Characteristic Permitting Coexistence Among Parasitoids of a Sawafly in Quebec Author(S): Peter W
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Characteristic Permitting Coexistence Among Parasitoids of a Sawafly in Quebec Author(s): Peter W. Price Source: Ecology, Vol. 51, No. 3 (May, 1970), pp. 445-454 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1935379 . Accessed: 29/08/2011 15:38 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology. http://www.jstor.org CHARACTERISTICS PERMITTING COEXISTENCE AMONG PARASITOIDS OF A SAWFLY IN QUEBEC PETER W. PRICE Forest Research Laboratory, Department of Fisheries and Forestry, Quebec 10, Quebec and Department of Entomology, Cornell University, Ithaca, N. Y. 14850 Abstract.-Six parasitoid species (Hymenoptera: 5 Ichneumonidae, 1 Eulophidae) coexist on cocoon populations of the Swaine jack pine sawfly, Neodiprion swainei Midd. in Quebec. Their distribution between plant communities was related to host availability except for Gelis urbanus (Brues) which has alternative hosts. Pleolophus basizonus (Grav.) was dominant at high host densities, tending to displace Pleolophus indistinctus (Prov.) which was dominant at low host densities. Mastrus aciculatus (Prov.) occupied dry open sites at high host densitiies, where other species were less numerous. The highest species diversity of cocoon parasitoids occurred at the ecotone around jack pine stands and at moderate host densities in increasing host populations. Furthermore, parasitoids attacking larvae were well represented in moderately dense host populations. As the host density increased, P. basizonus became so dominant that it reduced species diversity, particularly among the parasitoids attacking the host larvae. Species diversity can be predicted from the proportion of P. basizonus present. This parasitoid acts as an organizer species within the parasitoid complex because of its superior competitive ability. Parasitoids of larvae have a conservative strategy for host exploitation that appears adapted for utilizing low host populations. In contrast, the parasitoids attacking cocoons gain dominance only when host-finding is easy in high host populations. Factors which enable several species of parasi- fore all those species that attack the host in the toids to coexist in the same host population are cocoon stage, the cocoon parasitoid guild, were not well understood. Heatwole and Davis (1965) studied in detail. Through an understanding of have found that small differences in ovipositor the organization within this guild I have been able length enabled three species of Megarrhyssa (Ich- to assess some of its effects on the parasitoids that neumonidae) to exploit different segments of the attack larvae, the larval parasitoid guild. same host population. However, the general lack of knowledge concerning competitive interactions GUILD MEMBERSHIP AND LIFE HISTORIES between parasitoids led Turnbull and Chant Two members of the cocoon parasitoid guild (1961) to caution against the multiple introduc- were introduced from Europe: Pleolophus basi- order tion of parasitoids for biological control. In zonus (Grav.) (Ichneumonidae) during 1933- interactions, to gain insight on these competitive 1940 and Dahlbominus fuscipennis (Zett.) (Eu- pine I studied the parasitoids of the Swaine jack lophidae) during 1934-1948 (McGugan and Cop- dur- sawfly, Neodiprion swainei Midd. in Quebec pel 1962). These species were liberated against ing 1967 and 1968. the spruce sawfly, Diprion hercyniae (Htg.), but Differences in habitat selection provide one ob- have since become established in my study area vious means for species to partition a resource on N. swainei (McLeod and Martineau 1967). held in common. In such a case, competition will The remainder of the cocoon parasitoid guild occur only in those habitats that the species share. are indigenous nearctic ichenumonids: Therefore the distribution of the parasitoids on Pleolophus N. swainei was studied in the three major plant indistinctus (Prov.), Endcasys subclcvatus (Say), communities, across ecotones between these com- Mastrus aciculatus (Prov.) and Gelis urbanus munities, and at different levels of the host pop- (Brues). All species oviposit within the cocoon ulation. The differences in distribution provided case on the surface of the host. All species may a basis for understanding the various evolutionary behave as primary parasitoids and all but E. sub- strategies adopted by parasitoids that exploit the clavatus are known to act as hyperparasitoids on same host. both larval and cocoon parasitoids. The life his- My approach has been to divide the parasitoid tory of P. basizonus in Canada was described by complex on N. swainei into two natural categories Griffiths (1961) and that of D. fuscipennis by or "guilds" (Root 1967). The guild concept fo- Morris and Cameron (1935). No detailed studies cuses attention upon groups of species that are have been made on the other species. In the study most likely to compete with one another. There- area all species were active from June until Oc- 446 PETER PRICE Ecology, Vol. 51, No. 3 tober. The ichneumonids were bi-voltine and the is dominated by three communities: pure stands eulophid at least tri-voltine. of jack pine, Pinus divaricata (Ait.) Dumont with The parasitoids that attack host larvae are a floristically depauperate understory of the eri- treated as a guild without reference to individual coid shrubs, Kalmia angustifolia L., Ledum groen- species in this paper. The members of the guild landicum Oeder, and Vaccinium angustifolium were the ichneumonid wasps Exenterus amicto- Ait.; pure stands of black spruce, Picea mariana rius (Panz.), E. diprionis (Roh.), Lamachus sp., (Mill.) BSP., with a similar shrub layer; and an Olesicampe lophyri (Riley), and the tachinid fly, association of hardwoods with paper birch, Betula Spathimeigenia spinigera Tnsd. Although the papyrifera Marsh., and aspen poplar, Populus species in this guild attack the host larva, the adult tremuloides Michx., with maples, Acer rubrum L. parasitoids emerge from the host cocoon in the and Acer spicatum Lam., entering the community following year. While in the cocoon they are on rocky slopes and eskers. The understory of available for hyperparasitism by members of the the hardwood stands has a comparatively rich cocoon parasitoid guild. Therefore the cocoon floristic composition of which Aralia nudicaulis parasitoid guild has the potential to inflict con- L., Aster macrophyllus L. and Clintonia borealis siderable mortality on the larval parasitoid pop- (Ait.) Raf. are typical. Linteau (1955) described ulation. these associations as the Kalmia-Vacciniumw,Cla- All larval and cocoon parasitoids mentioned in donia-Vaccinium, and Cornus-Maianthemum site this study have been found in the same sawfly types respectively. population. Eight plots were selected in five different jack Development of N. swainei, the most abundant pine stands. All but plot 3 were located in closed- host available to these parasitoids in the study canopy stands of 45-yr-old trees where the moss, area, was described by Tripp (1965). Adult saw- Pleurozium schreberi (Brid.) Mitt. and the li- flies emerge from overwintered cocoons in the chens, Cladonia spp., dominated the ground cover. forest litter in late June and oviposit in jack pine South slopes, such as the one where plot 8 was foliage. The eggs hatch after about 1 month and located, tended to be drier with a ground cover the larvae feed gregariously in the tree canopy of Cladonia spp. and pine-needle litter. Plot 3 until mid-September when they fall to the ground was located in a small stand of 20-yr-old jack pine, and spin cocoons. After adult sawfly emergence, adjacent to an older stand (plot 2). The trees the cocoon parasitoids encounter a severe host were open-grown with large unshaded areas where shortage until the larvae drop from the trees, about sweet fern, Cornptonia peregrina (L.) Coult., and 10 weeks later. cow-wheat, Melampyrum lineare Desr., grew in- terspersed with Cladonia spp., dry pine-needle lit- STUDY AREAS ter and bare sand. The 12 study plots were located on the Lauren- Two plots, 5 and 11, were located in black spruce tian Shield, in the St. Maurice River watershed stands and two plots, 4 and 12, in hardwood stands. in the transition zone between the mixed hard- Each was typical of the communities already de- wood-conifer and pure conifer biotic provinces scribed, consisting of natural regeneration after (Marie-Victorin 1964) (Table 1). The local area logging operations about 35 yr ago. TABLE 1. Location and main characteristics of study areas North lat West long No. Nearest Dominant tree of Plot lake degrees min degrees min species cages Other features 1 Cousacouta 47 16 73 37 45 10 site cc yrit jack pinei Sandy plain, typical 2 47 16 73 37 " 25 Transect across plain 3 it 47 15 73 37 20 yr jack pine 9 Open canopy, much bare sand it 4 47 14 73 36 Birch, poplar 5 Hardwood regeneration 5 it 47 14 73 35 Black spruce 5 Dense stand, dry site 6 McLaren 47 11 73 31 45 5 cc yr jacki pine Exposed hilltop 7 47 11 73 31 " d " 11 Transect from sandy plain to rocky hillside it t i i 8 47 12 73 29 A d 10 Dry south slope, little ground cover It it 9 Chevalier 47 03 73 43 Adt8 Transect on undulating topography 10 Chienne 47 04 73 27 AdAdAd 5d Sandy plain, typical site 11 Traine 47 05 73 25 Black spruce 5 Dense stand, dry site 12 47 04 73 23 Birch, maple, poplar 5 Hardwood regeneration Late Spring 1970 COEXISTENCE AMONG PARASITOIDS 447 METHODS jack pine at Lake Cousacouta was sampled in The distribution of parasitoids in the study areas two ways.