Pink Gypsy Moth - aphis - US Department of Agriculture
Pink Gypsy Moth - aphis - US Department of Agriculture
Pink Gypsy Moth - aphis - US Department of Agriculture
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Mini Risk Assessment<br />
<strong>Pink</strong> <strong>Gypsy</strong> <strong>Moth</strong>, Lymantria mathura Moore<br />
[Lepidoptera: Lymantriidae]<br />
Erica E. Davis 1 , Sarah French 1 , & Robert C. Venette 2<br />
1-<strong>Department</strong> <strong>of</strong> Entomology, University <strong>of</strong> Minnesota, St. Paul, MN<br />
2-North Central Research Station, <strong>US</strong>DA Forest Service, St. Paul, MN<br />
September 29, 2005<br />
A<br />
B<br />
C<br />
D<br />
Figure 1. Lymantria mathura: (A&B) Adult female with pink hind wings; (C) adult male<br />
with yellow hind wings ; (D) larva on foliage <strong>of</strong> deciduous host. Images not to scale.<br />
[Images (A-C) courtesy <strong>of</strong> W. Wallner, <strong>US</strong>DA Forest Service,<br />
http://www.inspection.gc.ca/english/sci/surv/data/lymmate.shtml (D) David Mohn,<br />
www.forestryimages.org]<br />
Table <strong>of</strong> Contents<br />
Introduction......................................................................................................................... 2<br />
1. Ecological Suitability.................................................................................................. 2<br />
2. Host Specificity/Host Availability.............................................................................. 3<br />
3. Survey Methodology................................................................................................... 7<br />
4. Taxonomic Recognition.............................................................................................. 8<br />
5. Entry Potential ............................................................................................................ 8<br />
CAPS PRA: Lymantria mathura 1
6. Destination <strong>of</strong> Infested Material ................................................................................. 9<br />
7. Potential Economic Impact......................................................................................... 9<br />
8. Potential Environmental Impact ............................................................................... 10<br />
9. Establishment Potential............................................................................................. 10<br />
References......................................................................................................................... 11<br />
Appendix A. Geographic distribution.............................................................................. 15<br />
Appendix B. Host distribution ......................................................................................... 18<br />
Appendix C. Taxonomy and morphology ....................................................................... 20<br />
Appendix D. Threatened or endangered plants................................................................ 23<br />
Appendix E. Biology ....................................................................................................... 30<br />
Introduction<br />
The pink or rosy gypsy moth, Lymantria mathura Moore, is a major defoliator <strong>of</strong><br />
deciduous trees in the Palearctic, primarily in eastern Asia from India to the Russian Far<br />
East (Roonwal 1979b, Baranchikov et al. 1995, CAB 2004, EPPO 2005). Spurred by<br />
concerns surrounding L. mathura, the <strong>US</strong> <strong>Department</strong> <strong>of</strong> <strong>Agriculture</strong>-Animal and Plant<br />
Health Inspection Service, <strong>US</strong>DA Forest Service and Russian counterparts have<br />
developed an early warning system to alert <strong>US</strong> pest <strong>of</strong>ficials about periods <strong>of</strong> increased<br />
insect activity and prevent the introduction <strong>of</strong> this insect (Anon. 2001). <strong>US</strong> <strong>of</strong>ficials are<br />
also alerted when New Zealand finds a Russian freighter to be infested with this insect<br />
(<strong>US</strong>DA 2001b).<br />
Risks associated with L. mathura have been evaluated previously. In the Exotic Forest<br />
Pest Information System, L. mathura was considered to pose a very high risk to North<br />
America forests relative to other forest pests and pathogens, and this assessment was<br />
given with a very high degree <strong>of</strong> certainty (Rosovsky 2001). Gninenko and Gninenko<br />
(2002) proposed a scoring system to evaluate the relative propensity <strong>of</strong> different<br />
lymantriids to be moved by international shipping. These authors suggest that L.<br />
mathura is less likely than L. dispar or L. monacha to be moved by shipping, but it is<br />
more likely to be moved than 26 other species <strong>of</strong> Lymantriidae. Limited biological<br />
information about lymantriids <strong>of</strong> the Russian Far East, including L. mathura, complicates<br />
the assessment <strong>of</strong> risk (Gninenko and Gninenko 2002). The purpose <strong>of</strong> this mini-pest<br />
risk assessment is to further evaluate several factors that contribute to risks posed by<br />
L. mathura and apply this information to the refinement <strong>of</strong> sampling and detection<br />
programs.<br />
1. Ecological Suitability. Rating: Medium. Lymantria mathura is present<br />
throughout much <strong>of</strong> Asia. Appendix A provides a detailed list <strong>of</strong> the reported<br />
worldwide distribution <strong>of</strong> this insect. In general, L. mathura occurs in cool,<br />
temperate to warm climates with varying amounts <strong>of</strong> seasonal rainfall, and dry<br />
periods. The currently reported distribution <strong>of</strong> L. mathura suggests that the pest<br />
may be most closely associated with biomes characterized as: temperate broadleaf<br />
and mixed forests; temperate coniferous forests; tropical and subtopical dry<br />
broadleaf forests; and tropical and subtropical moist broadleaf forests. Of these<br />
biomes, only tropical and subtopical dry broadleaf forests do not occur in the <strong>US</strong>.<br />
CAPS PRA: Lymantria mathura 2
Consequently, we estimate that approximately 38% <strong>of</strong> the continental <strong>US</strong> would<br />
have a suitable climate for L. mathura (Fig. 2). See Appendix A for a more<br />
complete description <strong>of</strong> this analysis.<br />
Figure 2. Predicted distribution (yellow) <strong>of</strong> Lymantria mathura<br />
in the contiguous <strong>US</strong>.<br />
Figure 2 illustrates where L. mathura is most likely to encounter a suitable climate<br />
for establishment within the continental <strong>US</strong>. This prediction is based only on the<br />
known geographic distribution <strong>of</strong> the species. Because this forecast is based on<br />
coarse information, areas that are not highlighted on the map may have some<br />
chance <strong>of</strong> supporting populations <strong>of</strong> this exotic species. However, establishment<br />
in these areas is less likely than in those areas that are highlighted. Initial surveys<br />
should be concentrated in the higher risk areas and gradually expanded as needed.<br />
2. Host Specificity/Host Availability. Rating: Low/High. Lymantria mathura is<br />
not host specific; it is a polyphagous pest <strong>of</strong> taxonomically diverse deciduous<br />
trees that are common across the <strong>US</strong> (Appendix B). L. mathura reportedly feeds<br />
on more than 45 genera in 24 families. Table 1 summarizes hosts reported in the<br />
literature. Numerous accounts <strong>of</strong> preferential feeding are reported and vary<br />
widely [see Table 1 below, and Appendix E for more information on host<br />
selection] (Roonwal 1979b, Baranchikov et al. 1995).<br />
CAPS PRA: Lymantria mathura 3
Table 1. Host plants <strong>of</strong> Lymantria mathura:<br />
Hosts<br />
References<br />
alder (Alnus sp.)<br />
(Wallner et al. 1995, Yamazaki and<br />
Sugiura 2004)<br />
apple (Malus sp.) (Mohn 1993, Pucat and Watler 1997,<br />
Zlotina et al. 1998, Gries et al. 1999, CAB<br />
2004, Yamazaki and Sugiura 2004)<br />
apple, Chinese (Malus prunifolia (=“M. (Baranchikov et al. 1995)<br />
“pruniflora”)) 1, 2<br />
arjuna (Terminalia arjuna) 3<br />
(Beeson 1941, Roonwal 1953, Roonwal et<br />
al. 1962, Browne 1968, Roonwal 1979b,<br />
Pucat and Watler 1997, Rosovsky 2001,<br />
CAB 2004)<br />
ash (Fraxinus sp.) (Rosovsky 2001, CAB 2004)<br />
asna (Terminalia elliptica (= Terminalia (Roonwal 1979b)<br />
tomentosa))<br />
Australian red-cedar (Toona ciliata (= (Roonwal 1979b)<br />
Cedrela toona))<br />
beech (Fagus sp.) (Mohn 1993, Pucat and Watler 1997,<br />
Zlotina et al. 1998, Gries et al. 1999,<br />
Rosovsky 2001, CAB 2004)<br />
beech, American (Fagus grandifolia)² (Zlotina et al. 1998)<br />
beech, European (Fagus sylvatica)² (Zlotina et al. 1998)<br />
beleric (Terminalia belerica) (Roonwal 1979b)<br />
Bengal kino (Butea monosperma) (Roonwal 1979b)<br />
birch (Betula sp.)<br />
(Baranchikov et al. 1995, Wallner et al.<br />
1995, Zlotina et al. 1998, Rosovsky 2001,<br />
CAB 2004)<br />
blackboard tree (Alstonia scholaris) (Roonwal 1979b)<br />
Buddhas coconut (Pterygota alata (Roonwal 1979b)<br />
(=Sterculia alata))<br />
“Catania” sp. 1 (Lee and Lee 1996)<br />
Ceylon tea (Elaeodendron<br />
(Roonwal 1979b)<br />
(= “Eeodendron”) glaucum) 1<br />
cherry (Prunus sp.)<br />
(Pucat and Watler 1997, Zlotina et al.<br />
1998, CAB 2004)<br />
cherry, wild Himalayan (Prunus (Roonwal 1979b)<br />
cerasoides (= Prunus puddum))<br />
chestnut (Castanea sp.)<br />
(Zhang 1994, Lee and Lee 1996, Rosovsky<br />
2001, CAB 2004)<br />
chestnut, Chinese hairy (Castanea (Rosovsky 2001, CAB 2004)<br />
mollissima)<br />
chestnut, European (Castanea sativa) (Roonwal 1979b)<br />
china berry tree (Melia azedarach) (Roonwal 1979b)<br />
cottonwood (Populus sp.)<br />
(Baranchikov et al. 1995, Zlotina et al.<br />
1998)<br />
CAPS PRA: Lymantria mathura 4
Hosts<br />
References<br />
crabapple, Manchurian (Malus<br />
(Baranchikov et al. 1995)<br />
mandshurica ([=“mandjurica”)) 1<br />
dhaoda (Anogeissus lalifolia) (Roonwal 1979b)<br />
Douglas-fir (Pseudotsuga menziesii) (Rosovsky 2001, CAB 2004)<br />
duabanga (Duabanga grandiflora (=<br />
Duabanga sonneratioides))<br />
elm (Ulmus sp.)<br />
(Roonwal 1979b)<br />
(Baranchikov et al. 1995, Zlotina et al.<br />
1998)<br />
elm, Japanese (Ulmus davidiana) (Yurchenko and Turova 2002)<br />
fir (Abies sp.) (Rosovsky 2001, CAB 2004)<br />
fir, Manchurian (Abies nephrolepis (Zlotina et al. 1998)<br />
(=“nephroletis”)) 1, 2<br />
Formosan sweetgum (Liquidambar (Mohn 1993, Zhang 1994, Rosovsky 2001,<br />
formosana)<br />
CAB 2004)<br />
Grewia sapinda (Roonwal 1979b)<br />
haldu (Haldina cordifolia (= Adina (Roonwal 1979b)<br />
cordifolia))<br />
hickory (Carya sp.) (Rosovsky 2001, CAB 2004)<br />
hollock (Terminalia myriocarpa) 3 (Beeson 1941, Roonwal 1953, Roonwal et<br />
al. 1962, Browne 1968, Roonwal 1979b,<br />
Pucat and Watler 1997, Rosovsky 2001,<br />
CAB 2004)<br />
Indian banyan (Ficus benghalensis) (Roonwal 1979b)<br />
kadam (Neolamarckia cadamba (= (Browne 1968, Roonwal 1979b, Pucat and<br />
Anthocephalus cadamba))<br />
Watler 1997, Rosovsky 2001, CAB 2004)<br />
kamala (Mallotus philipinensis) (Roonwal 1979b)<br />
larch (Larix sp.)<br />
(Wallner et al. 1995, Rosovsky 2001, CAB<br />
2004)<br />
leechee (Litchi chinensis)<br />
(Singh 1954, Roonwal 1979b, Rosovsky<br />
2001, CAB 2004)<br />
linden, Manchurian (Tilia mandshurica) (Zlotina et al. 1998)<br />
longaan (Dimocarpus longan) (Mohn 1993)<br />
Manchurian nut (Yurchenko and Turova 2002)<br />
mango (Mangifera indica)<br />
(Singh 1954, Browne 1968, Roonwal<br />
1979b, Mohn 1993, Pucat and Watler 1997,<br />
Zlotina et al. 1998, Rosovsky 2001, CAB<br />
2004)<br />
monkey-jack tree (Artocarpus lacucha (Roonwal 1979b)<br />
(= Artocarpus lakoocha))<br />
mulberry, white (Morus alba) (Roonwal 1979b)<br />
oak (Quercus sp.)<br />
(Odell et al. 1992, Mohn 1993, Wallner et<br />
al. 1995, Lee and Lee 1996, Pucat and<br />
Watler 1997, Zlotina et al. 1998, Gries et<br />
al. 1999, Rosovsky 2001, CAB 2004,<br />
Yamazaki and Sugiura 2004)<br />
CAPS PRA: Lymantria mathura 5
Hosts<br />
References<br />
oak, banj (Quercus leucotrichophora (= (Beeson 1941, Roonwal 1953, Roonwal et<br />
Quercus incana)) 3<br />
al. 1962, Browne 1968, Roonwal 1979b,<br />
Pucat and Watler 1997, Rosovsky 2001,<br />
CAB 2004)<br />
oak, chestnut (Quercus prinus) 2 (Zlotina et al. 1998, Gries et al. 1999)<br />
oak, Chinese cork (Quercus variabilis) 2 (Zlotina et al. 1998)<br />
oak, Daimyo (Quercus dentata) (Wileman 1918)<br />
oak, Japanese evergreen (Quercus acuta) (Wileman 1918)<br />
oak, Konara (Quercus serrata (=Q.<br />
glandulifera)) 3<br />
oak, Mongolian (Quercus mongolica)<br />
oak, ring-cup (Quercus glauca) (Funakoshi 2004)<br />
oak, white (Quercus alba) 2 (Zlotina et al. 1998)<br />
(Wileman 1918, Beeson 1941, Roonwal<br />
1953, Roonwal et al. 1962, Browne 1968,<br />
Roonwal 1979b, Pucat and Watler 1997,<br />
Rosovsky 2001, CAB 2004)<br />
(Baranchikov et al. 1995, Zlotina et al.<br />
1998, Rosovsky 2001, Yurchenko and<br />
Turova 2002, CAB 2004)<br />
pear (Pyrus sp.)<br />
(Pucat and Watler 1997, Zlotina et al.<br />
1998, CAB 2004)<br />
pine (Pinus sp.)<br />
(Lee and Lee 1996, Rosovsky 2001, CAB<br />
2004)<br />
pine, Korean (Pinus koraiensis) 2 (Zlotina et al. 1998)<br />
pink-cedar (Acrocarpus fraxinifolius) (Roonwal et al. 1962, Roonwal 1979b)<br />
plum, Java (Syzigium cumini (=Eugenia<br />
jambolana)) 3<br />
(Beeson 1941, Roonwal 1953, Roonwal et<br />
al. 1962, Browne 1968, Roonwal 1979b,<br />
Pucat and Watler 1997, Rosovsky 2001,<br />
CAB 2004)<br />
(Roonwal 1979b)<br />
pongame oil tree (Millettia pinnata<br />
(=Pongamia glabra))<br />
Prunus sp. (Mohn 1993, Yamazaki and Sugiura 2004)<br />
rayana (Aphanamixis polystachya = (Roonwal 1979b)<br />
Amoora (=“Ammora”) rohituka) 1<br />
rose, Japanese (Rosa rugosa) 2 (Baranchikov et al. 1995)<br />
sal tree (Shorea robusta) 3<br />
(Beeson 1941, Roonwal 1953, Roonwal et<br />
al. 1962, Browne 1968, Roonwal 1979a, b,<br />
Pucat and Watler 1997, Rosovsky 2001,<br />
CAB 2004)<br />
sea buckthorn (Hippophae rhamnoides) 2 (Baranchikov et al. 1995)<br />
sumac (Rhus sp.) (Gries et al. 1999)<br />
Terminalia pyrifolia (Roonwal 1979b)<br />
walnut (Juglans sp.) (Rosovsky 2001, CAB 2004)<br />
walnut, Manchurian (Juglans<br />
mandshurica)<br />
waxtree, Japanese (Toxicodendron<br />
succedaneum (=Rhus succedanea))<br />
(Baranchikov et al. 1995, Zlotina et al.<br />
1998)<br />
(Wileman 1918)<br />
CAPS PRA: Lymantria mathura 6
Hosts<br />
willow (Salix sp.)<br />
References<br />
(Zlotina et al. 1998, Rosovsky 2001, CAB<br />
2004)<br />
willow, crack (Salix fragilis) 2 (Baranchikov et al. 1995)<br />
zelkowa (Zelkowa sp.) (Gries et al. 1999)<br />
zelkowa, Japanese (Zelkowa acuminata) (Wileman 1918)<br />
1. Likely mispelling in literature, or unrecognized name.<br />
2. Experimental hosts (Baranchikov et al. 1995, Zlotina et al. 1998)<br />
3. In 1954, following an outbreak in the New Forest Area (Western Sub-Himalayas), 185 tree<br />
species were observed with egg masses, 22 <strong>of</strong> these species were defoloiated, and 6 species<br />
(noted in the table) were heavily defoliated. L. mathura has historically demonstrated food<br />
preferences; depending on host availability some hosts may be chosen or avoided in the<br />
presence <strong>of</strong> more preferred species (Roonwal 1979b).<br />
See Appendix B for maps showing where various hosts are grown in the<br />
continental <strong>US</strong>.<br />
3. Survey Methodology. Rating: High. Several tools are available to assist with<br />
surveys for L. mathura. Pheromone-baited traps are particularly useful for<br />
regional surveys while visual inspections are necessary for conveyances that may<br />
be bringing L. mathura into an area. Inspectors should look for egg masses on<br />
any products originating from infested areas. Egg masses may be deposited on<br />
logs, nursery stock, forest products, or sea containers (Pucat and Watler 1997).<br />
Females prefer to deposit eggs on a rough surface (Roonwal 1979b).<br />
Sex pheromones for L. mathura have been identified and can be used for<br />
detection surveys. Early research (reviewed in Gries et al. 1999) indicated that<br />
males <strong>of</strong> L. mathura were attracted to cis-7,8-epoxy-2-methyloctadecane and 2-<br />
methyl-Z7-octadecene (Odell et al 1992).<br />
Males also demonstrated<br />
elctrophysiological responses to<br />
(Z3,Z6,Z9)-nonadecatriene and (9S,10R)-<br />
9,10-epoxy-Z3,Z6-nonadecadiene in<br />
extracts from abdominal tips <strong>of</strong> L.<br />
mathura females (Oliver et al. 1999).<br />
Subsequent research revealed that major<br />
sex pheromone components include a<br />
blend <strong>of</strong> (9R,10S)-cis-9,10-epoxy-Z3,Z6-<br />
nonadecadiene (named (+)-mathuralure)<br />
and (9S,10R)-cis-9,10-epoxy-Z3,Z6-<br />
nonadecadiene (named (-)-mathuralure) in<br />
a 1:4 ratio (Gries et al. 1999). Neither<br />
component is attractive alone (Gries et al.<br />
1999). Khrimian et al. (2004) explain that<br />
the enantiomer (-)-mathuralure is<br />
equivalent to the compound identified by<br />
Oliver et al. (1999) and provide a detailed<br />
protocol for the synthesis <strong>of</strong> (+)-<br />
Figure 3. Delta trap used<br />
for detecting lymantriids.<br />
[Image from <strong>US</strong>DA APHIS PPQ<br />
Archives, www.forestpests.org]<br />
CAPS PRA: Lymantria mathura 7
mathuralure and (-)-mathuralure in a 4:1 ratio. The pheromone is most effectively<br />
deployed using PVC-coated string dispensers with 64 µg pheromone per cm<br />
(Khrimian et al. 2004). Traps baited with (+)-disparlure will also attract male<br />
L. mathura (Odell et al. 1992).<br />
Pheromone lures have been used with Delta sticky traps (Fig. 3, Gries et al. 1999)<br />
or 3.8-L milk carton traps (Odell et al. 1992). Traps are generally hung 1.5-2 m<br />
[ca. 5-6.5 ft] above ground (Odell et al. 1992, Gries et al. 1999). To improve<br />
diffusion <strong>of</strong> the pheromone, traps have been suspended 0.6 m [2 ft] from the trunk<br />
<strong>of</strong> a tree on wooden stakes nailed to the tree (Odell et al. 1992). For research<br />
purposes, traps were placed 20-25 m apart (Gries et al. 1999), but standard<br />
protocols for detection <strong>of</strong> gypsy moth in uninfested states should be appropriate.<br />
Wallner et al. (1995) evaluated several light sources (e.g., diffuse coated sodium<br />
lamps; phosphor-coated, high-pressure mercury lamps, and blacklight lamps) and<br />
found that L. mathura were most attracted to blacklight. However, light traps are<br />
generally considered ineffective and impractical for regional monitoring <strong>of</strong> this<br />
insect (CAB 2004).<br />
4. Taxonomic Recognition. Rating: High. Lymantria mathura is not likely to be<br />
confused with other lymantrrids, particularly if a specimen is an adult or late<br />
instar larva (EPPO 2005). Eggs or neonates are incredibly difficult to distinguish,<br />
and molecular tools are being developed to aid with identification (Armstrong et<br />
al. 2003). Lymantria mathura might be confused with L. monacha (also exotic,<br />
not known to occur in the <strong>US</strong>) or L. dispar. See Appendix C for a more complete<br />
description <strong>of</strong> the morphology <strong>of</strong> L. mathura.<br />
5. Entry Potential. Rating: Low. Officers with <strong>US</strong>DA-APHIS and <strong>Department</strong> <strong>of</strong><br />
Homeland Security did not report an interception <strong>of</strong> L. mathura at <strong>US</strong> ports <strong>of</strong><br />
entry from 1985-2004 (<strong>US</strong>DA 2005). Two specimens, one L. dispar and one<br />
“Lymantria sp.,” were noted from infested sea containers in Wilmington, NC and<br />
Seattle, WA, respectively (<strong>US</strong>DA 2005). The container infested with L. dispar<br />
may have come from Germany, although the record questions this origin, while<br />
the container with Lymantria sp. came from the Russian Federation. These<br />
records may not reflect the true potential for entry <strong>of</strong> L. mathura. Lymantriids<br />
can be extremely difficult to identify, particularly as eggs and larvae.<br />
Interceptions <strong>of</strong> “Lymantriidae; species <strong>of</strong>” were reported much more frequently.<br />
Unidentified lymantriids were intercepted at least 112 times between 1985-2004<br />
(incomplete records complicate teh accuracy <strong>of</strong> this count) (<strong>US</strong>DA 2005); on<br />
average, 5.6 (±0.7 standard error <strong>of</strong> the mean) interceptions were reported<br />
annually. Most interceptions were associated with permit cargo (38%),<br />
international airline baggage (38%), and general cargo (17%) and were most<br />
commonly reported within the continental <strong>US</strong> from Los Angeles, CA (31%), JFK<br />
International airport, NY (25%), Dallas, TX (4%), Miami, FL (4%), and Long<br />
Beach, CA (4%). These ports are the first points <strong>of</strong> entry for infested material<br />
coming into the <strong>US</strong> and do not necessarily represent the final destination <strong>of</strong><br />
CAPS PRA: Lymantria mathura 8
infested material. Movement <strong>of</strong> potentially infested material is more fully<br />
characterized in the next section.<br />
Remarkably, a substantial number <strong>of</strong> unspecified lymantriid interceptions (<strong>US</strong>DA<br />
2005) were associated with cut flowers, for example Oncidium sp. (24%),<br />
Orchidaceae (7%), Dendrobium sp. (5%), and Astilbe sp. (2%). These plants are<br />
not known hosts for L. mathura, and it is possible that insects were strictly<br />
hitchhikers. However, only ~16% <strong>of</strong> the infested items came from a country<br />
known to have L. mathura. Thus, it seems unlikely that all or even most <strong>of</strong> the<br />
interceptions would have been <strong>of</strong> L. mathura.<br />
Even if all unidentified specimens <strong>of</strong> Lymantriidae had been L. mathura, this<br />
insect would still have an apparent low potential for entry, relative to other exotic<br />
insects. Although we assign a low rating to the potential for entry, we recognize<br />
that not all pathways for the introduction <strong>of</strong> forest pests have been studied with<br />
any detail. Consequently, a great deal <strong>of</strong> uncertainty is associated with the rating.<br />
6. Destination <strong>of</strong> Infested Material. Rating: Medium. When an actionable pest is<br />
intercepted, <strong>of</strong>ficers ask for the intended final destination <strong>of</strong> the conveyance. The<br />
shipments intercepted with L. dispar and “Lymantria sp.” were destined for North<br />
Carolina and Oregon, respectively (<strong>US</strong>DA 2005) Materials infested with<br />
“Lymantriidae” were destined for 17 <strong>of</strong> the contiguous United States. The most<br />
commonly reported destinations were California (38%), New York (24%), Texas<br />
(6%), Florida (6%), Georgia (3%), Illinois (3%), and Massachusetts (3%) (<strong>US</strong>DA<br />
2005). Some portion <strong>of</strong> each state identified as the intended final destination has<br />
a climate and hosts that would be suitable for establishment by L. mathura, yet<br />
probably very few <strong>of</strong> these interceptions involved L. mathura. Consequently,<br />
available data do not permit a confident evaluation <strong>of</strong> this element.<br />
7. Potential Economic Impact. Rating: High. Lymantria mathura larvae are<br />
gregarious defoliators, able to consume whole leaves and sometimes avoid tough<br />
veins in older foliage growth. Larvae may also feed on flowers and tender young<br />
shoots (Browne 1968, Roonwal 1979b). Damage <strong>of</strong> this nature can result in<br />
decline in overall growth and development, a reduction in yield or total crop loss<br />
(fruit crops), or even tree death (Singh 1954, Roonwal 1979b).<br />
In India, L. mathura is an economically important forest pest, which defoliates<br />
Shorea robusta, and several other deciduous forest and fruit tree species [see<br />
‘Host Specificity’]. Roonwal (1953, 1962, 1979b) states that outbreaks are<br />
periodic, and prior to the worst epidemic <strong>of</strong> this pest on record in India during<br />
1953, L. mathura was considered unimportant. In India, this severe outbreak<br />
occurred in Uttar Pradesh in the New Forest area <strong>of</strong> Dehra Dun (approximately<br />
610 m in altitude, in the western sub-Himalayas). The outbreak extended from<br />
the western sub-Himalayas to West Bengal, encompassing several adjacent forest<br />
divisions. In the Russian Far East, there has been only one reported outbreak in<br />
the Primorie region, where losses amounted to hundreds <strong>of</strong> hectares <strong>of</strong> deciduous<br />
CAPS PRA: Lymantria mathura 9
forests (Baranchikov et al. 1995). Damage to chestnut resulted from an outbreak<br />
<strong>of</strong> L. mathura in areas <strong>of</strong> Kyonggi province, Korea (Lee and Lee 1996).<br />
Establishment <strong>of</strong> L. mathura in the <strong>US</strong> could also adversely impact trade. This<br />
insect has been proposed as an A2 quarantine pest in Europe, a status reflecting its<br />
limited presence (EPPO 2005). Potentially infested products within the <strong>US</strong> could<br />
become the focus <strong>of</strong> domestic or international quarantines.<br />
8. Potential Environmental Impact. Rating: High. In general, newly established<br />
species may adversely affect the environment by reducing biodiversity, altering<br />
forest composition, disrupting ecosystem function, jeopardizing endangered or<br />
threatened plants, degrading critical habitat, or stimulating use <strong>of</strong> chemical or<br />
biological controls. Lymantria mathura is likely to affect the environment in<br />
many <strong>of</strong> these ways.<br />
Because L. mathura is known to adversely impact forest productivity and cause<br />
tree mortality with repeated outbreaks, this insect has the potential to directly and<br />
indirectly alter the structure and function <strong>of</strong> forests. Lymantria mathura has the<br />
potential to directly affect forest composition because it has a broad host range<br />
and feeds on foliage <strong>of</strong> primarily deciduous tree species [see ‘Host Specificity’].<br />
Indirect effects stem from the arrival and establishment <strong>of</strong> secondary<br />
organisms/pathogens, such as opportunistic fungi.<br />
Synthetic insecticides are an option, but in many natural settings, complex terrain<br />
limits the feasibility <strong>of</strong> this option, especially over large areas. However, as has<br />
been observed with L. dispar, formulations <strong>of</strong> endotoxin from Bacillus<br />
thuringiensis (e.g, Bt-k) may be applied aerially to localized populations (Myers<br />
and Hosking 2002). Bt is generally considered host specific (Lacey and Siegel<br />
2000), but some exceptions have been noted especially after repeated applications<br />
(Lacey and Siegel 2000, Boulton 2004). Biological control is a much more likely<br />
option (Rosovsky 2001). Previous experience with gypsy moth demonstrates that<br />
predators, parasitoids, and pathogens might be introduced. In previous years,<br />
generalist agents (e.g., Compsilura concinata) were introduced, <strong>of</strong>ten with<br />
significant impacts on non-target species (reviewed in Syrett 2002). Current<br />
protocols for the screening <strong>of</strong> agents limit the likelihood <strong>of</strong> these severe impacts<br />
to non-target species (reviewed in Hoddle and Syrett 2002).<br />
Lymantria mathura may also jeopardize threatened or endangered plants.<br />
Appendix D summarizes state and federally listed threatened or endangered plant<br />
species (<strong>US</strong>DA 2001a) found within plant genera known to be hosts (or potential<br />
hosts) for L. mathura. Plants listed in Appendix D might be suitable hosts for<br />
L. mathura, and thus, could be adversely affected by this insect.<br />
9. Establishment Potential. Rating: Medium. Large areas <strong>of</strong> the United States<br />
are predicted to have a suitable climate for establishment <strong>of</strong> L. mathura, but this<br />
area is only moderate compared with the entire area <strong>of</strong> the country. The host<br />
CAPS PRA: Lymantria mathura 10
status <strong>of</strong> many plants is largely inferred from the genera <strong>of</strong> plants attacked in<br />
Asia. Additional research is needed to confirm the susceptibility <strong>of</strong> <strong>US</strong> species.<br />
If host associations at the genus level continue to hold, several plants in the <strong>US</strong><br />
would be threatened. Many <strong>of</strong> these hosts naturally occur over broad geographic<br />
areas and in relatively high densities. Thus, the potential for establishment seems<br />
high, but our confidence in this assessment is, at best, moderate.<br />
See Appendix E for a more detailed description <strong>of</strong> the biology <strong>of</strong> L. mathura.<br />
References<br />
Anon. 2001. Russian Lymantria Project. <strong>US</strong>FS Forest Health Protection Intermountain<br />
Region. Available on-line at http://www.fs.fed.us/r1-<br />
r4/spf/fhp/publications/weis_final.htm. Accessed 22 August 2005.<br />
Armstrong, K. F., P. McHugh, W. Chinn, and F. E.R. 2003. Tussock moth species<br />
arriving on imported used vehicles determined by DNA analysis. New Zealand<br />
Plant Protection 56: 16-20.<br />
Baranchikov, Y., T. Vshivkova, and M. Montgomery. 1995. Suitability <strong>of</strong> foreign tree<br />
species for Lymantria mathura Moore. pp. 49. In S. L. C. Fosbroke and K. W.<br />
Gottschalk [eds.]. Proceedings, U.S. <strong>Department</strong> <strong>of</strong> <strong>Agriculture</strong> Interagency<br />
<strong>Gypsy</strong> <strong>Moth</strong> Research Forum, 17-20 January, 1995, Annapolis, Maryland. <strong>US</strong>DA<br />
Forest Service General Technical Report NE-213.<br />
Beeson, C. F. C. 1941. The ecology and control <strong>of</strong> the forest insects <strong>of</strong> India and the<br />
neighboring countries. Vasant Press, Dehro Dun.<br />
Boulton, T. J. 2004. Responses <strong>of</strong> nontarget Lepidoptera to Foray 48 B® Bacillus<br />
thuringiensis var. kurstaki on Vancouver Island, British Columbia. Environmental<br />
Toxicology and Chemistry 23: 1297-1304.<br />
Browne, F. G. 1968. Pests and diseases <strong>of</strong> forest plantation trees. Clarendon Press,<br />
Oxford.<br />
CAB. 2004. Crop Protection Compendium. CAB International. Available on-line at<br />
http://www.cabicompendium.org/cpc. Accessed 5 July 2005.<br />
EPPO. 2005. Data sheets on quarantine pests: Lymantria mathura. European and<br />
Mediterranean Plant Protection Organization. Available on-line at<br />
http://www.eppo.org/QUARANTINE/insects/Lymantria_mathura/DSLYMAMA.<br />
pdf. Accessed 29 September 2005.<br />
Funakoshi, S. 2004. Larvae <strong>of</strong> three Lymantria species found on Quercus glauca,<br />
including L. minomonis Matsumura (Lymantriidae). The Japan Heterocerist's<br />
Journal 231: 107-108.<br />
Gninenko, Y. I., and M. Y. Gninenko. 2002. Little known lymantriids <strong>of</strong> the Russian<br />
Far East - potential for movement to other countries <strong>of</strong> the Pacific region. Bulletin<br />
OEPP/EPPO Bulletin 32: 477-480.<br />
Gries, G., R. Gries, P. W. Schaefer, T. Gotoh, and Y. Higashiura. 1999. Sex<br />
pheromone components <strong>of</strong> pink gypsy moth, Lymantria mathura.<br />
Naturwissenschaften 86: 235-238.<br />
Hoddle, M. S., and P. Syrett. 2002. Realizing the Potential <strong>of</strong> Classical Biological<br />
Control, pp. 395-424. In G. J. Hallman and C. P. Schwalbe [eds.], Invasive<br />
CAPS PRA: Lymantria mathura 11
Arthropods in <strong>Agriculture</strong>: Problems and Solutions. Science Publishers, Inc.,<br />
Enfield (NH), <strong>US</strong>A.<br />
Khrimian, A., J. E. Oliver, R. C. Hahn, N. H. Dees, J. White, and V. C. Mastro.<br />
2004. Improved synthesis and deployment <strong>of</strong> (2S,3R)-2-(2Z,5Z-Octadienyl)-3-<br />
nonyloxirane, a pheromone <strong>of</strong> the pink moth, Lymantria mathura. Journal <strong>of</strong><br />
Agricutural and Food Chemistry 52: 2890-2895.<br />
Lacey, L. A., and J. Siegel. 2000. Safety and ecotoxicology <strong>of</strong> entomopathogenic<br />
bacteria, pp. 253-273. In J. F. Charles, A. Delécluse and C. Nielsen-LeRoux<br />
[eds.], Entomopathic Bacteria: From Laboratory to Field Application. Kluwer,<br />
Dordrecht.<br />
Lee, J. H., and H. P. Lee. 1996. Parasites and phenology <strong>of</strong> Lymantria mathura Moore<br />
(Lymantriidae: Lepidoptera) in Kyonggi Province, Korea. Korean Journal <strong>of</strong><br />
Entomology 26: 393-401.<br />
Lewis, F. B., W. E. Wallner, and W. D. Rollinson. 1984. Activity <strong>of</strong> Lymantriid NPVS<br />
from the People's Republic <strong>of</strong> China against North American Lymantria dispar.<br />
Entomophaga 29: 299-302.<br />
Mohn, D. L. 1993. Rosy gypsy moth (Lymantriidae Lymantria mathura - Moore, 1865).<br />
Light Creations. Available on-line at http://www.ccshk.org/DM/butterfly/Lymantrid/Lymantria-mathura.html.<br />
Accessed 22 August<br />
2005.<br />
Moore, F. 1865. On the Lepidopterous insects <strong>of</strong> Bengal. Proceedings <strong>of</strong> the Scientific<br />
Meetings <strong>of</strong> the Zoological Society <strong>of</strong> London: 755-.<br />
Myers, J. H., and G. Hosking. 2002. Eradication, pp. 293-307. In G. J. Hallman and C.<br />
P. Schwalbe [eds.], Invasive Arthropods in <strong>Agriculture</strong>: Problems and Solutions.<br />
Science Publishers, Inc., Enfield (NH), <strong>US</strong>A.<br />
Odell, T. M., C. Xu, P. W. Schaefer, B. A. Leonhardt, D. Yao, and X. Wu. 1992.<br />
Capture <strong>of</strong> gypsy moth, Lymantria dispar (L.), and Lymantria mathura (L.) males<br />
in traps baited with disparlure enantiomers and olefin precursor in the People's<br />
Republic <strong>of</strong> China. Journal <strong>of</strong> Chemical Ecology 18: 2153-2159.<br />
Oliver, J. E., J. C. Dickens, M. A. Zlotina, V. C. Mastro, and G. I. Yurchenko. 1999.<br />
Sex attractant <strong>of</strong> the rosy Russian gypsy moth (Lymantria mathura Moore).<br />
Zeitschrift fur Naturforschung C Biosciences 54: 387-394.<br />
Olson, D. M., E. Dinerstein, E. D. Wikramanayake, N. D. Burgess, G. V. N. Powell,<br />
E. C. Underwood, J. A. D'Amico, I. Itoua, H. E. Strand, J. C. Morrison, C. J.<br />
Loucks, T. F. Allnutt, T. H. Ricketts, Y. Kura, J. F. Lamoreux, W. W.<br />
Wettengel, P. Hedao, and K. R. Kassem. 2001. Terrestrial ecoregions <strong>of</strong> the<br />
world: a new map <strong>of</strong> life on earth. BioScience 51: 933-938.<br />
Pfeifer, T. A., L. M. Humble, M. Ring, and T. A. Grigliatti. 1995. Characterization <strong>of</strong><br />
gypsy moth populations and related species using a nuclear DNA marker. The<br />
Canadian Entomologist 127: 49-58.<br />
Pucat, A. M., and D. E. Watler. 1997. Lymantria mathura Moore: rosy (pink) gypsy<br />
moth. Canadian Food Inspection Agency, Plant Health Risk Assessment Unit.<br />
Available on-line at<br />
http://www.inspection.gc.ca/english/sci/surv/data/lymmate.shtml. Accessed 18<br />
August 2005.<br />
CAPS PRA: Lymantria mathura 12
Roonwal, M. L. 1953. Unusual population eruption <strong>of</strong> the moth, Lymantria mathura<br />
Moore, in autumn. Current Science 22: 384.<br />
Roonwal, M. L., P. N. Chatterjee, and R. S. Thapa. 1962. Experiments on the control<br />
<strong>of</strong> Lymantria mathura Moore (Lepidoptera, Lymantriidae) in the egg and larval<br />
stages in India, with general suggestions for its control. Zeitschrift fur<br />
Angewandte Entomologie 50: 463-475.<br />
Roonwal, M. L. 1979a. The willow, apple, and the sal defoliator. Indian Farming 29: 3-<br />
8.<br />
Roonwal, M. L. 1979b. Field-ecological studies on mass eruption, seasonal life-history,<br />
nocturnal feeding and activity rhythm, and protective behavior and coloration in<br />
the sal defoliator, Lymantria mathura (Lepidoptera: Lymantriidae), in sub-<br />
Himalayan forests. Records <strong>of</strong> the Zoological Survey <strong>of</strong> India 75: 209-236.<br />
Rosovsky, J. 2001. EXFOR Database Pest Report: Lymantria mathura. <strong>US</strong>DA Forest<br />
Service. Available on-line at<br />
http://www.spfnic.fs.fed.us/exfor/data/pestreports.cfm?pestidval=113&langdispla<br />
y=english. Accessed 4 August 2005.<br />
Schintlmeister, A. 2004. The taxonomy <strong>of</strong> the genus Lymantria Hüebner, (1819)<br />
(Lepidoptera: Lymantriidae). Quadrifina 7: 1-248.<br />
Sevastopulo, D. G. 1947. The early stages <strong>of</strong> Indian Lepidoptera. Journal <strong>of</strong> the Bombay<br />
Natural History Society 47: 26-.<br />
Singh, S. M. 1954. A note on serious damage to mango crop by Lymantria mathura<br />
Moore, in Doon Valley. The Indian Journal <strong>of</strong> Horticulture 11: 150.<br />
Syrett, P. 2002. New Restraints on Biological Control, pp. 363-394. In G. J. Hallman and<br />
C. P. Schwalbe [eds.], Invasive Arthropods in <strong>Agriculture</strong>: Problems and<br />
Solutions. Science Publishers, Inc., Enfield (NH), <strong>US</strong>A.<br />
<strong>US</strong>DA. 2001a. The PLANTS database, ver. 3.5. National Plant Data Center, <strong>US</strong>DA-<br />
Natural Resources Conservation Service. Available on-line at<br />
http://plants.usda.gov. Accessed.<br />
<strong>US</strong>DA. 2001b. Pest risk assessment <strong>of</strong> the importation into the United States <strong>of</strong><br />
unprocessed Eucalyptus logs and chips from South America. General Technical<br />
Report FPL-GTR-124. <strong>US</strong> <strong>Department</strong> <strong>of</strong> <strong>Agriculture</strong>, Forest Service, Forest<br />
Products Laboratory., Madison, WI.<br />
<strong>US</strong>DA. 2005. Port Information Network (PIN-309): quarantine status database. <strong>US</strong><br />
<strong>Department</strong> <strong>of</strong> <strong>Agriculture</strong>, Animal and Plant Health Inspection Service, Plant<br />
Protection and Quarantine, Riverdale, MD. Available on-line at restricted access<br />
database. Accessed 2003.<br />
<strong>US</strong>DA NRCS. 2004. The PLANTS database. National Plant Data Center, Baton Rouge,<br />
LA. Available on-line at http://plants.usda.gov. Accessed 11 September 2005.<br />
Wallner, W. E., L. M. Humble, R. E. Levin, Y. N. Baranchikov, and R. T. Carde.<br />
1995. Response <strong>of</strong> adult Lymantriid moths to illumination devices in the Russian<br />
Far East. J. Econ. Entomol. 88: 337-342.<br />
Wileman, A. E. 1918. Notes on Japanese Lepidotera and their larvae: part V. Philippine<br />
Journal <strong>of</strong> Science D Ethnology 13: 151-172.<br />
Yamazaki, K., and S. Sugiura. 2004. Gall-feeding habits in Lepidoptera <strong>of</strong> Japan. III:<br />
two leaf galls. Transactions <strong>of</strong> the Lepidopterological Society <strong>of</strong> Japan 55: 166-<br />
172.<br />
CAPS PRA: Lymantria mathura 13
Yurchenko, G. I., and G. I. Turova. 2002. Biology, behavior and outbreak patterns <strong>of</strong><br />
pink gypsy moth (Lymantria mathura Moore) in the Russian Far East. Chteniya<br />
Pamyati Alekseya Ivanovicha Kurentsova 12: 84-96.<br />
Zhang, B. C. 1994. Index <strong>of</strong> Economically Important Lepidoptera. CAB International,<br />
Wallingford, UK.<br />
Zlotina, M. A., V. C. Mastro, D. E. Leonard, and J. S. Elkinton. 1998. Survival and<br />
development <strong>of</strong> Lymantria mathura (Lepidoptera: Lymantriidae) on North<br />
American, Asian, and European tree species. J. Econ. Entomol. 91: 1162-1166.<br />
Zlotina, M. A., V. C. Mastro, J. S. Elkinton, and D. E. Leonard. 1999. Dispersal<br />
tendencies <strong>of</strong> neonate larvae <strong>of</strong> Lymantria mathura and the Asian form <strong>of</strong><br />
Lymantria dispar (Lepidoptera: Lymantriidae). Environmental Entomology 28:<br />
240-245.<br />
Zolotarenko, G. S., and V. V. Dubatolov. 1998. Lymantriidae collection <strong>of</strong> the Siberian<br />
Zoological Museum. Siberian Zoological Museum. Available on-line at<br />
http://szmn.sbras.ru/Lepidop/Lymantr.htm. Accessed 22 August 2005.<br />
CAPS PRA: Lymantria mathura 14
Appendix A. Geographic distribution and comparison <strong>of</strong> climate zones. To<br />
determine the potential distribution <strong>of</strong> a quarantine pest in the <strong>US</strong>, we first collected<br />
information about the worldwide geographic distribution <strong>of</strong> the species (Table A1).<br />
Using a geographic information system (e.g., ArcView 3.2), we then identified which<br />
biomes (i.e., habitat types), as defined by the World Wildlife Fund (Olson et al. 2001),<br />
occurred within each country or municipality reported An Excel spreadsheet<br />
summarizing the occurrence <strong>of</strong> biomes in each nation or municipality was prepared. The<br />
list was sorted based on the total number <strong>of</strong> biomes that occurred in each<br />
country/municipality. The list was then analyzed to determine the minimum number <strong>of</strong><br />
biomes that could account for the reported worldwide distribution <strong>of</strong> the species.<br />
Countries/municipalities with only one biome were first selected. We then examined<br />
each country/municipality with multiple biomes to determine if at least one <strong>of</strong> its biomes<br />
had been selected. If not, an additional biome was selected that occurred in the greatest<br />
number <strong>of</strong> countries or municipalities that had not yet been accounted for. In the event <strong>of</strong><br />
a tie, the biome that was reported more frequently from the entire species’ distribution<br />
was selected. The process <strong>of</strong> selecting additional biomes continued until at least one<br />
biome was selected for each country. Finally, the set <strong>of</strong> selected biomes was compared to<br />
only those that occur in the <strong>US</strong>.<br />
Table A1. Reported geographic distribution <strong>of</strong> Lymantria mathura:<br />
Locations<br />
References<br />
Asia (southeast) (Roonwal 1979b, Baranchikov et al. 1995)<br />
Bangladesh (Rosovsky 2001, CAB 2004)<br />
China (Wileman 1918, Zhang 1994, Pucat and Watler 1997,<br />
Gries et al. 1999, Rosovsky 2001, Khrimian et al. 2004)<br />
China (Beijing) (Lewis et al. 1984)<br />
China (Dunhua) (Lewis et al. 1984, Odell et al. 1992)<br />
China (Heilogjiang Province) (CAB 2004)<br />
China (Heilongjiang Province - Menjiagang) (Lewis et al. 1984, Odell et al. 1992)<br />
China (Hong Kong) (Mohn 1993, Rosovsky 2001, CAB 2004)<br />
China (Jiaohe) (Lewis et al. 1984)<br />
China (Jingpo Hu) (Lewis et al. 1984)<br />
China (Manchuria) (Wileman 1918, Pucat and Watler 1997)<br />
China (north) (Zlotina et al. 1998)<br />
China (northeast) (Wallner et al. 1995, Rosovsky 2001)<br />
China (Yabuli) (Lewis et al. 1984)<br />
China (Yunnan) (Schintlmeister 2004)<br />
India<br />
(Wileman 1918, Browne 1968, Zhang 1994, Pucat and<br />
Watler 1997, Zlotina et al. 1998, Gries et al. 1999,<br />
Rosovsky 2001, CAB 2004, Khrimian et al. 2004)<br />
India (Assam) (Beeson 1941, Roonwal 1953, Roonwal et al. 1962,<br />
Roonwal 1979a, b)<br />
India (Bengal - northeast) (Schintlmeister 2004)<br />
India (Darjeeling District - Tukdah) (Sevastopulo 1947)<br />
India (Darjeeling District) (Schintlmeister 2004)<br />
India (north)<br />
(Wileman 1918, Beeson 1941, Roonwal 1953, Roonwal et<br />
al. 1962, Schintlmeister 2004)<br />
CAPS PRA: Lymantria mathura 15
Locations<br />
References<br />
India (northwest) (Baranchikov et al. 1995)<br />
India (Uttar Pradesh) (Roonwal et al. 1962, Roonwal 1979a, b)<br />
India (Uttaranchal - Dehra Dun) (Roonwal 1953, Singh 1954, Roonwal 1979b)<br />
India (Uttaranchal - Doon Valley) (Roonwal 1979b)<br />
India (West Bengal - Buxa Forest Division) (Roonwal 1979b)<br />
Japan (Wileman 1918, Zhang 1994, Pucat and Watler 1997,<br />
Zlotina et al. 1998, Gries et al. 1999, Rosovsky 2001,<br />
Khrimian et al. 2004, Schintlmeister 2004)<br />
Japan (central) (Funakoshi 2004)<br />
Japan (Higo Province - Kosadake-machi) (Wileman 1918)<br />
Japan (Hokkaido Prefecture - Bibai) (Gries et al. 1999)<br />
Japan (Hokkaido Prefecture - Jozankei) (Wileman 1918)<br />
Japan (Honshu) (Schintlmeister 2004)<br />
Japan (Iwate Prefecture - Morioka) (Gries et al. 1999)<br />
Japan (Iyo Province - Ohoki) (Wileman 1918)<br />
Japan (Kishu Province - Koyasan) (Wileman 1918)<br />
Japan (Kyushu) (Wileman 1918)<br />
Japan (Musashi Province - Kawai, Dzushi) (Wileman 1918)<br />
Japan (Musashi Province - Tokyo) (Wileman 1918)<br />
Japan (Nagahama) (Schintlmeister 2004)<br />
Japan (Osaka Prefecture - Sakai City) (Yamazaki and Sugiura 2004)<br />
Japan (Ryukyu Islands) (Wileman 1918)<br />
Japan (Settsu Province - Kobe) (Wileman 1918)<br />
Japan (Shinano Province - Karuizawa) (Wileman 1918)<br />
Japan (Yokohama) (Wileman 1918)<br />
Kashmir (Wileman 1918)<br />
Korea (Wileman 1918, Rosovsky 2001)<br />
Korea (Kyonggi Province - Jinjung-Ri) (Lee and Lee 1996)<br />
Korea (Kyonggi Province - Songchon-Ri) (Lee and Lee 1996)<br />
Korea, Republic <strong>of</strong> (CAB 2004)<br />
Kurile Islands (Wileman 1918)<br />
Myanmar (formerly Burma) (Roonwal 1979b)<br />
Pakistan (Browne 1968, Pucat and Watler 1997, Rosovsky 2001)<br />
Russia (Khrimian et al. 2004)<br />
Russia (Amur) (Zolotarenko and Dubatolov 1998)<br />
Russia (eastern Siberia) (Wileman 1918)<br />
Russia (eastern) (Gries et al. 1999)<br />
Russia (Far East - Yakolevka) (Pfeifer et al. 1995)<br />
Russia (Far East)<br />
(Baranchikov et al. 1995, Wallner et al. 1995, Zlotina et<br />
al. 1998, Anon. 2001, Rosovsky 2001, CAB 2004)<br />
Russia (Nakhodka) (Anon. 2001)<br />
Russia (Primorsky Krai - Kavalerovo) (Zlotina et al. 1998, Zlotina et al. 1999)<br />
Russia (Primorsky Krai - Mineralni) (Wallner et al. 1995)<br />
Russia (Primorye Region - Barabash) (Oliver et al. 1999)<br />
CAPS PRA: Lymantria mathura 16
Locations<br />
Russia (Primorye Region)<br />
References<br />
(Baranchikov et al. 1995, Zolotarenko and Dubatolov<br />
1998)<br />
Russia (Siberia) (CAB 2004)<br />
Russia (Vladivostok) (Oliver et al. 1999, Anon. 2001, Rosovsky 2001)<br />
Russia (Vostochny) (Anon. 2001)<br />
Taiwan (Zhang 1994, Pucat and Watler 1997, Gries et al. 1999,<br />
Rosovsky 2001, CAB 2004, Schintlmeister 2004)<br />
Taiwan (Puli-Wushe) (Schintlmeister 2004)<br />
temperate broadleaf and mixed forest 2 (Schintlmeister 2004)<br />
temperate coniferous forest 2 (Schintlmeister 2004)<br />
tropical and subtropical dry broadleaf forest 2 (Schintlmeister 2004)<br />
tropical and subtropical moist broadleaf (Schintlmeister 2004)<br />
forest 2<br />
United States <strong>of</strong> America 1 (N. America; west (Baranchikov et al. 1995, CAB 2004)<br />
coast ports)<br />
1. Intercepted but not established (Baranchikov et al. 1995, CAB 2004).<br />
2. Refer to map by Schintlmeister for general locations; no scale provided (Schintlmeister 2004).<br />
CAPS PRA: Lymantria mathura 17
Appendix B. Host distribution (partial) for Lymantria<br />
mathura. The host status <strong>of</strong> all species has not<br />
necessarily been confirmed.<br />
Map 1. Apple (Malus domestica)<br />
Little, Atlas <strong>of</strong> United States Trees, 2004<br />
climchange.cr.usgs.gov/data/atlas/little/<br />
Map 2. American beech (Fagus grandifolia)<br />
Map 3. Cherry (Prunus avium)<br />
Little, Atlas <strong>of</strong> United States Trees, 2004<br />
climchange.cr.usgs.gov/data/atlas/little/<br />
Map 4. Douglas fir (Pseudotsuga menziesii)<br />
Map 5. Mango (Mangifera indica)<br />
CAPS PRA: Lymantria mathura 18
Little, Atlas <strong>of</strong> United States Trees, 2004<br />
climchange.cr.usgs.gov/data/atlas/little/<br />
Little, Atlas <strong>of</strong> United States Trees, 2004<br />
climchange.cr.usgs.gov/data/atlas/little/<br />
Map 6. Banj oak (Quercus leucotrichophora)<br />
Map 7. Chestnut oak (Quercus prinus)<br />
Little, Atlas <strong>of</strong> United States Trees, 2004<br />
climchange.cr.usgs.gov/data/atlas/little/<br />
Map 8. White oak (Quercus alba)<br />
Map 9. Pear (Pyrus communis)<br />
CAPS PRA: Lymantria mathura 19
Appendix C. Taxonomy and morphology <strong>of</strong> Lymantria mathura<br />
Synonyms<br />
Portheria mathura (Moore)<br />
Ocneria mathura (Moore)<br />
Lymantria aurora Butler<br />
Lymantria fusca Leech<br />
Lymantria mathura aurora Butler<br />
Diagnostic features<br />
For complete accuracy, the following morphological descriptions <strong>of</strong> L. mathura are<br />
quoted from Moore (1865) and Roonwal (1979b).<br />
Lymantria mathura<br />
“Lymantria mathura Moore (Lepidoptera : Lymantriidae) is a moderate sized moth...<br />
There is marked sexual dimorphism in size and colour. The male is smaller (wing<br />
expanse male: 35-50mm; female: 75-95mm), with the forewings brown and hindwings<br />
yellow. In females the forewings are white with dark markings, and the hindwings<br />
pink...”(Roonwal 1979b).<br />
Male<br />
“Upperside-fore wing greyish white, markings brown, with pale-brown interspaces; with<br />
two or three black and yellow spots at the base; two transverse subbasal irregular lines,<br />
between which is a broad band; a round spot within the cell and a blackish curved streak<br />
at its end; three transverse discal lunulated bands, the first broad, the others narrow; a<br />
marginal row <strong>of</strong> spots: hind wing dull yellow, with a blackish discal spot, narrow<br />
submarginal maculated band, and a marginal row <strong>of</strong> small spots. Underside dull yellow,<br />
suffused with pale brown between the veins, with darker-brown discal and marginal<br />
spots. Thorax white, with yellow and black spots. Abdomen yellow, tuft white, with<br />
dorsal, lateral, and a row beneath <strong>of</strong> black spots. Head at the sides, palpi in front, and legs<br />
yellow; palpi above and at the sides, and spots on the legs, black. Antennae brown.<br />
Expanse 2¼ inches” (Moore 1865).<br />
“Egg-masses and covering hairs”<br />
Egg masses are laid from ground-level up to about 18 m (60 ft.) <strong>of</strong> the trunk, but are most<br />
dense between the levels <strong>of</strong> 0.5 to 5 m. They are flat, <strong>of</strong> an ovoid-elongate or other<br />
shape, with irregular edges, and vary in extent from about 0.5 x 1 cm to 6 x 15 cm. From<br />
a distance the egg masses are visible as characteristic white, fluffy patches against the<br />
dark-coloured bark. Each egg-mass contains about 50 to 1,200 or more eggs which are<br />
laid 2 to 4 layers deep directly on the bark. An egg-mass is covered over with a nearly<br />
one-millimetre, white thick felt-like covering composed <strong>of</strong> long, white, silken hairs. (...<br />
these hairs are shed by the female from the anal tuft. ... ) The hairs are about 800-1200 µ<br />
long and 3.1-6.2 µ in diameter; one end is knob-like, the other pointed; a few such hairs<br />
are also mixed with the eggs. Freshly laid eggs are rounded, have a flat base, the<br />
CAPS PRA: Lymantria mathura 20
maximum and minimum diameters varying from 1 .13-1.19 mm and 0.86-0.92 mm<br />
respectively ” (Roonwal 1979b).<br />
“Egg-mass after hatching”<br />
After the majority <strong>of</strong> eggs have hatched, an egg mass presents a changed appearance.<br />
Firstly, the hair-covering which has hitherto (for several months in the case <strong>of</strong> the<br />
overwintering eggs) remained pure white, now becomes dull-coloured, a dirty cream,<br />
and, in a few cases, with irregular patches <strong>of</strong> pale buff. Secondly, the hair covering<br />
is pierced by numerous rounded holes <strong>of</strong> varying diameters (c. 0.5-3 mm) through which<br />
the newly hatched larvae have escaped. Beneath the thin, hole-pierced, hairy covering,<br />
there is a flat, hollow space containing the remnants <strong>of</strong> eggshells and a few remaining<br />
eggs which have not yet hatched” (Roonwal 1979b).<br />
Larvae<br />
“Three main colour forms are found in mature caterpillars, the following proportions<br />
being noticed in 1,613 caterpillars examined: grey-white 66 %, intermediate 11 %, and<br />
blackish brown 23 %. The details <strong>of</strong> colour are described below briefly.<br />
Form I (Grey-white) : Ground colour dirty white tinged with grey. Dorsal : Head white<br />
with numerous black or brown spots; frons with a longitudinal median black streak; rest<br />
<strong>of</strong> body grey-white, with numerous fine dots forming paired patches. A transverse<br />
yellow-brown streak present between pro- and mesothorax, and another in middle <strong>of</strong><br />
metathorax: abdominal warts blackish; paired lateral papules on abdomen white, with<br />
tufts <strong>of</strong> long white and brown hairs. Long pencil-like plumes <strong>of</strong> hairs on head and on, end<br />
<strong>of</strong> abdomen black. Ventral: Brownish pink; legs and prolegs brown, the latter with a<br />
black patch externally.<br />
Form II (Intermediate): Dorsal: Ground colour pale brown, with a median white patch on<br />
abdominal terga 4 and 5. Ventral: As in Form I.<br />
Form III (Blackish brown): Dorsal: Ground-colour dark brown to almost black; numerous<br />
black spots visible in brown larvae but merged with ground-colour in darker ones; several<br />
small white dots present on abdominal terga 4 to the last, and large white patches on terga<br />
4-6. Ventral: Ashy, suffused with a little pink in the median parts; rest as in Form I.<br />
In the masses <strong>of</strong> caterpillars on tree trunks the various colour types are mixed on<br />
individual trees; this fact has a protective value by making detection by enemies difficult”<br />
(Roonwal 1979b).<br />
“The size ... characteristics <strong>of</strong> the six larval stages are given below briefly...<br />
Stage I. Length 3 mm; head-width 0.5 mm. Generally black dorsally; meso- and<br />
metathorax and segment 5 <strong>of</strong> abdomen brown; legs black; prolegs pale brown with a<br />
black patch externally.<br />
CAPS PRA: Lymantria mathura 21
Stage II. Length 5 mm; head-width 0.7 mm. Generally black dorsally; meso- and<br />
metathorax greyish; last abdominal segment pale brown with blackish tinge; rest as in<br />
Stage I.<br />
Stage III. Length 13 mm; head-width 1.5 mm. Head brown; body black above, paler<br />
below; thoracic terga with yellow-brown spots; legs black, prolegs brown with a black<br />
external patch.<br />
Stage IV. Length 20 mm; head-width 2.5 mm. Head above either black (brown distally)<br />
or pale green with black dots; sides brown; body black with white warts; meso- and<br />
metathorax with brown stripes anteriorly; legs and prolegs as in Stage III.<br />
Stage V. Length 30-40 mm; head-width 3.5 mm. Head above brown to grey, speckled<br />
with black; body black with many minute white spots; pro- and mesothorax with a<br />
transverse brown streak at the distal edge; ninth abdominal segment with a pair <strong>of</strong><br />
prominent dorsal white spots; legs and prolegs reddish brown, the latter with a<br />
large black patch externally.<br />
Stage VI. Length 60-85 mm; head-width 5-6 mm. With sexual dimorphism, females<br />
being longer (males: 60-65 mm, females: 70-85 mm). Colour pattern similar to Stage V,<br />
but in ground-pattern three types recognizable, viz., grey-white, blackish-brown and<br />
intermediate (vide infra). Older larvae well “camouflaged” against tree trunks” (Roonwal<br />
1979b).<br />
The pupa<br />
“The pupa is <strong>of</strong> the ‘obtect adecticus type,’ and the appendages are firmly soldered to the<br />
body. It is buff to dark brown, about 20-36 mm long, and shows sexual dimorphism; the<br />
female pupa is paler, larger and heavier than the male, as follows:<br />
Female: Buff to pale brown. Length (including hair tufts) 30-36 mm; maximum<br />
width 10-14 mm. Weight 0.88 gm (average <strong>of</strong> 18 pupae).<br />
Male: Very dark chocolate brown, Length (including hair tufts) 15-25 mm;<br />
maximum width 6-8 mm. Weight 0.14 gm (average <strong>of</strong> 53 pupae)” (Roonwal<br />
1979b).<br />
CAPS PRA: Lymantria mathura 22
Appendix D. Threatened or endangered plants potentially affected by Lymantria mathura.<br />
Lymantria mathura has the potential to adversely affect threatened and endangered plant species. Because L. mathura is not known to<br />
be established in the <strong>US</strong> and threatened and endangered plant species do not occur outside the <strong>US</strong>, it is not possible to confirm the host<br />
status <strong>of</strong> these rare plants from the scientific literature. From available host records, L. mathura is known to feed on species within the<br />
following families: Anacardiaceae, Apocynaceae, Betulaceae, Celastraceae, Combretaceae, Dipterocarpaceae, Elaeagnaceae,<br />
Euphorbiaceae, Fabaceae, Fagaceae, Hamamelidaceae, Juglandaceae, Lythraceae, Malvaceae, Meliaceae, Moraceae, Myrtaceae,<br />
Oleaceae, Pinaceae, Roseaceae, Rubiaceae, Salicaceae, Sapindaceae, Tiliaceae, and Ulmaceae. From these host records, we infer that<br />
threatened or endangered plant species which are closely related to known host plants might also be suitable hosts (Table D1) (<strong>US</strong>DA<br />
NRCS 2004). For our purposes closely related plant species belong to the same genus.<br />
Table D1: Threatened and endangered plants in the conterminous U.S. that are potential hosts for<br />
Lymantria mathura.<br />
Documented/Reported Hosts<br />
Threatened and/or Endangered Plant Protected Status 1<br />
Scientific Name Common Name Federal State<br />
Abies sp., A. nephrolepis Abies balsamea balsam fir CT (E)<br />
A. fraseri Fraser fir TN (T)<br />
Alnus sp. Alnus incana ssp. rugosa speckled alder IL (E)<br />
A. viridis ssp. crispa mountain alder MA (T)<br />
PA (E)<br />
Betula sp. Betula alleghaniensis yellow birch IL (E)<br />
B. minor dwarf white birch ME (E)<br />
NY (E)<br />
B. nana [= B. glandulosa] dwarf birch ME (E)<br />
NH (T)<br />
NY (E)<br />
B. nigra river birch NH (T)<br />
B. papyrifera var. cordifolia mountain paper birch TN (E)<br />
B. populifolia gray birch IL (E)<br />
CAPS PRA: Lymantria mathura 23
Table D1: Threatened and endangered plants in the conterminous U.S. that are potential hosts for<br />
Lymantria mathura.<br />
Documented/Reported Hosts<br />
Threatened and/or Endangered Plant Protected Status 1<br />
Scientific Name Common Name Federal State<br />
B. pumila bog birch IA (T)<br />
MA (E)<br />
NH (E)<br />
NY (T)<br />
OH (T)<br />
B. pumila var. glandulifera bog birch VT (E)<br />
B. uber Virginia roundleaf birch T VA (E)<br />
Carya sp. Carya aquatica water hickory KY (T)<br />
C. cordiformis bitternut hickory ME (E)<br />
C. laciniosa shellbark hickory MD (E)<br />
NY (T)<br />
C. myristiciformis nutmeg hickory NC (T)<br />
C. pallida sand hickory AR (T)<br />
IL (E)<br />
IN (T)<br />
C. texana black hickory IN (E)<br />
Castanea sp., C. mollissima,<br />
C. sativa<br />
Castanea dentata American chestnut KY (E)<br />
MI (E)<br />
C. pumila chinkapin KY (T)<br />
NJ (E)<br />
Fraxinus sp. Fraxinus pr<strong>of</strong>unda pumpkin ash MI (T)<br />
NJ (E)<br />
PA (E)<br />
F. quadrangulata blue ash IA (T)<br />
WI (T)<br />
Juglans sp., J. mandshurica Juglans cinerea butternut TN (T)<br />
Larix sp. Larix laricina tamarack IL (T)<br />
MD (E)<br />
CAPS PRA: Lymantria mathura 24
Table D1: Threatened and endangered plants in the conterminous U.S. that are potential hosts for<br />
Lymantria mathura.<br />
Documented/Reported Hosts<br />
Malus sp., M. mandshurica,<br />
M. prunifolia<br />
Threatened and/or Endangered Plant Protected Status 1<br />
Scientific Name Common Name Federal State<br />
Malus angustifolia southern crabapple FL (T)<br />
IL (E)<br />
M. glaucescens Dunbar crabapple NY (E)<br />
Morus alba Morus rubra red mulberry CT (E)<br />
MA (E)<br />
MI (T)<br />
VT (T)<br />
Pinus sp., P. koraiensis Pinus banksiana jack pine IL (E)<br />
NH (T)<br />
VT (T)<br />
P. echinata shortleaf pine IL (E)<br />
P. pungens Table Mountain pine NJ (E)<br />
P. resinosa red pine CT (E)<br />
IL (E)<br />
NJ (E)<br />
P. virginiana Virginia pine NY (E)<br />
Populus sp. Populus balsamifera balsam poplar IL (E)<br />
OH (E)<br />
PA (E)<br />
P. heterophylla swamp cottonwood CT (E)<br />
MA (E)<br />
MI (E)<br />
NY (T)<br />
Prunus sp., P. cerasoides Prunus alleghaniensis Allegheny plum MD (T)<br />
NJ (E)<br />
PA (T)<br />
P. americana American plum NH (T)<br />
VT (T)<br />
P. angustifolia Chickasaw plum NJ (E)<br />
CAPS PRA: Lymantria mathura 25
Table D1: Threatened and endangered plants in the conterminous U.S. that are potential hosts for<br />
Lymantria mathura.<br />
Documented/Reported Hosts<br />
Quercus sp., Q. acuta, Q. alba,<br />
Q. dentata, Q. glauca,<br />
Q. leucotrichophora,<br />
Q. mongolica, Q. prinus,<br />
Q. serrata, Q. variabilis<br />
Threatened and/or Endangered Plant Protected Status 1<br />
Scientific Name Common Name Federal State<br />
Prunus geniculata scrub plum E FL (E)<br />
P. maritima beach plum MD (E)<br />
ME (E)<br />
PA (E)<br />
P. maritima var. gravesii Grave’s plum CT (E)<br />
P. nigra Canadian plum IA (E)<br />
P. pumila sandcherry AR (T)<br />
TN (T)<br />
P. pumila var. depressa eastern sandcherry NY (T)<br />
P. pumila var. pumila Great Lakes sandcherry NY (E)<br />
P. pumila var. susquehanae [= P. pumilla var. Sesquehana sandcherry OH (T)<br />
cuneata]<br />
Quercus acerifolia mapleleaf oak AR (T)<br />
Q. bicolor swamp white oak ME (T)<br />
Q. coccinea scarlet oak ME (E)<br />
Q. falcata southern red oak OH (T)<br />
PA (E)<br />
Q. hinckleyi Hinckley oak T TX (T)<br />
Q. ilicifolia bear oak VT (E)<br />
Q. imbricaria shingle oak NJ (E)<br />
Q. lyrata overcup oak NJ (E)<br />
Quercus macrocarpa bur oak CT (E)<br />
Q. muehlenbergii [= Q. prinoides] chinkapin oak IN (E)<br />
Q. nigra water oak NJ (E)<br />
Q. oglethorpensis Oglethorpe oak GA (T)<br />
CAPS PRA: Lymantria mathura 26
Table D1: Threatened and endangered plants in the conterminous U.S. that are potential hosts for<br />
Lymantria mathura.<br />
Documented/Reported Hosts<br />
Threatened and/or Endangered Plant Protected Status 1<br />
Scientific Name Common Name Federal State<br />
Quercus phellos willow oak IL (T)<br />
NY (E)<br />
PA (E)<br />
Q. prinus [= Q. montana] chestnut oak IL (T)<br />
ME (T)<br />
Q. shumardii Shumard’s oak MD (T)<br />
PA (E)<br />
Q. sinuata var. sinuata [= Q. durandii] bastard oak AR (T)<br />
Q. texana [= Q. nuttallii] Texas red oak IL (E)<br />
Rosa rugosa Rosa acicularis prickly rose IA (E)<br />
IL (E)<br />
MA (E)<br />
NH (E)<br />
VT (E)<br />
R. acicularis ssp. sayi prickly rose NY (E)<br />
R. blanda smooth rose MD (E)<br />
OH (T)<br />
R. minutifolia Baja rose CA (E)<br />
R. nitida shining rose NY (E)<br />
Rhus sp. Rhus aromatica var. arenaria fragrant sumac IN (T)<br />
R. michauxii false poison sumac E FL (E)<br />
GA (E)<br />
NC (E)<br />
Salix sp., S. fragilis Salix arctophila northern willow ME (E)<br />
S. argyrocarpa Labrador willow ME (E)<br />
NH (T)<br />
S. bebbiana Bebb willow MD (E)<br />
CAPS PRA: Lymantria mathura 27
Table D1: Threatened and endangered plants in the conterminous U.S. that are potential hosts for<br />
Lymantria mathura.<br />
Documented/Reported Hosts<br />
Threatened and/or Endangered Plant Protected Status 1<br />
Scientific Name Common Name Federal State<br />
Salix candida sageleaf willow ME (T)<br />
OH (T)<br />
PA (E)<br />
S. caroliniana costal plain willow OH (T)<br />
PA (E)<br />
S. cordata [= S. syrticola] heartleaf willow IL (E)<br />
NY (E)<br />
WI (E)<br />
S. eriocephala [= S. cordata] Missouri River willow FL (E)<br />
IN (T)<br />
S. exigua narrowleaf willow CT (T)<br />
MD (E)<br />
S. floridana Florida willow FL (E)<br />
GA (E)<br />
S. herbacea snowbed willow ME (T)<br />
NH (T)<br />
NY (E)<br />
S. interior sandbar willow ME (E)<br />
S. lucida shining willow IA (T)<br />
MD (E)<br />
S. myricoides bayberry willow ME (E)<br />
S. pedicellaris bog willow CT (E)<br />
IA (T)<br />
NJ (E)<br />
OH (E)<br />
PA (E)<br />
S. pellita satiny willow NH (T)<br />
WI (E)<br />
CAPS PRA: Lymantria mathura 28
Table D1: Threatened and endangered plants in the conterminous U.S. that are potential hosts for<br />
Lymantria mathura.<br />
Documented/Reported Hosts<br />
Threatened and/or Endangered Plant Protected Status 1<br />
Scientific Name Common Name Federal State<br />
Salix petiolaris meadow willow OH (T)<br />
PA (E)<br />
S. planifolia diamondleaf willow ME (T)<br />
MI (T)<br />
NH (T)<br />
VT (T)<br />
WI (T)<br />
S. pyrifolia balsam willow NY (T)<br />
S. sericea silky willow AR (E)<br />
S. serissima autumn willow IL (E)<br />
IN (T)<br />
PA (T)<br />
S. sessilifolia northwest sandbar<br />
WA (T)<br />
willow<br />
S. uva-ursi bearberry willow ME (T)<br />
NY (T)<br />
VT (E)<br />
Tilia mandshurica Tilia americana var. heterophylla [= T. American basswood IL (E)<br />
heterophylla]<br />
Toxicodendron succedaneum Toxicodendron rydbergii western poison ivy OH (E)<br />
T. vernix poison sumac KY (E)<br />
Ulmus sp., U. davidiana Ulmus thomasii rock elm IL (E)<br />
NY (T)<br />
OH (T)<br />
1. E= Endangered; T=Threatened.<br />
CAPS PRA: Lymantria mathura 29
Appendix E. Biology <strong>of</strong> Lymantria mathura<br />
Population phenology<br />
Lymantria mathura is bivoltine (Beeson 1941, Browne 1968, Roonwal 1979b,<br />
Baranchikov et al. 1995, Lee and Lee 1996). Roonwal (1953, 1962, 1979b) provides a<br />
thorough review <strong>of</strong> the phenology and behavior <strong>of</strong> L. mathura following the most<br />
damaging outbreak on record for this pest in 1953 in Uttar Pradesh, India. The first<br />
generation occurs between April and October. Eggs are laid between mid-April and mid-<br />
June and hatch in 3-4 weeks. Larvae and pupae occur from early June to late September,<br />
and from late July to late October, respectively. In the second or overwintering<br />
generation, eggs are laid between early September to mid-October, and embryos are<br />
developed within 6 weeks. This generation overwinters as developed embryos, and eggs<br />
hatch in the spring between February and early April, depending on temperature.<br />
Incubation requires160-178 days, or a shorter duration in warmer temperatures. Of 426<br />
field-collected pupae in the overwintering generation in the New Forest area, 58% were<br />
male. The pupal stage occurs within 10-11 days for this generation.<br />
In outbreak years, L. mathura tends to lay eggs on many tree species, including nonhosts.<br />
Lymantria mathura eggs were laid on 185 different host species, and <strong>of</strong> these, 22<br />
tree species were later defoliated by feeding larvae, and 6 species were heavily defoliated<br />
[see ‘Host Specificity’]. Lymantria mathura has historically demonstrated food<br />
preferences, but these preferences depend on which hosts are available (Roonwal 1979b,<br />
Baranchikov et al. 1995). The selection <strong>of</strong> a location for egg deposition may also depend<br />
on the presence or density <strong>of</strong> other egg masses, host preference, and the extent <strong>of</strong> feeding<br />
that has already occurred on a host (Roonwal 1979b).<br />
Stage specific biology<br />
Several papers discuss periods <strong>of</strong> development for L. mathura and related species,<br />
however there are no known temperature developmental thresholds for L. mathura in<br />
published literature obtained to date (Anon. 2001).<br />
Adult<br />
Flight has been observed between 1-3 a.m. in far east Russia. Flight activity is not well<br />
known for this species, but is thought to coincide with peak flight activity <strong>of</strong> two closely<br />
related species, L. dispar and L. monacha (Anon. 2001). Males are scarcely seen and die<br />
about a week before females. Females congregate in groups <strong>of</strong> 6 or more near the egg<br />
masses and become inactive after laying eggs. They do not fly or feed before dying<br />
(Roonwal 1979b).<br />
Egg<br />
Between 50-1,200 eggs are laid in white, distinctive silky hair-covered masses on trunks<br />
and large branches <strong>of</strong> deciduous hosts (Browne 1968, Roonwal 1979b). Eggs are laid<br />
from the base <strong>of</strong> a tree trunk to a height <strong>of</strong> about 18 m (60 ft.), and most egg masses tend<br />
to occur at a height between 0.5 to 5 m (Roonwal 1979b).<br />
CAPS PRA: Lymantria mathura 30
During an outbreak in the New Forest area <strong>of</strong> the western sub-Himalayas in 1953-54,<br />
between 1-223 egg masses were reported on 405 trees over an area <strong>of</strong> 10 km 2 . After eggs<br />
hatch, the egg mass becomes darker in color. The group <strong>of</strong> newly hatched larvae remain<br />
near the hair-covered mass for 2-3 weeks. It is not known whether the larvae recieve<br />
some nutritive benefit from the mass prior to feeding on foliage (Roonwal 1979b).<br />
Larva<br />
There are 6 larval stages or instars and 5 molts. Larvae are gregarious defoliators,<br />
devouring entire leaves, sometimes avoiding tough veins in older growth. Larvae may<br />
also feed on flowers and tender young shoots (Browne 1968, Roonwal 1979b). Early<br />
stages <strong>of</strong> larvae move about more freely than later instars (Roonwal 1979b). Late instar<br />
larvae have demonstrated regular periods <strong>of</strong> daily dispersal activity (Roonwal 1979b,<br />
Zlotina et al. 1999). Roonwal (1979b) observed that caterpillars remained still and at rest<br />
for much <strong>of</strong> the day and migrated to the tree crown to feed at night. Prior to dusk,<br />
caterpillars exhibited a characteristic twisting body movement, then crawled to the tree<br />
crown at a rate <strong>of</strong> approximately 65.5 cm/min. Feeding occurs from dusk to near dawn,<br />
followed by a rapid descent to the trunk. Larval densities may approach can average was<br />
1,338 / tree (1,140-1,671), or an average <strong>of</strong> 629 larvae (510-836) per square meter<br />
(Roonwal 1979b). Density on the host trunk reached a maximum at 5 PM, just prior to<br />
the evening migration to the crown. Early instar L. mathura larvae are thought to possess<br />
the ability to disperse in a similar manner as other related species, by dropping on a<br />
trailing silk thread and utilizing air and wind currents to “balloon” to other locations<br />
(Zlotina et al. 1999).<br />
Zlotina et al. (1999) studied dispersal rates, settling velocities and diel dispersal activity<br />
<strong>of</strong> L. mathura and L. dispar larvae. Lymantria mathura larvae showed a higher dispersal<br />
tendency than L. dispar. Unlike L. dispar, larval dispersal tendency was inversely related<br />
to larval weight, with lighter individuals having a greater propensity to disperse (Zlotina<br />
et al. 1999). Settling velocity <strong>of</strong> L. dispar larvae was significantly higher compared to<br />
that <strong>of</strong> L. mathura. This result suggests that larvae <strong>of</strong> L. mathura may disperse farther<br />
via wind than L. dispar. Dispersal activity was influenced by time <strong>of</strong> day. Activity <strong>of</strong><br />
L. mathura larvae began to increase after 11AM and peaked at 5 PM , while L. dispar<br />
activity began to increase after noon , and peaked at 4 PM (Zlotina et al. 1999). This<br />
result indicates that L. mathura has a greater window <strong>of</strong> activity than L. dispar.<br />
Collectively, these results imply that ballooning <strong>of</strong> neonate larvae from infested ships<br />
could be an important means for the arrival.<br />
When population density is high, parasitism by Hymenoptera and polyhedral viral disease<br />
may result in high mortality <strong>of</strong> larvae and pupae (Roonwal 1979b).<br />
Pupa<br />
Pupation <strong>of</strong>ten occurs in groups <strong>of</strong> 40-50 in protected areas <strong>of</strong> branches, in leaf litter at<br />
the base <strong>of</strong> trees, or on the back or underside <strong>of</strong> signs or other objects (Browne 1968,<br />
Roonwal 1979b).<br />
CAPS PRA: Lymantria mathura 31