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Observations of Sagebrush Gall Morphology and Emergence of Rhopalomyia Pomum (Diptera: Cecidomyiidae) and Its Parasitoids

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Observations of Sagebrush Gall Morphology and Emergence of Rhopalomyia Pomum (Diptera: Cecidomyiidae) and Its Parasitoids Western North American Naturalist Volume 64 Number 3 Article 5 8-27-2004 Observations of sagebrush gall morphology and emergence of Rhopalomyia pomum (Diptera: Cecidomyiidae) and its parasitoids Ruth A. Hufbauer Colorado State University, Fort Collins Follow this and additional works at: https://scholarsarchive.byu.edu/wnan Recommended Citation Hufbauer, Ruth A. (2004) "Observations of sagebrush gall morphology and emergence of Rhopalomyia pomum (Diptera: Cecidomyiidae) and its parasitoids," Western North American Naturalist: Vol. 64 : No. 3 , Article 5. Available at: https://scholarsarchive.byu.edu/wnan/vol64/iss3/5 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Western North American Naturalist 64(3), © 2004, pp. 324–330 OBSERVATIONS OF SAGEBRUSH GALL MORPHOLOGY AND EMERGENCE OF RHOPALOMYIA POMUM (DIPTERA: CECIDOMYIIDAE) AND ITS PARASITOIDS Ruth A. Hufbauer1 ABSTRACT.—Galls induced by insects are specialized plant tissues thought to provide a suitable microclimate and high-quality food for insect development. Galls are also hypothesized to provide protection from predators, and particu- larly from parasitoids, because larger galls may be too deep for parasitoid oviposition (enemies hypothesis). However, galls may actually increase the risk of parasitism by making the location of gallformers more apparent (apparency hypothesis). Rhopalomyia pomum (Diptera: Cecidomyiidae) forms soft, lobed galls on big sagebrush (Artemisia triden- tata: Asteraceae). The volume of the largest galls can be 600X that of the smallest. Here I explore the relationships of the number of lobes per gall and gall volume with the emergence of R. pomum and its parasitoids. I collected 159 galls from 4 locations in southern Utah in spring 2002, measured them, and monitored emergence from each. Lobing was not related to midge emergence (F1,150 = 2.35, P = 0.13) but was positively associated with parasitoid emergence (F1,150 = 15.27, P < 0.001), suggesting that early season parasitoids attacking before gall development may contribute to lobe for- mation by disrupting cues from eggs or larvae to the plant, or that flies in lobed galls are more accessible to oviposition by late season parasitoids. More midges emerged from larger galls than from smaller galls (F1, 150 = 22.0, P < 0.001), but gall size was not related to parasitoid emergence (F1,150 = 0.3, P = 0.6), providing no support for either the enemies or apparency hypothesis. Key words: gall, Rhopalomyia, Artemisia, Cecidomyiidae, sagebrush, parasitoid, parasitism. Gall-forming insects stimulate their host “enemies hypothesis” (Askew 1965, Hawkins plant to develop tumor-like growths that pro- and Gagné 1989, Cornell 1990, Schultz 1992, vide them with food and shelter (Abrahamson Schonrogge et al. 1996, Stone and Cook 1998, and Weis 1997). Morphology of the galls Tscharntke et al. 2001). Chemical composition, depends upon the species that induces it, and size, and hardness are several characteristics characteristics of galls are often used in iden- of galls that may influence rates of attack by tification of gallformers (e.g., Gagné 1989). parasitoids (Cornell 1983). Gall size seems Abrahamson and Weis (1997) argue that galls particularly important, as parasitoids may not can be seen as adaptations of the insects that be able penetrate large galls deeply enough induce them rather than of the plant. How- with their ovipositor to reach the gallformers, ever, there is ongoing debate about the adap- leading to lower rates of parasitism of insects tive nature of insect galls. Price et al. (1987) in larger galls (e.g., Askew 1965, Price and described the competing hypotheses and Clancy 1986, Plakidas and Weis 1994, Abra- argued cogently that galls function to provide hamson and Weis 1997). However, presence of superior nutritional resources and a more suit- a gall may also make insects more apparent to able microclimate for gallformers. Subsequent their natural enemies, as pointed out by research on nutritional quality of galls (Hartley Hawkins and Gagné (1989). Increasing gall and Lawton 1992, Hartley 1998, Schonrogge size may increase apparency of galls, poten- et al. 2000), water potential in galls (Fay et al. tially increasing the risk of parasitism rather 1993), and distribution of galls in arid and than decreasing it (Price et al. 1987), similar to mesic biomes (Fernandes and Price 1992, Price the idea that the apparency of a plant influ- et al. 1998) lends support to those conclusions. ences the risk of herbivore attack (Feeny However, there is ongoing debate over whether 1976). For simplicity I call this explanation for galls also act as a barrier to predation and par- a positive association between gall apparency asitism, what Price et al. (1987) describe as the and natural enemy attack the “apparency 1Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, CO 80523-1177. 324 2004] GALL SIZE, GALL FLIES, AND PARASITOIDS 325 hypothesis.” Support for this idea comes from shrub of great ecological importance through- Hawkins and Gagné (1989), who found that out its range in western North America (Mon- cecidomyiid midges with more apparent galls sen and Shaw 2000). In the Great Basin, sage- have more species of parasitoids associated brush-dominated vegetation accounts for with them than cecidomyiids with less appar- approximately 46% of all habitats, and on the ent galls, leaf rolls, or other unapparent modes Colorado Plateau it accounts for approxi- of feeding. However, this pattern does not mately 12% of all habitats (West 1979). There always hold because no significant differences are 3 parapatric subspecies of A. tridentata, in parasitism between galling and nongalling each of which uses slightly different environ- species of grass-feeding chalcids were found ments (McArthur and Sanderson 1999, Mon- by Tscharntke et al. (2001). Lack of relationship sen and Shaw 2000). Artemisia tridentata is between gall size and parasitism may be due tolerant to browsing (e.g., Messina et al. 2002) to parasitism occurring during the “window of and provides important forage and habitat for vulnerability” before the galls grow around the many vertebrates. In addition, it is host to spe- gallformers (Washburn and Cornell 1979, Craig cialized herbivorous insects including many et al. 1990, Briggs and Latto 1996). gall-forming midges. For example, in Idaho, Cecidomyiid gall midges are among the 26 species of gall midges develop on the 3 sub- most common and speciose of the gall-forming species of big sagebrush (Jones et al. 1983). insects. They are an ancient group found world- Rhopalomyia pomum Gagné (Cecidomyi- wide (Gagné 1989) that includes economically idae), a gall midge specific to A. tridentata, is important pests (e.g., Mayetiola destructor Say, among the more abundant galling species found the Hessian fly) and biological control agents throughout most of the range of its host plant of weeds (Peschken et al. 1989, Ehler and Kin- (Gagné 1989). It forms galls on all 3 subspecies sey 1993, Hinz and Müller-Schärer 2000, of A. tridentata and on hybrids between them Skuhravá and Hinz 2000, Sobhian et al. 2000). (Graham et al. 2001). It is univoltine: eggs are Although some cecidomyiids are well studied, laid on leaves in late spring, galls form in late the biology and ecology of others are poorly summer and larvae develop and overwinter understood, and new species and even genera within them, pupation occurs in the galls, and continue to be described at a rapid pace (Fedo- pupae break through the gall surface the fol- tova 2000, Gagné 2002, Kolesik et al. 2002). lowing spring prior to adult emergence (Jones Rhopalomyia pomum Gagné (Cecidomyi- et al. 1983). The galls are globular to lobed, idae), a common gall-forming midge of big soft and spongy, reddish, pale, or green, and sagebrush (Artemisia tridentata Nutt., Aster- covered in short trichomes (Fig. 1; Gagné 1989). aceae), is unusual among gallformers in that it induces a wide range of gall morphologies. METHODS Rhopalomyia pomum galls can have from 1 to many lobes (Jones et al. 1983) and can range Galls were collected on 7 May 2002 from from 0.35 cm to over 3 cm in diameter, with approximately 20 plants at 3 locations in Gar- volumes varying 600-fold (see below). The phys- field County, Utah (Table 1). The first 2 loca- iological mechanisms leading to this variation tions were in the Dixie National Forest, north in gall morphology are not known. Here I of Bryce Canyon National Park, and the 3rd employ the naturally occurring wide range of location was near the Lower Box entrance of gall morphologies to explore associations be- tween R. pomum gall size and lobing on para- the Box Death Hollow Wilderness. At that 3rd sitism in this system. I am particularly interested location, 2 collections were made: (1) from a in determining whether there is either a nega- population of sagebrush spanning about 1 ha tive or a positive association between gall size in a dry wash, and (2) from 1 of 4 heavily and parasitism, which would lend support to the galled plants along the roadside. To sample enemies hypothesis or the apparency hypothe- galls of all sizes without bias, I collected all of sis, respectively. the galls from each of the shrubs sampled, a total of 159 galls. STUDY SYSTEM I counted the number of lobes on each gall, and then I measured each gall across its Artemisia tridentata Nutt. (big sagebrush, widest dimension (x) and perpendicular to that Asteraceae) is an erect, aromatic evergreen (y). Gall volume was approximated using the 326 WESTERN NORTH AMERICAN NATURALIST [Volume 64 TABLE 1. Geographical coordinates of sample locations, elevations, and sample sizes (N).
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