Photoperiodic Response of Abrostola Asclepiadis (Lepidoptera: Noctuidae), a Candidate Biological Control Agent for Swallow- Worts (Vincetoxicum, Apocynaceae)
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The Great Lakes Entomologist Manuscript 2340 Photoperiodic Response of Abrostola asclepiadis (Lepidoptera: Noctuidae), a Candidate Biological Control Agent for Swallow- worts (Vincetoxicum, Apocynaceae) Lindsey R. Milbrath Margarita Dolgovskaya Mark Volkovitsh René F.H. Sforza Jeromy Biazzo Follow this and additional works at: https://scholar.valpo.edu/tgle Part of the Entomology Commons This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at [email protected]. Milbrath et al.: Abrostola asclepiadis Photoperiod Response 2019 THE GREAT LAKES ENTOMOLOGIST 71 Photoperiodic Response of Abrostola asclepiadis (Lepidoptera: Noctuidae), a Candidate Biological Control Agent for Swallow-worts (Vincetoxicum, Apocynaceae) Lindsey R. Milbrath1*, Margarita Dolgovskaya2, Mark Volkovitsh2, René F.H. Sforza3, Jeromy Biazzo1 1 United States Department of Agriculture, Agricultural Research Service, Robert W. Holley Center for Agriculture and Health, 538 Tower Road, Ithaca, NY 14853, USA 2 Zoological Institute, Russian Academy of Sciences, 199034, St. Petersburg, Russia 3 USDA-ARS European Biological Control Laboratory, CS 90013 Montferrier-sur-Lez, 34988 St Gély du Fesc, France *Corresponding author: (email: [email protected]) Abstract A biological control program is in development for two swallow-wort species (Vince- toxicum, Apocynaceae), European vines introduced into northeastern North America. One candidate agent is the defoliator Abrostola asclepiadis (Denis and Schiffermüller) (Lepi- doptera: Noctuidae). The moth reportedly has up to two generations in parts of its native range. We assessed the potential multivoltinism of Russian and French populations of the moth by rearing them under constant and changing photoperiods, ranging from 13:11 to 16:8 hour (L:D). The French population was also reared outdoors under naturally-changing day lengths at a latitude similar to northern New York State. Less than six adult moths emerged, with one exception, for any photoperiod treatment. We expect A. asclepiadis to be univoltine if it were to be released into North America, limiting its potential impact on swallow-worts. It should therefore be given a lower priority for release. Keywords: Cynanchum rossicum, Cynanchum louiseae, swallow-wort, weed biolog- ical control, photoperiod Understanding the potential efficacy (Apocynaceae-subfamily Asclepiadoideae) of a candidate weed biological control agent are European twining vines introduced into has become more prominent in recent years, North America in the mid- to late-1800s. especially pre-release studies of an agent’s They have become invasive over the last 40 impact on plant performance (e.g., Grevstad years in a variety of natural areas and pe- et al. 2013, Reddy and Mehelis 2015). Such rennial cropping systems, ranging from open assessments have been proposed for prior- fields to forest understories. The primary itizing agents for further study or release, infestations occur in southeastern Canada in order to enhance the success rate of weed and the northeastern United States, with biological control programs (Balciunas and increasing concerns in the Midwest, includ- Coombs 2004, McClay and Balciunas 2005). ing states and provinces of the Great Lakes The seasonal duration and amount of dam- Basin (DiTommaso et al. 2005, Averill et age that an agent might inflict on a targeted al. 2011, USDA/NRCS 2019). Both species invasive plant will depend in part on the are long-lived, herbaceous perennials that number of agent generations (voltinism) spread via wind-dispersed seeds. Floral and produced in the area of release. The induc- faunal communities are negatively affected tion of diapause in an insect population is a by swallow-worts, including the globally-rare key component determining the number of alvar ecosystems of the Lake Ontario basin generations and seasonality of insect activ- (DiTommaso et al. 2005, Ernst and Cap- ity. For many insects of temperate zones, puccino 2005). Broad-spectrum herbicides photoperiod is a primary cue for diapause have been the main management tool, but induction (Tauber et al. 1986). are not necessarily desired in natural areas. Mechanical control is generally unsatisfac- Black swallow-wort [Vincetoxicum tory in reducing swallow-wort densities and nigrum (L.) Moench = Cynanchum loui- cover (DiTommaso et al. 2013). A biological seae Kartesz and Gandhi] and pale swal- control program has been in development low-wort [V. rossicum (Kleopow) Barbar. = for several years, but to date only the moth Cynanchum rossicum (Kleopow) Borhidi] Hypena opulenta (Christoph) (Lepidoptera: Published by ValpoScholar, 1 Submission to The Great Lakes Entomologist 72 THE GREAT LAKES ENTOMOLOGIST Vol. 52, Nos. 3–4 Erebidae) has been released in Canada dage, France (44°42.033’ N, 05°37.000' E). (2013) and more recently in the U.S. (Weed Larvae were reared on V. hirundinaria at et al. 2011b, Young and Weed 2014, Milbrath the European Biological Control Laboratory and Biazzo 2016). (EBCL), Montferrier-sur-Lez, France, under One candidate biological control agent a 10:14 hour (L:D) photoperiod to induce is the defoliating moth Abrostola asclepiadis diapause and pupae were shipped to the (Denis and Schiffermüller) (Lepidoptera: senior author. Voucher specimens have been Noctuidae). It is widely distributed in deposited with the Cornell University Insect Europe in open field to shaded habitats. Collection, Department of Entomology, Itha- In some years and locations it defoliates ca, NY (under Lot Number 1263). entire stands of Vincetoxicum hirundina- Diapausing pupae were overwintered ria Medik., its primary host plant (Förare at 5°C and a 10:14 hour (L:D) photoperiod for 1995, Leimu and Lehtilä 2006). The moth up to 9 months. Pupae were then transferred overwinters as a pupa in leaf litter and has to a chamber set at 14:10 hour (L:D) and one-two generations per year. Two genera- 25:20°C for adult emergence, mating, and tions have been reported in some areas of oviposition. Two groups of adult moths were central Europe (Förare 1995) that would held in 60 3 60 3 60 cm aluminum-mesh cag- have similar latitudes to northern New York es containing honey-water and potted pale and southern Ontario. Abrostola asclepiadis or black swallow-wort plants for oviposition. appears to be host specific to Vincetoxicum Leaves with egg masses were collected daily spp. (Hazlehurst 2011). However, a single (< 24 hour post-oviposition) to minimize the defoliation of black or pale swallow-wort time at a non-experimental photoperiod. has a limited effect on the plants (Weed et Initial diapause test. Neonate lar- al. 2011a, Milbrath et al. 2019). A bivoltine vae (1st laboratory generation) were placed population of A. asclepiadis that can damage individually into ventilated 0.25 liter plastic the plants over a greater duration of the vials and randomly assigned to custom-made season is therefore desired for release into programmable chambers set to 20 or 25˚C North America. Many noctuid species have a and one of six photoperiod treatments facultative diapause, in which the number of at each temperature (hours, L:D): 13:11, generations produced depends in part on the 16:8, 13:11 to 16:8 fast, 16:8 to 13:11 fast, response of populations at a given latitude 13:11 to 16:8 slow, or 16:8 to 13:11 slow. A to environmental cues such as photoperiod short day of 13 hours represents natural (Tauber et al. 1986, Saulich et al. 2017). day lengths in the North Caucasus in ear- No diapause-induction studies have been ly-April or early-September, and a long day conducted with A. asclepiadis. of 16 hours exceeds the longest summer Our objective was to determine the day length (sunrise to sunset, Fig. 1). For potential number of generations of different the fast photoperiod change, larvae were populations of A. asclepiadis that could oc- transferred to the alternate photoperiod at cur in the invasive range of swallow-worts. approximately the third instar or halfway Populations were exposed to photoperiods through their larval development (day 11 at they would typically experience during the 20°C, day 8 at 25˚C). For the slow change, growing season in the northeastern U.S. day length was increased (or decreased) daily and southeastern Canada. The results were in a stepwise fashion over a 5 day (20°C) or to be used in concert with separate impact 3 day (25°C) period. Ten larvae were used studies and a population matrix model for per treatment combination for each of the swallow-worts (Milbrath et al. 2018, 2019) to two Russian populations for a total of 240 predict the efficacy of this candidate agent. larvae. Larvae were fed excised leaves from Vincetoxicum sp. plants that had also been Materials and Methods collected from the North Caucasus region. The larvae were checked daily and leaves Insect cultures. Eggs of Russian A. were replaced. Upon pupation, pupae were asclepiadis were collected 29 May–6 June maintained under their same experimental 2013 from Vincetoxicum spp. leaves in the rearing conditions for an additional 40 days, Russian North Caucasus, near Kislovodsk which is over twice the expected time for (43°56.400' N, 42°41.734' E) and Borgus- emergence from non-diapausing pupae (L. tanskaya (44°02.0436' N, 42°30.161' E). Eggs Milbrath, unpublished data). Live pupae were transported to the Zoological Institute, that had not emerged as adults at the end Russian Academy of Sciences, St. Petersburg of this period were considered to be in dia- for the first diapause experiment (see Ini- pause. Temperature, photoperiod and source tial diapause test). Diapausing pupae were population were treated as a single combined subsequently shipped to the senior author factor and tested against the categorical data for additional experiments described below. of diapause (yes or no) using the G-test with Eggs of French A.