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Intake rates and the functional response in shorebirds (Charadriiformes) eating macro-invertebrates
- John D. Goss-Custard, Andrew D. West, Michael G. Yates, Richard W. G. Caldow, Richard A. Stillman, Louise Bardsley, Juan Castilla, Macarena Castro, Volker Dierschke, Sarah. E. A. Le. V. dit Durell, Goetz Eichhorn, Bruno J. Ens, Klaus-Michael Exo, P. U. Udayangani-Fernando, Peter N. Ferns, Philip A. R. Hockey, Jennifer A. Gill, Ian Johnstone, Bozena Kalejta-Summers, Jose A. Masero, Francisco Moreira, Rajarathina Velu Nagarajan, Ian P. F. Owens, Cristian Pacheco, Alejandro Perez-Hurtado, Danny Rogers, Gregor Scheiffarth, Humphrey Sitters, William J. Sutherland, Patrick Triplet, Dave H. Worrall1, Yuri Zharikov, Leo Zwarts, Richard A. Pettifor
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- Journal:
- Biological Reviews / Volume 81 / Issue 4 / November 2006
- Published online by Cambridge University Press:
- 24 July 2006, pp. 501-529
- Print publication:
- November 2006
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As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type II (‘disc equation’) formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching.
A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (<150/m−2). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote.
A multivariate analysis of 468 ‘spot’ estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81% of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3%), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6%, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93% of cases.
We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested.
As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shorebird ecology and behaviour to estimate intake rate without the need for conventional time-consuming field studies, including species for which it has not yet proved possible to measure intake rate in the field.
Motility of the reticulum and rumen of sheep given juice-extracted pasture
- L. M. McLeay, D. C. Kokich, H-U. Hockey, T. E. Trigg
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- Journal:
- British Journal of Nutrition / Volume 47 / Issue 1 / January 1982
- Published online by Cambridge University Press:
- 24 July 2007, pp. 79-85
- Print publication:
- January 1982
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1. Sheep were fed on different diets of juice-extracted herbage to determine what effect juice-extraction had on reticulo-rumen motility.
2. The frequency of A and B sequences of contraction of the reticulo-rumen were recorded during eating, rumination and inactivity for continuous periods of 24–72 h by using integrated electromyograms obtained from electrodes implanted in the musculature of the reticulum and cranial dorsal rumen.
3. Animals were fed on herbage in which approximately 200 g/kg dry matter had been removed in juice extracted from ryegrass (Lolium perenne), white clover (Trifolium repens), mixed ryegrass–white clover and lucerne (Medicago saliva).
4. Over all the frequency of A sequences of contraction did not differ in animals fed on pressed herbage or the unpressed material from which it was derived, although it was slower during rumination on some of the pressed material. In contrast, the frequency of B sequences was higher on the pressed material. The frequencies of contraction of A and B sequences in animals fed on pressed herbage was related to the activity of the animals in the order eating > rumination > inactivity.
5. Changes in reticulo-rumen motility due to juice extraction were small and the frequencies of A and B sequences of contraction in sheep fed on pressed herbage were in the range encountered in ruminants consuming more conventional foods.