Superparasitism as an ESS: To reject or not to reject, that is the question.
Nelly van der Hoeven1* and Lia Hemerik2.
*: Present address: ECOSTAT, Vondellaan 23, 2332 AA Leiden, The Netherlands
- Institute of Theoretical Biology, University of Leiden, P.O. Box 9516, 2300 RA Leiden, The Netherlands
- Department of Population Biology, University of Leiden, P.O. Box 9516, 2300 RA Leiden, The Netherlands
* Corresponding Author:
Nelly van der Hoeven,
2332 AA Leiden,
tel.: (+)31-(0)71-5315011; fax: (+)31-(0)842-116988;
A stochastic model is formulated to determine the optimal strategy for a solitary parasitoid which has discovered an already parasitized host. The model assumes that the parasitoid can count both the number of eggs already present in a host and the number of conspecifics searching in the same patch. The survival probability of an egg is assumed to depend on the total number of eggs in a host. The decision to (super)parasitize depends both on the degree to which the discovered host already is parasitized and on the number of conspecific females searching in the same patch. We consider both the case that egg laying does not involve any costs for the parasitoid and the case that it involves some marginal costs. Uniform behaviour of all the conspecific parasitoids in a patch, i.e. laying one additional egg in all encountered larvae containing a particular number of eggs, appears to be a pure evolutionary stable strategy (ESS). If either the probability that a parasitoid emerges from a host decreases with an increasing degree of parasitism, at least from a particular number of eggs onwards, or if parasitism involves marginal costs, the maximum number of eggs for which it is still profitable to superparasitize a host once more is limited. This number increases with the number of conspecifics searching in the patch. Large marginal costs (i.e. the expected gain of not parasitizing now) decrease the profit of superparasitism. For newly emerged parasitoids the rejection of an already parasitized host is not advantageous as long as the marginal costs of parasitism are small, because the host can never contain an egg of its own.
Journal of Theoretical Biology 146: 467-482, 1990
Selection from publications of Nelly van der
Is it safe to pool the Blank Control Data with the Solvent Control data? N. van der Hoeven,
Ecotoxicol. Environm. Safety, 73:1480-1483, 2010
Multi-criteria decision analysis of test endpoints for detecting the effects of endocrine active substances in fish full life cycle tests.
M. Crane, M. Gross, P. Matthiessen, G.T. Ankley, S. Axford, P. Bjerregaard, R. Brown, P. Chapman, M. Dorgeloh, M. Galay-Burgos, J. Green, C. Hazlerigg, J. Janssen, K. Lorenzen, J. Parrott, H. Rufli, C. Schäfers, M. Seki, H.C. Stolzenberg, N. van der Hoeven, D. Vethaak, IJ. Winfield, S. Zok & J. Wheeler
Integr Environ Assess Manag., 6: 378-389, 2010
Exposure analysis of bisphenol A in surface water systems in North America and Europe. (PMID:19746705)
G.M. Klecka, C.A. Staples, K.E. Clark, N. van der Hoeven, D.E. Thomas & S.G. Hentges
Environmental Science & Technology, 43: 6145-6150, 2009
The Minimum Significant Difference at the NOEC calculated with a non-parametric test.
Hoeven, N. van der,
In: Proceedings of the 30th Anniversary Meeting of the Netherlands Society of Toxicology, june 2009. p. 122
Calculation of the Minimum Significant Difference at the NOEC using a non-parametric test.
Hoeven, N. van der,
Ecotoxicol. Environm. Safety, 70: 61-66, 2008
Does bisphenol a induce superfeminization in Marisa cornuarietis?
Part I: Intra- and inter-laboratory variability in test endpoints. Forbes, V.E., H. Selck, A. Palmqvist, J. Aufderheide, R. Warbritton, N. Pounds, R. Thompson, N. van der Hoeven & N. Caspers
Ecotoxicol. Environm. Safety, 66: 309-318, 2007
Does bisphenol A induce superfeminization in Marisa cornuarietis?
Part II: Toxicity test results and requirements for statistical
power analyses. Forbes, V.E., J. Aufderheide, R. Warbritton, N. van der Hoeven & N. Caspers
Ecotoxicol. Environm. Safety, 66: 319-325, 2007
Statistical issues in fish life-cycle tests with many endpoints.
Hoeven, N. van der & D.R. Dietrich,
Abstract and poster for SETAC Europe, May 2005, Lille
The probability to select the correct model using likelihood-ratio based criteria in choosing between two nested models of which the more extended one is true.
Hoeven, N. van der,
Journal of Statistical Planning and Inference, 135: 477-486, 2005
Effects of bisphenol A on adult fathead minnow (P. promelas) gonadal histology: a 42-day exposure study.
Dietrich, D.R., J. Wolf, A.R. Brown, J.E. Caunter, N. van der Hoeven & U. Friederich,
Abstract and poster for the Cluster workshop on Ecological relevance of chemically induced endocrine disruption in wildlife. University of Exeter, july 2004.
The Netherlands working group on Statistics and Ecotoxicology: Statistics and Models for Risk Assessment.
Hoeven, N. van der,
In: Proceedings of the Jubilee Annual Meeting of the Netherlands Society of Toxicology, june 2004. p. 114
Current issues in statistics and models for Ecotoxicological Risk Assessment.
Hoeven, N. van der,
Acta Biotheoretica 52: 201-217, 2004
Balancing statistics and ecology: on the lumping of experimental data for model selection.
Hoeven, N. van der, L. Hemerik & P.A. Jansen. In: T.A.C. Reydon & L. Hemerik (Eds): Current themes in Theoretical Biology: A Dutch Perspective. pp 233-265. Springer, Dordrecht, The Netherlands, 2004.
Using marine bioassays to classify the toxicity of Dutch harbour sediments. Stronkhorst, J., C. Schipper, J. Brils, M. Dubbeldam, J. Postma & N. van der Hoeven
Environmental Toxicology and Chemistry, 22: 1535-1547, 2003
What can egg distributions of solitary parasitoids tell us about the information the parasitoid has and uses for its oviposition decisions? Hemerik, L.,N. van der Hoeven &
J. J.M. van Alphen, Acta Biotheoretica, 50: 167-188, 2002
Statistical tests and power analysis for three in-vivo bioassays
to determine the quality of marine sediments. Hoeven, N. van der, B. J. Kater &
J. F. Pieters, Environmetrics 13: 281-293, 2002
Significance tables for the exact variance test for the Poisson
distribution with alternative underdispersion. N. van der Hoeven & L Hemerik,
Environmental and Ecological Statistics 9: 201-213, 2002
Estimating the 5-percentile of the species sensitivity
distribution without any assumptions about the distribution. N. van der Hoeven,
Ecotoxicology 10: 25-34, 2001
Power analysis for the NOEC: What is the probability to detect
small toxic effects on three different species using the appropriate standardized test
protocols? N. van der Hoeven, Ecotoxicology 7: 355-361, 1998
The acute toxicity of selected alkylphenols on young and adult
Daphnia magna. A.A.M. Gerritsen, N. van der Hoeven & A. Pielaat,
Ecotoxicol. Environm. Safety 39: 227-232. 1998
The ecotoxicity and the biodegradability of lactic acid, alkyl
lactate esters and lactic acid salts. C.T. Bowmer, R.N. Hooftman, A.O. Hanstveit,
P.W.M. Venderbosch & N. van der Hoeven, Chemosphere 37:
How to measure no effect? Part I: Towards a new measure of
chronic toxicity in ecotoxicology. Introduction and workshop results. N. van der Hoeven,
F. Noppert & A. Leopold, Environmetrics 8: 241-248, 1997
How to measure no effect? Part III: Statistical aspects of
NOEC, ECx and NEC estimates. N. van der Hoeven, Environmetrics 8: 255-261, 1997
The effect of chlorpyrifos on individuals of Daphnia
pulex in laboratory and field. N. van der Hoeven & A.A.M. Gerritsen,
Environm. Toxicol. Chem. 16: 2438-2447, 1997
A model based on soil structural aspects describing the fate of
genetically modified bacteria in soil. N. van der Hoeven & J.D. van Elsas,
Ecological Modelling 89: 161-173, 1996
Competition between cohorts of juvenile Daphnia
magna. E.L. Enserink, N. van der Hoeven, M. Smith, M. van der Klis & M.A.
van der Gaag, Archiv für Hydrobiologie 136: 433-454, 1996
Reliability of quantitative toxicity test results: from
experimental control to data processing. Enserink, E.L. & N. van der Hoeven,
The Science of the Total Environment, suppl. 1993, Proceedings of the Second
European Conference on Ecotoxicology, eds. W. Slooff & H. de Kruijf, p. 699-704,
LC50 estimates and their confidence intervals. The case that
only one test concentration has partial effect. N. van der Hoeven, Water
Research 25: 401-408, 1991
Effects of toxicants on individuals and populations of Daphnia,
a simulation study. N. van der Hoeven, Comparative Biochemistry and
Physiology 100C: 283-286, 1991
Effect of 3,4-dichloroaniline and metavanadate on Daphnia
populations. N. van der Hoeven, Ecotoxicology and Environmental Safety
20: 53-70, 1990
Salmonella test: Relation between mutagenicity and number of
revertant colonies. N. van der Hoeven, S.A.L.M. Kooijman & W.K. de Raat,
Mutation Research 234: 289-302, 1990
Superparasitism as an ESS: to reject or not to reject, that is
the question. N. van der Hoeven & L. Hemerik, J. of Theoretical Biology
146: 467-482, 1990
Population consequences of a physiological model for individual
development S.A.L.M. Kooijman, N. van der Hoeven & D.C. van der Werf,
Functional Ecology 3: 325-336, 1989
Oscillations in Daphnia populations. N. van der
Hoeven, A.M. de Roos & S.A.L.M. Kooijman, Econieuws 2, 7-8,
The population dynamics of Daphnia at constant food supply: a
review, re-evaluation and analysis of experimental series from the literature. N. van
der Hoeven, Netherlands Journal of Zoology 39: 126-155, 1989
Random elements in a population model based on individual
development. N. van der Hoeven, in: Ecodynamics, Proc. Int. Workshop at Jülich,
FRG, 19-20 Oct. 1987. Eds. W. Wolff, C.J. Soeder & F.R. Drepper, pp 333-342,
Research Notes in Physics, Springer Verlag., 1988