Description, Life History, and Parasitism of a New Species of Delphacid Planthopper (Hemiptera: Fulgoroidea)
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SYSTEMATICS Description, Life History, and Parasitism of a New Species of Delphacid Planthopper (Hemiptera: Fulgoroidea) 1 2 JAMES T. CRONIN AND STEPHEN W. WILSON Ann. Entomol. Soc. Am. 100(5): 640Ð648 (2007) ABSTRACT Delphacodes scolochloa Cronin & Wilson (Hemiptera: Fulgoroidea) is a newly discov- ered delphacid planthopper that feeds and oviposits exclusively on the stems of sprangletop, Scolochloa festucacea (Willd.) Link (Poaceae: Pooidae), in the prairie pothole region of northeastern North Dakota. D. scolochloa is bivoltine. It is also wing dimorphic, but populations are composed predom- inantly of ßightless brachypters. Macropters incur a substantial cost in terms of reduced longevity and fecundity relative to brachypters. The parasitoid complex attacking D. scolochloa consists of two egg parasitoids, Anagrus nigriventris Girault and Anagrus columbi Perkins (Hymenoptera: Mymaridae), and one undescribed dryinid. The proportion of eggs parasitized averages 21%, and egg parasitization is density independent. KEY WORDS Anagrus nigriventris, egg parasitoid, planthopper, prairie potholes, Scolochloa festucacea Grass-feeding delphacids have been the subject of ex- L. (Kiritani 1979, Wilson and Claridge 1985). These tensive ecological research, primarily because several delphacids also serve as hosts to mymarid, strep- species are vectors of pathogens of economically impor- sipteran, and dryinid parasitoids, an assemblage of tant grasses (Wilson 2005). In North American grass- parasitoids very comparable to that found for many lands and salt marshes, researchers have focused on fau- planthopper pest species (e.g., Oˆ take 1970, Bentur et nal surveys (e.g., Hendrickson, 1930, Wilson et al. 1993, al. 1982, Fowler et al. 1991). The study of hostÐpara- Hamilton 2002), systematics and morphology (e.g., Wil- sitoid interactions in species of grassland delphacids son 1982, Miller and Wilson 1999), life histories (e.g., may provide insight or potential biocontrol agents for Holder and Wilson 1992), and mating behavior (e.g., the suppression of related pest species. Heady and Denno 1991, Heady and Wilson 1990). Del- Here, we describe a new species of delphacid pl- phacids also have been used to address a broad range of anthopper and provide data on its biology, host range, ecological questions regarding plantÐherbivore interac- rate of macroptery, and population dynamics. We also tions (e.g., Tallamy and Denno 1979, Denno 1983, Denno identify the parasitoid species that attack the plan- et al. 1986, Wilson et al. 1994), life history theory (e.g., thopper, quantify rates of parasitism, and test whether Denno and Dingle 1981, Denno and McCloud 1985, the proportion parasitized among host plant patches Huberty and Denno 2006), population dynamics (e.g., depends on host density. Denno 1983, Strong 1989, Reeve et al. 1994), interspeciÞc competition (Denno and Roderick 1992, Denno et al. Materials and Methods 2000, Ferrenberg and Denno 2003), predatorÐprey in- teractions (Do¨bel and Denno 1994, Cronin et al. 2004, Study Location and Host Plant. Collection of spec- Fagan and Denno 2004), hostÐparasitoid interactions imens and census of planthopper and parasitoid pop- (e.g., Stiling and Strong 1982; Cronin and Strong 1990, ulations occurred in the prairie pothole region of 1993, 1999; Cronin 2003b), food web structure (e.g., northeastern North Dakota (Petersburg, Nelson Denno et al. 2002, Finke and Denno 2003, Gratton and County; 48.02097Њ N, 98.12230Њ W). This formerly gla- Denno 2003), dispersal and metapopulation dynamics ciated area is characterized by numerous seasonal and (Cronin 2003a, Haynes and Cronin 2006), and landscape permanent ponds (van der Valk 1989), and it is an ecology (Haynes and Cronin 2003, 2006; Cronin and important breeding habitat for waterfowl (Batt et al. Haynes 2004). 1989). Among the wetland plants bordering these Some grass-feeding delphacids are ecologically sim- ponds is the native sprangletop or common river grass, ilar to pests such as the brown planthopper, Nilapar- Scolochloa festucacea (Willd.) Link (Poaceae: Pooi- vata lugens (Stål.), a serious pest of rice, Oryza sativa dae). Sprangletop is circumpolar in distribution and in North America ranges from northern Iowa and Ne- 1 Corresponding author: Department of Biological Sciences, braska to southcentral Canada and west to Oregon Louisiana State University, Baton Rouge, LA 70803 (e-mail: (Gould 1968, Clayton and Renvoize 1986). Ecologi- [email protected]). 2 Department of Biology, University of Central Missouri, Warrens- cally, it is an emergent hydrophyte, and it is found burg, MO 64093. where water levels ßuctuate from 0 to 150 cm above 0013-8746/07/0640Ð0648$04.00/0 ᭧ 2007 Entomological Society of America September 2007 CRONIN AND WILSON:BIOLOGY OF NEW SPECIES OF DELPHACID PLANTHOPPER 641 Fig. 1. D. scolochloa. (A) Adult brachypterous female with dryinid parasitoid sac. (B) Macropterous and brachypterous females. (C) Egg clusters on S. festucacea stems that have by parasitized by Anagrus spp. (plant epidermis has been removed to expose eggs in the upper egg cluster). the soil surface (Sculthorpe 1967). Sprangletop often Planthopper Taxonomy. The specimens collected forms pure stands several meters deep along the pond from S. festucacea represent a new species, and they margins and as small clonal patches further in open are described below. Descriptions are based on nine shallow water (Smith 1973; J.T.C., unpublished data). males and 34 females. Using an ocular micrometer, Population Census. Three semipermanent ponds, body length was measured from the apex of the vertex broadly bordered by sprangletop, were used in this to posterior aspect of the abdomen. Thoracic length study (Ponds 100, 109, and N29-1). The ponds were was measured along the midline from the anterior 1Ð3 ha in size and were separated from each other by aspect of the pronotum to the posterior aspect of the 1.8Ð5.6 km. Shortly after spring thaw, mid-May 1998, mesonotal scutellum as the metanotum is obscured by the sprangletop bordering each pond was sampled the forewings. To describe and illustrate the male and with a sweepnet. Three sets of 10 sweeps each were female genitalia, the abdomen was removed and conducted per pond. Sweep samples were transported placed in a 10% KOH solution overnight, then washed on ice to the laboratory, and planthoppers were in- in distilled water; genitalia were stored in genitalia spected to determine stage (nymph or adult), sex, capsules Þlled with glycerol (Wilson 2005). The ter- wing morph (brachypters versus macropters; Fig. 1, A minology used in describing the female genitalia fol- and B), and presence of parasitoids protruding from lows Heady and Wilson (1990). between the abdominal segments (dryinid larval sac Host Range of D. scolochloa. To ascertain the host or strepsipteran cephalothorax) (Fig. 1A). Surveys range of the planthopper, we caged planthopper adults were conducted at 2-wk intervals until the end of the onto potted stems of the 11 dominant plant species at our summer when sprangletop had completely senesced Þeld sites. These plant species were the grasses spangle- (early September). The survey was continued each top, switch grass (Panicum virgatum L.), prairie season for 3 yr. cordgrass (Spartina pectinata Link), and smooth brome For each planthopper generation, after adults had (Bromus inermis Leyss.); common reed [Phragmites aus- become scarce, we conducted a census of planthopper tralis (Cav.) Trin]; sedges (Carex sp.); rushes (Scirpus eggs. Twenty-Þve sprangletop stems with planthopper sp.); cattails (Typha spp.); Canada goldenrod (Solidago eggs were haphazardly collected from each pond. canadensis L.); smart weed (Polygonum sp.); and thistle Stems were left to incubate for3datroom temper- (Circium sp.). Two common grass crops, corn (Zea mays ature (stem bases were submerged in water) to ensure L.) and wheat (Triticum aestivum L.), also were included that recently parasitized hosts could be identiÞed. in this study. Young, nonßowering shoots of the noncrop Stems were subsequently dissected under a stereo- plants were excavated from the Þeld in early June 1999 scopic dissecting microscope (10ϫ) to expose plan- and potted in Pro-mix BX soil (20-cm-diameter pots). thopper eggs inserted just beneath the stem epidermis Corn was grown from seed (Burpee Seed Co., Warm- (Fig. 1C). Planthopper eggs were identiÞed as either inster, PA, Early Choice Hybrid SE variety), and wheat healthy (developing white embryos or as cast chori- was obtained as seedlings from a local farmer. Both crop ons, indicating successful emergence), parasitized plants were grown in 20-cm pots by using Pro-Mix BX. (red immatures, or host chorions with circular exit Clear Mylar sheeting, formed into tubes, were inserted holes, indicating successful emergence), or dead over the plants. The tops of the cages were constructed (deßated or brown eggs). A subset of planthopper of mosquito netting hot-glued to the Mylar. Twenty-Þve nymphs and adults and egg parasitoids were preserved recently eclosed adult planthoppers were added to each in 70% ethanol for later identiÞcation of species. cage, and there were three caged plants per plant spe- 642 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 100, no. 5 cies. Cages were inspected daily for live planthoppers and evidence of feeding (excretion of honeydew). After all planthoppers had died, the plants were carefully in- spected for planthopper eggs. Longevity and Fecundity of Wing-Dimorphic Fe- males. Several hundred late