
Page 185 and 186: Nereis Arenicola tipuia earthwo'm M
Page 187 and 188: take rate. We might therefore expec
Page 189 and 190: prey species, using parallel slopes
Page 191 and 192: 5.0 4.0 1-3.0 a & • % * • * •
Page 193 and 194: time during the feeding period. As
Page 195 and 196: winter. Similarly, handling time in
Page 197 and 198: 5.0 - f-4.0 S 3.0 in I 20 a 2 a | 1
Page 199 and 200: situation arrived in the western pa
Page 201 and 202: PREY PROFITABILITY AND INTAKE RATE
Page 203 and 204: • ' • 14 PREY PROFITABILITY AND
Page 205 and 206: Notes lo appendix: PREY PROFITABILI
Page 207: Chapter 10 WHY OYSTERCATCHERS HAEMA
Page 210 and 211: 2 4 6 8 10 time on feeding area (h)
Page 212 and 213: 80- 60- o 80 60- 40- 20- INTAKE RAT
Page 214 and 215: est of bod\ behind: an estimated 22
Page 216 and 217: INTAKE RATE AND PROCESSING RATE IN
Page 218 and 219: Wanink 1993). The energy content of
Page 220 and 221: (7) Age All studies dealt with adul
Page 222 and 223: irds over long periods. As an examp
Page 224 and 225: Discussion There is no difference i
Page 226 and 227: comparison between the weight of ih
Page 228 and 229: . 1', L ! •
Page 230 and 231: Introduction PREDICTING SEASONAL AN
Page 234 and 235: PREDICTING SEASONAL AND ANNUAL FLUC
Page 240 and 241: 1000 r 1 II" 1 - r^Jan'80 PREDICTIN
Page 252 and 253: IOO BO 60 40 20 0 PREDICTING SEASON
Page 265 and 266: WHY KNOT TAKE MEDIUM-SIZED MACOMA W
Page 267 and 268: in Zwarts & Esselink 1989). The len
Page 269 and 270: shell length (mm) FIR. 3. Peringia.
Page 271 and 272: fused, specimens larger than this w
Page 273 and 274: 50 100- s s I — f = S. plana, n=2
Page 275 and 276: area is twice as large as the touch
Page 277 and 278: 10 15 lower size threshold (mm) Fig
Page 279 and 280: lime (Hughes 1979). This would furt
Page 281 and 282: WHY KNOT TAKE MEDIUM-SIZED MACOMA T
Page 283 and 284:
Chapter 13 ANNUAL AND SEASONAL VARI
Page 285 and 286:
VARIATION IN FOOD SUPPLY OF KNOT AN
Page 287 and 288:
Two sites (N and M in Fig. 2) were
Page 289 and 290:
20 30 VARIATION IN FOOD SUPPLY OF K
Page 291 and 292:
20 l i r'l ' i •o j^y^T n ^ ' " ^
Page 293 and 294:
July ' Aug ' Sept Fig. 9. Proportio
Page 295 and 296:
available. There must, however, be
Page 297 and 298:
Chapter 14 SEASONAL TREND IN BURROW
Page 299 and 300:
BURROWING AND FEEDING IN NEREIS SEA
Page 301 and 302:
Table 2. Results of a 2-way analysi
Page 303 and 304:
June 1981 June 1982 is * N.MUD •
Page 305 and 306:
8 10 I 12 JZ 5. -g 1*» •e o i 16
Page 307 and 308:
6 - n=!080 5 • g 4 CP > i 3 CO CD
Page 309 and 310:
1985) and Curlew (/wails ,v, Lsseli
Page 311:
Chapter 15 VERSATILITY OF MALE CURL
Page 314 and 315:
Smid! 1951. Muus 1967, Wolff 1973,
Page 316 and 317:
0.6 0.8 1.0 1.2 1.4 1.6 jaw lengih
Page 318 and 319:
Na oi both Nrieck Fig. 7. Numenius
Page 320 and 321:
6 8 10 12 14 16 18 worm length (cm)
Page 322 and 323:
too (Fig. 11 A). Indeed, the averag
Page 324 and 325:
VERSATILITY OF CURLEWS FEEDING ON N
Page 326 and 327:
total time spent al the surface. Th
Page 328 and 329:
2 4 . • 2.2 • 30 .-.2.0 to Q. 1
Page 331 and 332:
PREY DEPLETION BY OYSTERCATCHER AND
Page 333 and 334:
the rule: the observed lower limit
Page 335 and 336:
to locate the clams after they had
Page 337 and 338:
Curlew, by bury ing some hundreds o
Page 339 and 340:
PREY DEPLETION BY OYSTERCATCHER AND
Page 341 and 342:
SAMENVATTING 345
Page 343 and 344:
Voorgeschiedenis In 1960 verscheen
Page 345 and 346:
maken. Als gebied A een grotere voe
Page 347 and 348:
omdat ze nog vrijwel niets wegen. l
Page 349 and 350:
kunnen opzuigen. of diep leven en z
Page 351 and 352:
van de uitgerekte sifo over het wad
Page 353 and 354:
het toen niet gemakkelijk hebben ge
Page 355 and 356:
doende nonnetjes van 10 tot 15 mm l
Page 357 and 358:
aangevoerde voedsel en de wulp past
Page 359 and 360:
zijn dan twee jaar en wulpen geen s
Page 361 and 362:
was dan ook dank/ij de inzet van al
Page 363 and 364:
REFERENCES 367
Page 365 and 366:
Allen RL. 1983. Feeding behaviour o
Page 367 and 368:
BreyT. 1989. Der Einfluss physikali
Page 369 and 370:
latie tol chironoiiiiilen en de wai
Page 371 and 372:
huch der Vogel Milteleuropas, Band
Page 373 and 374:
llolmann II. & H. Huerschelmann 196
Page 375 and 376:
l-cndrem D.W. 1984. Flocking, feedi
Page 377 and 378:
Pekkarinen M. 1984. Regeneration of
Page 379 and 380:
lion of Mussels Mytilus edulis hy O
Page 381 and 382:
lulal Dal esiuanes: a comparison of
show all

waders and their estuarine food supplies - Vlaams Instituut voor de ...

Create successful ePaper yourself

Delete template?

Save as template?