Coleoptera: Buprestidae) in Northeast China, with Notes on Two Species of Parasitic Coleoptera
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329 Natural enemies of emerald ash borer (Coleoptera: Buprestidae) in northeast China, with notes on two species of parasitic Coleoptera Xiao-Yi Wang,1 Liang-Ming Cao, Zhong-Qi Yang, Jian J. Duan, Juli R. Gould, Leah S. Bauer Abstract—To investigate natural enemies of emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), in northeastern China, we conducted field surveys of ash (Fraxinus Linnaeus (Oleaceae)) trees in semi-natural forests and plantations at variable EAB densities from 2008 to 2013. Our surveys revealed a complex of natural enemies including eight hymenopteran parasitoids and two apparently parasitic Coleoptera, woodpeckers, and several undetermined mortality factors. Parasitoid complex abundance and its contribution to EAB mortality varied with the time of year, type of ash stands, and geographic regions. The egg parasitoid Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) and the larval parasitoid Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) were frequently observed in Jilin, Liaoning, and Heilongjiang provinces and in Beijing, but not in Tianjin. Spathius agrili Yang (Hymenoptera: Braconidae), however, was more prevalent near Beijing and further south in Tianjin. Larvae of two species of apparently parasitic beetle, Tenerus Laporte (Coleoptera: Cleridae) species and Xenoglena quadrisignata Mannerheim (Coleoptera: Trogossitidae), were also recovered attacking overwintering EAB in Liaoning Province, with Tenerus species being a dominant mortality agent (~13%). Our findings support the need to consider the geographic origin of insect natural enemies for EAB biocontrol, as well as an expanded foreign exploration for EAB natural enemies throughout its native range in Asia. Introduction ecosystems (Cappaert et al. 2005; Poland and McCullough 2006; Herms and McCullough Emerald ash borer (EAB), Agrilus planipennis 2014; Bauer et al. 2015). Most recently, Orlova- Fairmaire (Coleoptera: Buprestidae), is an invasive Bienkowskaja (2014) proposed that EAB infes- insect pest of ash (Fraxinus Linnaeus; Oleaceae) tations detected in the European region of Russia trees, introduced to North America from Asia would likely expand throughout Europe. Early (Haack et al. 2002). Since initial detection in EAB eradication efforts in North America were Michigan, United States of America in 2002, this unsuccessful, and chemical control is expensive pest has been found in 24 other states and two and environmentally unacceptable in natural Canadian provinces, killing tens of millions of forests. Therefore, efforts to manage EAB now ash trees and causing severe damage to forested emphasise the use of classical biological control Received 18 October 2014. Accepted 6 July 2015. First published online 7 December 2015. X.-Y. Wang,1 L.-M. Cao, Z.-Q. Yang, Key Laboratory of Forest Protection, State Forestry Administration, Research Institute of Forest Ecology, Environment and Protection, Chinese Academy of Forestry, Beijing 100091, China J.J. Duan, United States Department of Agriculture, Agricultural Research Service, Beneficial Insects Introduction Research Unit, Newark, Delaware 19713, United States of America J.R. Gould, United States Department of Agriculture, Animal and Plant Health Inspection Service, Plant Protection and Quarantine, Center for Plant Health Science and Technology, Buzzards Bay, Massachusetts 02542, United States of America L.S. Bauer, United States Department of Agriculture, Forest Service, Northern Research Station, Lansing, Michigan 48910, United States of America 1Corresponding author (e-mail: [email protected]). Subject editor: Dylan Parry doi:10.4039/tce.2015.57 Can. Entomol. 148: 329–342 (2016) © 2015 Entomological Society of Canada 330 Can. Entomol. Vol. 148, 2016 (Bauer et al. 2008, 2015; Duan et al. 2010, 2013; introduced parasitoids attacking EAB in North Abell et al. 2014). This well accepted, long-term, America, the results of recent field studies have and sustainable approach to the management of also revealed that some native parasitoids of invasive species, involves the introduction and wood-boring larvae, such as species of Atanycolus establishment of co-evolved natural enemies from Förster (Hymenoptera: Braconidae) and Spathius the native range of an invasive pest (Van Driesche Nees (Hymenoptera: Braconidae), have acquired et al. 2008; Van Driesche 2014). EAB as a novel host (Cappaert and McCullough Following the invasion of North America by 2009, 2010; Duan et al. 2009, 2010, 2011, 2012a; EAB, field surveys of EAB natural enemies were Marsh et al. 2009; Taylor et al. 2012; Bauer et al. conducted in low to heavily infested ash stands in 2014, 2015). However, these parasitoids are plantations, city parks, and in semi-natural rural apparently opportunists, more specific to a niche and urban forests in the native range of EAB in than to host. Moreover, they are associated with northeast China, Russian Far East, and South high density EAB populations in some heavily Korea (Liu et al. 2003, 2007; Duan et al. 2012c; infested areas and may not significantly suppress Bauer et al. 2014, 2015). These field surveys led EAB populations at the leading edge of the to the discovery and description of several species invasion or in the aftermath forest where EAB of hymenopteran parasitoids: the egg parasitoid densities are still low (Bauer et al. 2014, 2015; Oobius agrili Zhang and Huang (Hymenoptera: Duan et al. 2014). Woodpecker predation was Encyrtidae) (Zhang et al. 2005), and the larval also higher at sites with more dense EAB popu- parasitoids: Spathius agrili Yang (Hymenoptera: lations (Jennings et al. 2013). Thus, co-evolved Braconidae) (Liu et al. 2003; Yang et al. 2005), natural enemies associated with low pest densities Tetrastichus planipennisi Yang (Hymenoptera: in the native range of EAB may be more effective Eulophidae) (Yang et al. 2006), Sclerodermus at suppressing the target population than generalist pupariae Yang and Yao (Hymenoptera: and opportunistic natural enemies that capitalise on Bethylidae) (Yang et al. 2012) from China, and pests during outbreaks. Spathius galinae Belokobylskij and Strazenac Here we report on EAB natural enemy surveys (Hymenoptera: Braconidae) from Russian Far conducted from 2008 to 2013 in EAB-infested East and South Korea (Belokobylskij et al. 2012; ash stands in northeastern China where EAB Duan et al. 2012c). Other hymenopteran para- densities were generally low in native ash trees sitoids of Agrilus Curtis species known to attack (mainly Fraxinus mandshurica Ruprecht) or high EAB in Asia and Europe are summarised in Bauer in exotic ash trees, mainly F. velutina Torrey. et al. (2015). Based on the results of our surveys, we assessed In an effort to manage EAB in the United States the relative abundance and rate of EAB stage- of America, introductions of three parasitoid specific mortalities caused by different groups of species from China, O. agrili, S. agrili, and natural enemies in relation to different host ash T. planipennisi, began in Michigan in 2007 and tree species, geographic regions, and seasonality subsequently expanded to other states (Federal in China. Register 2007; Bauer et al. 2008, 2014, 2015). By 2009, the introduced parasitoids were recovered at many of these early release sites, prompting the Materials and methods United States Department of Agriculture to initi- ate an EAB biocontrol programme and construct a Study sites parasitoid-rearing facility, leading to the expan- Field surveys were conducted in north and sion of EAB biocontrol to other EAB-infested northeast China at six study sites in the two regions of North America (Bauer et al. 2015). provincial cities of Beijing and Tianjin and in the Researchers evaluating EAB population provinces of Liaoning, Jilin, and Heilongjiang dynamics at long-term study sites in Michigan from 2008 to 2013 (Fig. 1, Table 1). Native have discovered that only two species, O. agrili F. mandshurica and exotic F. velutina trees grow- and T. planipennisi, have successfully established ing in parks, semi-natural forests or monoculture stable and increasing populations (Duan et al. plantations were sampled for EAB eggs and larvae. 2013, 2014; Abell et al. 2014). Besides the Because the EAB can develop in one or two years © 2015 Entomological Society of Canada Wang et al. 331 Fig. 1. Sample locations in China. The native range of EAB in Asia as reported in the literature, and regions of Asia where EAB natural enemies were surveyed prior to this report. The red dots indicate EAB study sites surveyed for natural enemies from 2008 to 2013, with the numbers corresponding to the following locations: 1. Dagang, Tianjin; 2. Haidian, Beijing; 3. Fengcheng, Liaoning; 4. Shenyang, Liaoning; 5. Changchun, Jilin; 6. Shangzhi, Heilongjiang. Numbers on the top line of graph frame are east longitude degrees, while numbers on the right side are north latitude degrees. Service layer credits: United States National Park Service. Data source for the native range of EAB: https://sites.google.com/site/eduardjendek/world-distribution-of-agrilus-planipennis. Map created by United States Department of Agriculture, Forest Service, Northeastern Area State and Private Forestry, Office of Knowledge Management (Durham, New Hampshire, United States of America). depending on climate, surveys of EAB larvae and both univoltinism and semivoltinism occurs in associated mortality factors were conducted in Liaoning Province. In contrast, a majority of EAB either spring or autumn, or both seasons every populations exhibit