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1 September 2008 Survey for Potential Insect Biological Control Agents of Ligustrum sinense (Scrophulariales: Oleaceae) in China
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Abstract

A systematic survey of Chinese privet foliage, stems, seeds, and roots for associated phytophagous insects was conducted in China during 2005 and 2006 in order to establish basic information about the insect communities that Chinese privet harbors and to evaluate the abundance and damage caused by these insects. A total of 170 phytophagous insect species in 48 families and 5 orders were collected from Chinese privet in China. The insects belong to 4 feeding guilds: foliage, sap, stem, and root feeders. The impact of foliage feeders varied by site and over time. The mean percent defoliation of Chinese privet over all sites and years was 20.5 ± 8.2%, but ranged as high as 48%.

Chinese privet, Ligustrum sinense Lour., is an invasive exotic weed in the United States where it is a perennial, semi-evergreen shrub or small tree that grows to 10 m in height. This species is of great concern in the southeastern United States (Faulkner et al. 1989; Stone 1997), but it ranges from Texas to Florida, and as far north as the New England states (The Nature Conservancy 2004; University of Connecticut 2004). In addition, it has been introduced into Puerto Rico and Oregon (USDA-NRCS 2002).

Chinese privet is native to China, Vietnam, and Laos (The Nature Conservancy 2004; Wu & Raven 2003). In China it is found in the provinces south of the Yangzi River (Qui et al. 1992). It can be found between 200 and 2600 m in elevation where it occurs in mixed forests, valleys, along streams, thickets, woods, and ravines (Wu & Raven 2003; Qui et al. 1992).

Ligustrum sinense has been studied in China for its chemical composition and the medicinal value of its bark and leaves (OuYang 2003; OuYang & Zhou 2003). It has never been recorded as a noxious weed in either agricultural or forest settings. Chinese privet is grown nationwide as an ornamental hedge plant, for its berries used in brewing, and for oils extracted from seeds and used in soap making (Qui et al. 1992).

Chinese privet was introduced into the United States in 1852 (Coates 1965; Dirr 1990) as an ornamental shrub, for hedgerows (USDA-NRCS 2002), and sometimes as single specimens for its foliage and profusion of small white flowers (Dirr 1990; Wyman 1973). It is a forage plant for deer in the southeastern U.S. (Stromayer et al. 1998a; Stromayer et al. 1998b; The Nature Conservancy 2004). According to Small (1933), the species was escaping from cultivation in southern Louisiana by the 1930s. A survey of appropriate herbaria reveals collection records from Georgia as early as 1900. Based on herbarium records the species became naturalized and widespread in the southeast and eastern U.S. during the 1950s, 60s, and 70s (USDA-NRCS 2002).

Chinese privet is widely believed to drastically reduce native plant biodiversity because of its ability to shade out native vegetations (USDI Fish and Wildlife Service 1992; Merriam & Feil 2002) and form dense, monospecific stands that dominate the forest understory (Dirr 1990). Recent surveys in the southeastern United States show Chinese privet completely covers 0.9 million acres and colonies of varying densities can be found on another 17.6 million acres (Rudis et al. 2006). In 1998, the U.S. Department of Agriculture listed privet as one of 14 species with the potential to adversely affect management objectives in North Carolina’s National Forests. Similarly, the Florida Exotic Pest Plant Council lists Chinese privet as a Category 1 invasive species (FLEPPC 2007). More recently, The Nature Conservancy ranked Chinese privet as having high potential to disrupt the ecological balance (NatureServe 2006).

In addition to privet’s impact on natural landscapes, it can be directly harmful to humans. The flower of Chinese privet is toxic to humans causing symptoms such as nausea, headache, abdominal pain, vomiting, diarrhea, weakness, and low blood pressure and body temperature (USDA-NRCS 2002). Where Chinese privet occurs in abundance, floral odors may cause respiratory irritation (Westbrooks & Preacher 1986).

Repeated mowing and cutting will control the spread of L. sinense, but may not eradicate it (Tennessee Exotic Pest Plants Council 1996). Although modern herbicides, including glyphosate, effectively kill privet (Tennessee Exotic Pest Plants Council 1996; The Nature Conservancy 2004; Madden & Swarbrick 1990; Harrington & Miller 2005), environmental concerns will limit use of herbicides on public land or in sensitive areas. Faulkner et al. (1989) reported that in experimental trials of prescribed burning, there was no significant difference in the abundance of Ligustrum sinense in burned vs. unburned plots.

Plants unimportant in their native habitat may reach damaging levels when released from control by important natural enemies through introduction into new geographic areas (Van Driesche & Bellows 1996). This suggests that exploration for Chinese privet natural enemies in China might detect species suitable for use in a classical biological control program in the U.S. Chinese privet is considered a good candidate for classical biological control because it has no known biological control agents capable of lowering its pest status in North America (The Nature Conservancy 2004), and no native Ligustrum spp. occur there. Johnson & Lyon (1991) list at least 27 species of insects or mites that feed on Ligustrum spp. in the United States, however, none suppress populations of this plant in forests. In contrast, based on published records, China appears to have a rich complex of natural enemies that attack Ligustrum spp. (Zheng et al. 2004) and the potential for finding a biological control agent is high. However, little is known about their relative abundance and impact on their host plant. Therefore, a cooperative program was initiated in 2005 to survey for natural enemies of Chinese privet in China, with the goal of finding potential biocontrol agents. Here we report results of systematic surveys conducted in 2005 and 2006, and provide a list of the phytophagous insects found. We provide basic information about the insect community on Chinese privet in China and identify species that may have potential as biological control agents in the US. Preliminary information on impacts of different feeding guilds on Chinese privet in China is also reported.

Materials and Methods

Survey Sites

Our surveys focused primarily on 6 sites in Huangshan city (118.16, 29.43, elevation approximately 200 m) in Anhui Province, China, because the climate-matching program Climex (Hearne Scientific Software, Melbourne, Australia) indicated that this was the province most similar in climate to the southeastern United States (Sun et al. 2006). The seventh site was established in Guiyang (Huaxi, 106.40, 26.25, elevation approximately 1096 m), Guizhou Province, which is another area where Chinese privet is prevalent in China, but it is much further south and warmer than the Anhui sites.

In order to collect the most natural enemies of privet, our survey sites were selected to include habitats varying from natural areas to semi-natural and planted sites. These were as follows:

(1) Pure natural sites included 1 site in Lingnan adjacent to the Jiulong Natural resource conservation area in Xiuning County in the most southern region of Anhui Province; a second site on an unnamed island in the suburb of Huangshan city; and a third site in Guiyang. Chinese privet in these 3 sites grew naturally mixed with many other plant species and most grew relatively tall (over 3 m).

(2) Semi-natural sites included 1 site in Zhanchuan, a small town south of Huangshan city, and a second site at the Institute of Forestry (IOF) located in the north of Huangshan city. Chinese privet plants in these sites grew semi-naturally but were near agricultural lands where they were more likely to be disturbed by local residents.

(3) Planted sites included 1 site in She County east of Huangshan city. The other site was in the center of Huangshan city. Chinese privets in these sites were abundant and planted as ornamental shrubs along roads. All plants were small periodically pruned shrubs less than 1.5 m tall.

Systematic Sampling

At each site, 10 Chinese privet plants were randomly selected for sampling, marked with stakes and surrounded by a circle of colored tape and a sign to prevent human disturbance and for ease of relocation. Privet plants in Zhanchuan, IOF and She county in Anhui Province were surveyed from May to Aug, 2005 and from Apr to Oct, 2006. Privet on the island near Huangshan city was sample from Jun to Aug 2005 when sampling was discontinued because it was cleared for development. Privet in Lingnan, Huangshan city, and Guiyang were surveyed for 1 year from Apr to Sep 2006. At all sites, surveys were conducted at 10-day intervals during the survey periods. Collection of insects feeding on Chinese privet was accomplished by hand-picking, aspirating, and sweep-netting each sample plant 30 times. In some cases, cages were placed over branches to capture insects as they emerged. When immature insects were found, they were collected in a plastic bag together with the plant part on which they were feeding and returned to our lab to be reared to the adult stage to confirm the species.

Most insect species were identified by Professor Yang Chuncai (Anhui Agricultural University). Some Chrysomelidae were identified by Professor Wang Shuyong (Institute of Zoology, Chinese Academy of Sciences [CAS]), some Lepidoptera larvae were identified by Professor Wang Linyao (Institute of Zoology, CAS), and some Homoptera were identified by Professor Liang Aiping (Institute of Zoology, CAS). Pseudaulacaspis pentagona (Targioni-Tozzetti) was identified by Professor Xie Yingping (Shanxi University). Others were sent to the Zoological Museum of the Institute of Zoology, CAS, where they were distributed to appropriate taxonomists for identification. All phytophagous insects were evaluated based on their frequency of occurrence on Chinese privet, stage of development, and collecting site. Information on insect host range was obtained from the references “Economic Insect Fauna of China”, edited by the Editorial Committee of Fauna Sinica, Academia Sinica (Chou et al. 1985; Ge 1966; Tan et al. 1985; Yu et al. 1996; Liu 1963; Liu & Bai 1977; Wang 1980; Zhang 1985; Zhang 1995; Zhao & Chen 1980).

Foliage Damage

Defoliation rate was estimated by averaging defoliation of 120 leaves per plant. Samples consisted of 10 leaves randomly selected from 3 layers (high, middle, and lower layer) and 4 cardinal directions (east, south, west, and north) in each layer, for a total of 12 sampling locations on each plant. Defoliation was estimated by placing leaves on transparent graph paper with a 1-mm2 grid and measuring total leaf area and leaf area removed.

Stem Damage

Altogether, 900 plants of Chinese privet were investigated for signs of insect feeding within stems, oviposition, and damage at all survey sites. Stem damage was described by attributes, including physical shape, the distance of the damage from the ground, the diameter of stem with boring hole and so on.

Root and Seed Damage

Fifty roots were dug from randomly selected sample sites and examined for root damage. Adult insects found feeding on roots were collected for identification and larvae were returned to the laboratory with pieces of root for rearing to the adult stage. In order to detect insects feeding in seed, 500 immature seeds were collected randomly from survey sites and half were dissected in the laboratory. Also, 200 panicles with 25 to 58 mature seeds per panicle plus the remaining immature seeds were collected and placed in glass containers with fine gauze lids in order to collect adult seed-feeders emerging from them.

Results

The phytophagous insects associated with Chinese privet in China are listed in Table 1. In all, 170 species in 5 orders and 48 families were collected from Chinese privet in Anhui and Guizhou Province from 2005 to 2006. Insects were found in 4 different feeding guilds: foliage, sap, stem, and root feeders. Among them, 95.9% of insects were collected from privet leaves, 1 species was found feeding in stems and 6 species were root feeders. In contrast, only 27 species of insects feed on Ligustrum spp. in the U.S. Table 1 also includes an estimate of host range for each insect based on published reports.

Foliage-feeding Insects

Among the foliage-feeding insects in China, Argopistes tsekooni (Coleoptera: Chrysomelidae), Leptoypha hospita (Hemiptera: Tingidae) and an unidentified sawfly appeared to have the greatest impact on the plant. The extent of defoliation varied among sites, seasons and years (Fig. 1 and Fig. 2). In Zhanchuan, defoliation remained relatively constant throughout the sampling period fluctuating only slightly from 20% to 28% (Fig. 1A). Defoliation was highest at the She county site, averaging over 50% in late Jul 2005 (Fig. 1B). At the IOF site, defoliation ranged from about 15% in early May to about 27% in mid-Aug 2005. Defoliation in 2006 was generally higher at the IOF site, averaging about 34% for the year (Fig. 1C). Guiyang, a natural area, had the lowest defoliation of all sites averaging 1.6% in 2006 (Fig. 2). Lingnan had an average defoliation of 16% for 2006. Defoliation at this site ranged from a high of about 16% in late Apr to a low of less than 5% in Sep. Defoliation at the Huangshan city site, a site consisting of planted privet, generally declined over the season from a high of ca. 30% in mid-May to a low of ca. 8% in early Oct. A natural site on a nearby island had a defoliation rate of about 30% as well (Fig. 3), suggesting that defoliation was similar in an area regardless of site condition. The mean percent defoliation for Chinese privet per year among all sites and all years of study was 20.5 ± 8.2%.

Stem Damage

Phassus excrescens (Lepidoptera: Hepialoidae) was the only stem borer found feeding on Chinese privet in our survey where 5.3% (48 of 900 privets) of the plants were damaged by it. Larvae of the insect were collected from trunks of Chinese privet where they bored in the xylem causing galls. While feeding, they created an off white mass consisting of silk, excrement, and wood scraps that covered the entrance to the larval gallery. Borer entrance holes were 29.77±1.95 cm (n = 48) from the ground and the average diameter of attacked stems was 2.27±0.72cm (n = 48).

Root and Seed Damage

Six species of insects were found feeding on roots of Chinese privet. All 6 fed on fine roots or the root surface. Observations of the above ground plant health gave no indication root-feeders were present. No seed-feeders were found in either immature or mature seeds.

Discussion

In order to collect the most natural enemies of privet, survey sites were selected to include diverse habitats varying from natural areas to semi-natural and planted sites. Anhui province was selected as the primary survey area, because it was the best climatic match to the southeastern United States. Climatic matching is important for conventional biological control to insure the selected agents are adapted to the climate where they will be released (Andres et al. 1976; Harley & Forno 1992). Guizhou province was another important survey area because it is near the center of the range of Chinese privet in China.

Chinese privet is a common ornamental shrub but not a noxious weed in China, suggesting that natural enemies suppress populations. We found 170 phytophagous insect species on Chinese privet in China. Most were foliage-feeding insects despite phenolic compounds in privet leaves that likely provide some protection against damage from generalist herbivores (Swearingen et al. 2002). In the United States, Johnson & Lyon (1991) list at least 27 species of insects or mites that feed on Ligustrum spp., however, none suppress populations of this plant in forests. Most are not specialist on Chinese privet so it seems likely that the diverse and abundant insect fauna in China is important in regulating Chinese privet populations in its native habitat. Other factors also may be involved such as disease organisms not include in this survey. Phytophagous insects were collected by hand picking or net sweeping. Most were determined as feeding on Chinese privet by observations made during surveys in the field and through the literature. However, other insects that were not privet feeders and only occasionally rested on the plants were likely included in sweep net samples. To distinguish among them, we made notes in Table 1 showing which were confirmed as feeding on privet during our surveys (√), were collected during our surveys and recorded as privet feeders in the literatures (u), or collected from Chinese privet, but were not confirmed as privet feeders by personal observation or in the literature (s). We confirmed 81 feeding on privet by personal observation (Y. Z. Zhang), 88 were collected from privet and were reported in the literature as feeding on privet, and 1 species was collected but could not be confirmed by either method. Table 1 provides the most comprehensive listing of phytophagous insect feeding on Chinese privet to date.

When screening potential biological control agents for invasive weeds, their host range is one of the most important factors because only host specific agents will be considered for release to control invasive weeds. Polyphagous species were included in Table 1 to provide a complete listing with no attempt to differentiate good candidates for testing.

We included all insects found on privet, not just the most common ones, because some insects that are rare in their native country and suppressed by their own natural enemies are effective biological control agents when released from their own population regulating fauna.

Due to host specificity and the severe damage it caused on Chinese privet (Zhang et al., unpublished data), A. tsekooni may be the most promising biological control agent. It was apparent in our field surveys that defoliation rates were high when populations of A. tsekooni were large. Also, preliminary host specificity tests suggest its host range is restricted to Ligustrum spp. (Zhang et al., unpublished data). Examples of flea beetle as biological control agents of exotic weeds include Altica carduorum Guer. (Chrysomelidae: Coleoptera) on Cirsium arvense (L.) Scop. (Asteraceae) (Wan et al. 1996) and Agascicles hygrophila Selman et Vogt in China for control of Alternanthera philoxeroides (Mart.) Griseb., a global virulent weed from South America (Julien et al. 1995). Argopistes tsekooni feeds on most members of the genus Ligustrum. However, since no indigenous Ligustrum spp. occur in the U.S. and all Ligustrum spp. in the U.S. are listed as invasive weeds (Miller et al. 2004), A. tsekooni is a potential biological control that warrants further testing.

Leptoypha hospita could be another promising biocontrol agent because it has a limited host range in the Oleaceae (Li 2001) and often occurred in high numbers on Chinese privet in our sample areas. Likewise, the unidentified sawfly may also be an important defoliator. Thus far we have been unable to rear adults for identification so we cannot fully evaluate its potential for biological control. However, we have not observed it attacking other plant species during our field surveys.

Feeding by P. excrescens weakened the trunk resulting in breakage or, in some cases, the stems died as a result of girdling by the larvae. However, it has a broad host range and is considered an important pest of many plant species. Therefore, it is unlikely that it could be developed as a biological control agent.

Defoliation of privet varied widely among sites. The highest defoliation was recorded at the She County site which was a planted site. The lowest was on privet at the Guiyang site a natural area in Guizhou Province. That site was selected because it was near the center of the range of Chinese privet and, therefore, likely to have high number of phytophagous insects. We are uncertain why privet defoliation was low at this site but it may be the result of high numbers of defoliator natural enemies, or the more widely scattered and shaded privet population we sampled at that location. Defoliation of privet at the Lingnan site in Anhui Province, another natural area, was also relatively low compared with the She County site, or the Zhanchuan and IOF sites which we classified as semi-natural. The average 20% defoliation rate of L. sinense in China for all sites combined demonstrates that defoliating insects have a large impact on privet even when their own populations are being regulated by natural enemies. These results suggest that in the absence of natural enemies some of these insects may be effective biocontrol agents.

Acknowledgments

This research is part of an ongoing Sino-US Chinese privet biological control cooperative program funded by the USDA-Forest Service, Southern Research Station, Research Work Unit 4552, and the Natural National Science Foundation of China (30525009, 30621003). We are grateful to Ding Jianqing and Wei Wei for helpful comments on survey plan; Fang Fang, Li Li and Chen Yuhui for field assistance. We are also grateful to the many taxonomists who helped with identification of our specimens.

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Appendices

Fig. 1.

Seasonal Chinese privet defoliation at: (A) Zhanchuan (a semi-natural site), (B) She County (a planted site), and (C) IOF (a semi-natural site) in Anhui Province. Means ± SME are shown.

i0015-4040-91-3-372-f01.gif

Fig. 2.

Seasonal Chinese privet defoliation in 2006 in Guiyang, a natural site in Guizhou Province, Lingnan, a natural site in Anhui Province and Huangshan city, a planted site in Anhui Province. Means±SME are shown.

i0015-4040-91-3-372-f02.gif

Fig. 3.

Seasonal Chinese privet defoliation on an island natural site in Huangshan city in Anhui Province. Means ± SME are shown.

i0015-4040-91-3-372-f03.gif

Table 1.

Phytophagous insects collected from Chinese privet in China during 2005 and 2006 with notes on relative abundance, stage of development observed, method of confirmation of L. sinense as a host, plant part attacked, and host range.

i0015-4040-91-3-372-t101.gif

Table 1.

(Continued) Phytophagous insects collected from Chinese privet in China during 2005 and 2006 with notes on relative abundance, stage of development observed, method of confirmation of L. sinense as a host, plant part attacked, and host range.

i0015-4040-91-3-372-t102.gif

Table 1.

(Continued) Phytophagous insects collected from Chinese privet in China during 2005 and 2006 with notes on relative abundance, stage of development observed, method of confirmation of L. sinense as a host, plant part attacked, and host range.

i0015-4040-91-3-372-t103.gif

Table 1.

(Continued) Phytophagous insects collected from Chinese privet in China during 2005 and 2006 with notes on relative abundance, stage of development observed, method of confirmation of L. sinense as a host, plant part attacked, and host range.

i0015-4040-91-3-372-t104.gif
Yan-Zhuo Zhang, James L. Hanula, and Jiang-Hua Sun "Survey for Potential Insect Biological Control Agents of Ligustrum sinense (Scrophulariales: Oleaceae) in China," Florida Entomologist 91(3), (1 September 2008). https://doi.org/10.1653/0015-4040(2008)91[372:SFPIBC]2.0.CO;2
Published: 1 September 2008
JOURNAL ARTICLE
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