Patch departure mechanisms and optimal host exploitation in an insect parasitoid

1. Patch-leaving decisions are of utmost importance in determining parasitoid foraging success. Parasitoids are known to use both marks left by hosts (chemical or otherwise) and ovipositions to assess host availability and to decide when to leave a host patch. 2. Previous studies have shown that, depending on the species, ovipositions either increase (an incremental mechanism) or decrease (a decremental mechanism) the patch residence times of parasitoids. Reports in the literature conflict on which mechanism is used by Venturia canescens, a parasitoid of pyralid moth larvae. 3. We hypothesize that, as a consequence of saturation in the capacity of the parasitoid to discriminate between host densities at high host numbers, V. canescens uses a decremental mechanism at low host numbers and an incremental one at high host numbers. We call this a ‘switching mechanism’. 4. Our experiments show that even if discrimination capacity saturates, V. canescens uses a decremental mechanism over a wide range of host densities. 5. The distribution of hosts in different fruits species under field conditions suggests a switching mechanism would not evolve in natural situations. 6. A model of patch departure in V. canescens is constructed and tested using an independent set of experiments. The model suggests that the patch leaving mechanism in V. canescens is a stochastic decremental one. As might be expected from Weber's Law, the initial leaving tendency is a convex decreasing function of kairomone concentration. The leaving tendency increases exponentially with the time spent in the patch without ovipositing. Ovipositions cause a sudden increase in leaving tendency. 7. Simulations suggest that a decremental mechanism would be out-competed by either one indifferent to ovipositions or an incremental one, only when travel times are much larger than those that are likely to occur in the field.

[1]  Richard F. Green,et al.  Stopping Rules for Optimal Foragers , 1984, The American Naturalist.

[2]  J. Waage,et al.  Foraging for patchily-distributed hosts by the parasitoid, Nemeritis canescens , 1979 .

[3]  P. Haccou,et al.  INFORMATION PROCESSING BY FORAGERS: EFFECTS OF INTRA-PATCH EXPERIENCE ON THE LEAVING TENDENCY OF LEPTOPILINA HETEROTOMA , 1991 .

[4]  C. Bernstein,et al.  Patch-marking and optimal search patterns in the parasitoid Venturia canescens , 1996 .

[5]  D. Gordon,et al.  Factors determining the functional response of the parasitoid Venturia canescens , 1989 .

[6]  M. Strand,et al.  BEHAVIORAL RESPONSE OF THE PARASITOID CARDIOCHILES NIGRICEPS TO A KAIROMONE , 1982 .

[7]  J. V. Lenteren,et al.  The leaving tendency of the parasitoid Encarsia formosa foraging for whitefly on tomato leaflets. , 1993 .

[8]  J. M. Nelson,et al.  Flexible patch time allocation by the leafminer parasitoid, Opius dimidiatus , 1995 .

[9]  T. Ahmad The Influence of Ecological Factors on the Mediterranean Flour Moth, Ephestia kuhniella and its Parasite, Nemeritis canescens , 1936 .

[10]  Y. Iwasa,et al.  Prey Distribution as a Factor Determining the Choice of Optimal Foraging Strategy , 1981, The American Naturalist.

[11]  L. Shaltiel,et al.  The use of kairomones for foraging decisions by an aphid parasitoid in small host aggregations , 1998 .

[12]  Jérôme Casas,et al.  A probabilistic model for the functional response of a parasitoid at the behavioural time scale , 1993 .

[13]  Per Kragh Andersen,et al.  Testing Goodness of Fit of Cox's Regression and Life Model , 1982 .

[14]  P. Haccou,et al.  Effects of intra-patch experiences on patch time, search time and searching efficiency of the parasitoid Leptopilina clavipes (Hartig). , 1993 .

[15]  G. Salt The hosts of Nemeritis canescens a problem in the host specificity of insect parasitoids , 1976 .

[16]  J. C. Lenteren,et al.  Behavioural Aspects of the Functional Responses of a Parasite (Pseudeucoila Bochei Weld) To Its Host (Drosophila Melanogaster) , 1977 .

[17]  E. Wajnberg,et al.  Genetic variation in patch time allocation in a parasitic wasp , 1999 .

[18]  J. Kalbfleisch,et al.  The Statistical Analysis of Failure Time Data , 1980 .

[19]  D. Rogers THE ICHNEUMON WASP VENTURIA CANESCENS: OVIPOSITION AND AVOIDANCE OF SUPERPARASITISM , 1972 .

[20]  P. Haccou,et al.  The Influence of Larval Dispersal in the Cinnabar Moth (Tyria jacobaeae) on Predation by the Red Wood Ant (Formica polyctena): An Analysis Based on the Proportional Hazards Model , 1985 .

[21]  Richard F. Green,et al.  Stochastic Models of Optimal Foraging , 1987 .

[22]  C. Cloutier Searching Behavior of the Aphid Parasitoid Aphidius nigripes (Hymenoptera: Aphidiidae) Foraging on Potato Plants , 1990 .

[23]  A. Kacelnik,et al.  A count-down mechanism for host search in the parasitoid Venturia canescens , 1995 .

[24]  James N. McNair,et al.  Optimal Giving-Up Times and the Marginal Value Theorem , 1982, The American Naturalist.

[25]  C. Bernstein,et al.  State dependent superparasitism in a solitary parasitoid: egg load and survival , 1997 .

[26]  F. Galis,et al.  PATCH TIME ALLOCATION AND PARASITIZATION EFFICIENCY OF ASOBARA TABIDA, A LARVAL PARASITOID OF DROSOPHILA , 1983 .

[27]  Lia Hemerik,et al.  Patch exploitation by the parasitoids Cotesia rubecula and Cotesia glomerata in multi‐patch environments with different host distributions , 1998 .

[28]  G. Rowe,et al.  Adaptive patterns in the avoidance of superparasitism by solitary parasitic wasps , 1987 .

[29]  W. Thorpe,et al.  Olfactory Conditioning in a Parasitic Insect and Its Relation to the Problem of Host Selection , 1937 .

[30]  J. McNamara Optimal patch use in a stochastic environment , 1982 .

[31]  D. Cox Regression Models and Life-Tables , 1972 .

[32]  S. Corbet Mandibular Gland Secretion of Larvae of the Flour Moth, Anagasta kuehniella, contains an Epideictic Pheromone and elicits Oviposition Movements in a Hymenopteran Parasite , 1971, Nature.

[33]  E. Charnov Optimal foraging, the marginal value theorem. , 1976, Theoretical population biology.