United States Department of Agriculture

Marketing and Regulatory Programs

Animal and

Plant Health Inspection
Service 

# Field Release of the Gall Wasp, **Aulacidea**
**subterminalis** (H ymen optera: Cynip idae)

, for Biological
Control of Invasive Hawkweeds ( **Hieracium** spp.) in the Continental
United States

Environmental Assessment,
S eptember 2010 United States Department of Agriculture

Marke ting and Regulatory Programs

Animal and

Plant Health Inspection
Service

# Field Release of the Gall Wasp,

**Aulacidea subterminalis** (Hymenoptera: Cynipidae), for Biological Control of Invasive

Hawkweeds ( **Hieracium** spp.) in the
Continental United States

Environmental Assessment, September 2010

Agency Contact: Shirley Wager

- Page, Branch Chief

P est Permitting
Plant Protection and Quarantine Animal and Plant Health Inspection Service U.S. Department of Agriculture 4700 River Road, Unit 133 Riverdale, MD 20737

– 1236

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## Table of Contents

I

. Purpose and Need for the Proposed Action

........................... 1

II

. Alternatives

.............................................. ............................. 2

I II. Affected Environment

.............................. ............................. 5

I V. Environmental Consequences

................. ........................... 12

V. Other Issues

............................................. ........................... 20

VI

. Agencies, Organizations, and Individuals C onsulted

........... 22

VII. References

............................................... ........................... 23

Appendix 1

............................................... ........................... 29 Appendix 2

............................................... ........................... 38 Appendix 3

............................................... ........................... 46 Appendix 4

............................................... ........................... 481 

## I. Purpose and Need for the Proposed
Action

The U.S. Department of Agriculture (USDA), Animal and Plant Health Inspection Service (APHIS)

, Plant Protection and Quarantine (PPQ),

Permit Unit is proposing to issue permits for release of the hawkweed gall

wasp
, Aulacidea subterminalis (H ymen optera: Cyn pidae). Th e agent would be used by the applicant for the biological control of the hawkweeds Hieracium pilosella, H. aurantiacum, H. floribundum, and H. flagellare in

the continental United States . Before permits are issued for release of A.

subterminalis

, the APHIS

– PPQ Permit Unit must analyze the potential impacts of the release of this agent into the continental United States.

This environmental assessment 1 (EA) has been prepared, consistent with USDA, APHIS' National Environmental Policy Act of 1969 (NEPA) implementing procedures (Title 7 of the Code of Federal Regulations (CFR), part 372). It examines the potential effects on the quality of the human environment that may be associated with the release of A.

subterminalis to control infestations of hawkweeds within the continental

United States . This EA considers the potential effects of th e proposed action and its alternatives, including no action.

The applicant’s purpose for releasing A. subterminalis is to reduce the severity of infestations of invasive hawkweeds in the United States

. There are many species of non -native, invasive hawkw eeds in North America. These species originated in Europe, which is the native range to a massive complex of species, subspecies, and varieties of Hieracium . Invasive hawkweeds were probably introduced into the eastern U nited States and Canada during the 1800s. Several hawkweed species are considered noxious in many western States . Introduced hawkweeds are highly competitive and relatively free of insects and pathogens in North America. These species out compete native and desirable vegetation, forming near monocultures and limiting economic use of infested land.

Existing management options for management of invasive hawkweeds are ineffective, expensive, temporary, and have nontarget impacts. For these

reasons
, the applicant has a need to release A. subterminalis, a hostspecific, biological control organism for the control of invasive hawkweeds, into the environment.

1 Regulations implementing the National Environmental Policy Act of 1969 (42 United States Code 4321 et seq. ) pro vide that an environmental assessment “shall include brief discussions of the need for the proposal, of alternatives as required by section 102(2)(E), of the environmental impacts of the proposed action and alternatives, and a listing of agencies and persons consulted.” 40 CFR § 1508.9.2 

## II . Alternatives

This section will explain the two alternatives available to the PPQ Permit

Unit —no action and issuance of permits for environmental release of A.

subterminalis . Although the PPQ Permit Unit’s alternatives are limited to a decision on whether to issue permits for release of A. subterminalis

, other methods available for control of hawkweeds are also described.

These c ontrol methods are not decisions to be made by the PPQ Permit

Unit , and their use is li kely to continue whether or not permits are issued

for environmental release of A. subterminalis , depending on the efficacy of

A. subterminalis to control hawkweeds . Th ese are methods presently being used to control hawkweeds by public and private concerns.

A third alternative was considered, but will not be analyzed further. Under

this third alternative, the PPQ Permit Unit would have issue d permit s for

the field rele ase of A. subterminalis ; however, the permit s would contain special provisions or requirements concerning release procedures or mitigating measures. No issues have been raised that would indicate special provisions or requirements are necessary

.



### A. No A ction

Under the no action alternative, the PPQ Permit Unit would not issue permit s for the field release of A. subterminalis for the control of invasive

hawkweeds . The release of this biological control agent would not take place. The following methods are presently being used to control hawkweeds; these methods will continue under the “No Action”

alternative and will likely continue even if permits are issued for release of

A. subterminalis , depending on the efficacy of the organism to control

invasive hawkweeds.

Hawkweeds may be controlled by using herbicides. Herbicides

, such as 2,4-D, clopyralid, aminopyralid, and picloram have resulted in the greatest degree of control when applied at the normally recommended rate for perennial pasture weeds (Noe l et al. , 1979; 1992a ; Wilson et al., 1997; 2005; 2006). Studies conducted at the University of Idaho showed that over 50 percent control was achieved for 6 years following treatment with

a 1 pint per acre rate of clopyralid (Lass and Callihan , 1992). Si milar

results were achieved usi ng picloram at the rate of 0.25 to 0.5 pounds of active ingredient per acre

. Other herbicides either failed to control meadow hawkweed or suppression was for less than 3 years (Lass and

Callihan , 1992; Miller et al. , 1998). More recently , aminopyralid (MilestoneTM) has been recommended as an effective herbicide ( http://www.msuextension.org/ruralliving/Dream/PDF/hawk.pdf last

accessed March 23, 2010 ). 1. Chemical
Control3 Mechanical co ntrol of invasive hawkweeds has had limited success. Digging the plants or otherwise disturbing the stolons, rhizomes, or roots only serves to spread the weed because plants can grow from buds on small root, stolon, and rhizome fragments. A stolon is a sh oot that bends to the ground or that grows horizontally above the ground and produces

roots and shoots at the nodes. A rhizome is similar to a stolon and is a horizontal, usually underground, stem that often sends out roots and shoots

from its nodes. Dis turbance by machinery spreads the weeds across the field. Local disturbances caused by grazing livestock, ungulates, and rodents also enhance the rate of spread of hawkweed. In lawns, mowing

does not kill invasive hawkweeds because the low -lying stolons and rhizomes are missed by the mower blades. Although mowing prevents seed production by removing flowering stems, repeated mowing encourages faster vegetative spread (Callihan et al. , 1997).

When perennial grasses, legumes, and other beneficial forbs are present in the plant community, fertilizers can help control hawkweed by increasing the competitive ability of the more desirable species. This may be particularly important on rangelands and pastures because these lands are generally not priority areas for supplemental fertilization

, and soil nitrogen levels may be inadequate for optimal grass health. Marked reductions of hawkweed density and vigor have been obtained by fertilizer treatments in the United States (Reader and Watt , 1981), Canada (Hay and Ouellette

, 1959), and New Zealand (Scott et al., 1990) when competing vegetation is

present. However , Reader and Watt (1981) found that repeated fertilizer treatments had no effect on dense patches of hawkweed that contained few grasses or other forbs . Depending on soil productivity and grass condition, a single nitrogen application may be sufficient for grasses to competitively suppress hawkweed growth for 3 to 5 years (Reader, 1990).

Five species of Hieracium

- specific insects have been tested and released for the biological control of hawkweeds in New Zealand. They are:

(1.) Aulacidea subterminalis (Insecta: Hymenoptera: Cynipidae)— hawkweed gall wasp ; (2.) Cheilosia psilophthalma ( Insecta: Diptera: Syrphidae) —hawkweed

crown -feeding hover fly; (3.) Cheilosia urbana (Insecta: Diptera: Syrphidae) —hawkweed rootfeeding hover fly; (4.) Macrolabis pilosellae (Ins ecta: Diptera: Cecidomyiidae) —hawkweed stem gall midge; and

(5.) Oxyptilus pilosellae (Insecta : Lepidoptera: Pterophoridae)— hawkweed plu me moth

.

None of these insects have been released in North America. An additional

Aulacidea wasp
, besides A. subterminalis

, is being considered as a 2. Mechanical
Control

3. Cultural
Control

4. Biological
Control4 bio logical control candidate for release in North America

. Aulacidea pilosellae , a wasp that forms galls on stems, leaves , and stolons of hawkweeds, i s currently undergoing testing and appears to be host- specific (Littlefield et al. 20 08)

.



### B

. Issue P ermit s for E nvironmental R elease of

**A. subterminalis**

Under this alternative, the PP Q Permit Unit would issue permits for the

field release of the gall wasp

, A. subterminalis , for the control of invasive

hawkweeds . Th ese permit s would contain no special provisions or requirements concerning release procedures or mitigating measures.

Bio logical Control A gent I nformation

Phylum: Arthropoda

Class: Insecta Order: Hymenoptera Superfamily: Cynipoidea Family: Cynipidae Subfamily: Cynipinae Tribe: Aylacini

Genus: Aulacidea Species : A. subterminalis Niblett

a. Native R ange

Aulacidea subterminalis is fairly common and widespread on H. pilosella in Northern Europe (Syrett et al., 1998). Fauna Europaea (2007) records

A. subterminalis as present in Britain, France, and Germany. In its native habitat, A. subterminalis occurs on H. pilosella inhabiting arid, nutrientpoor grasslands, poorly fertilized upland pastures, inland dunes, and manmade heaths (Ellenberg , 1988).

b. Other A reas of I ntroduction

A. subterminalis is one of five insect bio logical control agents introduced

to New Zealan d for suppression of H. pilosella (Syrett et al., 2 001). In New Zealand, A. subterminalis was approved for import and release in 1998, and was first released in 1999. It has since been released at 99 sites throughout the North and South Island, with establishment confirmed at approximately 37 percent of these sites (Aspinall , 2006; Smith et al., 2007). The failure of many releases was due to drought conditions following the release or site disturbance due to cropping or herbicide treatment (Smith et al., 2007). 1. Taxonomy

2. Geographical Range of **A.**

**subterminalis**5 c. Expected A ttainable R ange of **A. subterminalis** in North
America

A. subterminalis is fairly common and widespread on H. pilosella in Northern Europe and it is also establishing well in more moist areas of New Zealand, but is not well adapted to survive in dry or droughty environments where plants may senesce early ( Littlefie ld et al., 2008). Climate matching using CLIMEX Version 1.1 (CSIRO Australia) suggests that the wasp’s potential range in North America should match much of the distributions of the targeted Hieracium species (Littlefie ld et al., 2008).

A. subterminalis is parthenogen et ic ( a form of asexual reproduction in which females produce eggs that develop without fertilization by males ). Eggs are laid into the stolen tips. Eggs a re milky -white in color, 0.24 millimeters ( mm ) long and 0.09 mm wide and have a stalk appro ximately 1.18 mm long. Larvae develop inside the developing stolon-tip gall. Galls are abnormal outgrowths of plant tissues and can be caused by various parasites, including insects . Galls are about 4.5 mm in diameter (Klöppel

et al.

, 2003), but gall size is dependent upon the number of larval chambers

present. Galls become evident approximately five weeks after the plant is

expos ed to the female wasp . G alls contin ue to enlarge for an additional

5 weeks (Littlefield et al. , 2008 ). Mature larvae are somewhat fusiform (tapered at each end, spindle- shaped) , with a distinctive, unpigmented

head. The wasp overwinters in the larval stage. Pupation occurs in April, and adult wasps emerge from the galls from the beginning of May to the

end of June in Switzerland. A. subterminalis has one generation per year. Adults are generally short lived; up to a week under ambient laboratory

conditions.



## III

. Affected Environment

A. Target Weeds

(1.) Scientific name: Hieracium pilosella L. Family: Asteraceae Synonyms: none

Common name: Mouse ear hawkweed

(2.) Scientific name: Hieracium aurantiacum L. Family: Asteraceae Synonyms: H. brunneocroceum Pugsley. Common name: Orange hawkweed, Grim the Collier 3. Life History of

**A.**

**subterminalis**6 (3.) Scientific name: Hieracium floribundum Wimm. & Grab. Family: Asteraceae Synonyms: none Common name: King devil hawkweed

(4.) Scientific name: Hieracium x flagellare Willd. ( caespitosum x pilosella ) Family: Asteraceae Synonyms: Hieracium x duplex

, H. x macrostoloum Varieties: cernuiforme, flagellare, glatzense Common name: Whiplash hawkweed

Introduced hawkweeds are perennial plants that contain a milky sap and are generally stoloniferous (plants that bear or form stol ons). Hieracium piloselloides and H. glomeratum are the only non-stoloniferous introduced species. Each young plant consists of a rosette of 5 to 12 hairy leaves arising from a short, thick rhizome with a shallow, fibrous root system. The flowers are ye llow or orange ( H. aurantiacum is the only orangeflowered species). New plants begin as seedlings or as leaves sprouting from stolons, rhizomes (Thomas and Dale, 1974), and/or roots (Peterson

, 1979). Juvenile plants consist of low- lying rosettes that de velop into flowering plants as erect, slender stems arise from the rosette center.

All of the invasive hawkweeds reproduce asexually by apomixis (asexual seed production) and by vegetative spread by stolons, rhizomes, and root

buds. Orange hawkweed sen ds out from three to eight long, slender stolons along the soil surface. Meadow hawkweed also produces long, slender stolons, but some of its vegetative structures grow as shallow, underground rhizomes. Stolons and rhizomes, initiated from buds at the base of rosette leaves, begin to grow when the plant initiates flowering. Once established, vigorous stolon growth quickly expands the colony, forming dense patches that can have as many as 3,200 plants per square yard. The slender, leafy stolons elongate through the summer and form daughter rosettes at their tips (Thomas and Dale, 1974). Hawkweeds regrow each year from short, below-ground rhizomes. In H. piloselloides

, axillary buds develop into secondary flowering shoots rather than stolons.

Invasive hawkweeds also occasionally produce seeds sexually by periodic pollination and outcrossing. At lower elevations, flowering occurs around

mid

- June and seeds ripen by early August. Mature seeds can germinate as soon as they are released from the plant (St ergios , 1976). Studies have shown that seeds are viable in the soil for up to 7 years (Panebianco and

Willemsen , 1976). Invasive hawkweeds have relatively short life cycles, completing a generation within 4 months.

Although most new hawkweed infestation s are probably started by seeds, expansion of established populations is mostly vegetative. Thomas and7 Dale (1974) found that only 1 percent of new plants in a H. floribundum population were derived from seedlings. Studies in eastern Canada

showed that s eeds are not carried far by the wind; although minute barbs along ribs on the seeds enable them to stick to hair, fur, feathers, clothing, and vehicles, and can be carried long distances.



### B . Areas A ffected by H awkweeds

In their native ranges, hawkweeds occur in the northern, central, and eastern portions of Europe as ruderal species (plants that grows in rubbish, poor land, or waste) of pastures, roadside cutbanks, abandoned fields, and meadows (Skalinska , 1976). In most cases they are found in small, isolated pockets. Their highest densities are found on recently disturbed areas and they do not persist as dominant members of the early successional community (Skalinska, 1976). From Europe, hawkweeds spread to North America , New Zealand, Australia, Chil e, and Argentina.

In North America, invasive hawkweeds are prim arily weeds of moist

pastures, forest meadows, and mesic (moderately moist) rangeland. Based on current infestations, habitats most susceptible to invasion range from the lowlands of the northern Pacific Coast to elevations of 5,000 feet or

more in the Mountain S tates (Littlefiel d et al. , 2008). Sites most vulnerable to invasion include roadsides, mountain meadows, clearings in forest zones, permanent pastures, hayfields, cleared timber units, and abandoned farmland where the soil is well drained, coarse textured, and moderately low in organic matter. Consequently, elk habitat, recreation areas, and pristine mountain meadows in areas that have a climate similar to that in their native range are particularly susceptible. Although introduced hawkweeds can grow in open woodlands, they do not tolerate shade very well. Across the United States and Canada, introduced hawkweeds are closely associated with habitats that can support oxeye daisy ( Ch rysanthemum leucanthemum ), sulfur cinquefoil ( Potentilla

recta ), spotted knapweed ( Cen taurea maculosa ), gray goldenrod ( Solidago

nemoralis ), wild carrot ( Daucus carrota ), dandelion ( Taraxacum officinale ), and Kentucky bluegrass ( Poa pratensis ) (Thomas and Dale, 1974; Maycock and Guzikowa, 1984). None of the introduced hawkweed species are found in the natural grasslands or shrub-steppe of the northern Intermountain West, and are not expected to become problem weeds in any dry habitat usually associated wi th western rangelands. Neither meadow nor orange hawkweed appears to survive in annually tilled cropland.

There are many species of non -native, invasive hawkweeds in North

America. These species originated from Europe, which is the native range

to a m assive complex of species, subspecies, and varieties of Hieracium

. Invasive hawkweeds were probably introduced into the eastern United

States and Canada during the 19th century. Two important subgenera 1. Native and

Introduced

R ange of

Hawkweeds

2. Known and
Potential

D istribution of

Hawkweeds in

North America8 distinguish the group of invasive species, subgenera Pilosella and

Hieracium . Species in these subgenera vary widely, both morphologically and in their invasion history and species distribution. The majority of species belong to subgenus Pilosella , including the species considered as targets for A. subterm inalis . ( See figure 1 for a proposed phylogeny (taxonomic grouping) of North American Hieracium species. )

Figure 1. Proposed taxonomic grouping of North American Hieracium species by Gaskin and Wilson (2007).9 The present distributions of these e xotic species in the United States and

Canada are provided in t able 1

.

Table 1 . Invasive, Nonnative **Hieracium** species, from subgenus **Pilosella**

, in the United States and Canada (from Wilson et al.

, 2006).

Species Distribution

H. arvicola Nageli et Peter CAN : NB

H. aurantiacum L. CAN : AB, BC, NB, NL, NS, ON, PE, QC,
SK

USA : AK, AR, CA, CO, CT, FL, ID, IN, I A

, MA, MD, ME, MI, MN, MT, NC, NH, NJ, NY, OH, OR, PA, RI, SD, VA, VT, WA, WV, WI, WY

H. bauhini Schult CAN : BC

USA : CT, ID, MA, MN, NH, NY, VT, WA

H. brachiatum Bertol. ex DC. CAN : QC

USA : NY

H. caespitosum Dumort CAN: AB, BC, NB, NL, NS, ON, PE, QC

USA : CT, DE, GA, ID, IL, IN, KY, MA, MD, ME, MI, MN, MT, NC, NH, NJ, NY, OH, OR, PA, RI, SC, TN, VT, VA, WA, WV, WI

H. derubellum Gottschl. et Schuhw. [ H. atramentarium act.] CAN : QC

USA : MI, NY, WI

H. flagellare Willd. [incl. H. macrostolonum ] CAN : BC, NB, NL, NS, ON, PE, QC

USA : CT, ID, MA, ME, MI, NH, NY, PA, VA, VT

H. floribundum Wimm. & Grab. CAN : BC, NB, NL, NS, ON, PE, QC

USA : CT, ID, MA, MD, ME, MN, MT, NH, NJ, NY, OH, OR, RI, VA, VT, WA, WV

H. fuscoatrum Nageli & Peter USA : CT, NY, RI

H. glomeratum Froel. CAN : BC, ID, WA

H. lactuella Wallr. CAN : NS

H. pilosella L. CAN : BC, NB, NL, NS, ON, PE, QC, Saint Pierre and Miquelon

USA : CT, DE, GA, MA, MD, ME, MI, MN, NC, NH, NJ, NY, OH, OR, PA, RI, TN, VA, VT, WA, WV

H. piloselliflorum Nageli et Peter [incl. H. apatelium ] CAN : NB, NL, NS, ON, PE, QC

H. piloselloides Vill. [incl. H. preaaltum ] CAN: BC, NB, NL, NS, ON, PE, QC

USA: CT, DE, GA, IL, IN, IA, MA, ME, MI, MN, MT, NC, NH, NY, OH, PA, RI, SC, VA, VT, WI, WV

H. stoloniflorum Waldst. e t Kit. CAN : QC10 Hieracium a urantiacum (orange hawkweed) was introduced into Vermont in 1875 as an ornamental and within 25 years spread throughout much of northeastern United States and southeastern Canada (Voss and Böhlke, 1978). Within 25 years it had spread throughout much of New England, west to Michigan (Voss and Böhlke, 1978) and into Canada from New Brunswick to Ontario (Britton and Brown, 1970). It now occurs throughout the eastern seaboard and into the Midwest, extending west to Minnesota and Iowa, and south to Virginia and North Carolina (Johnson and Thomas, 1978). First recorded in Spokane, Washington, in 1945 (Marion Ownbey Herbarium, Wash. State Univ.), H. aurantiacum has been collected from gardens in Nevada County, California (Munz and

Keck , 1959), on the east slope of the Rocky Mountains in Colorado (Weber , 1990), from coastal Oregon and Washington (Rickett, 1973), and in southwestern British Columbia (Guppy, 1976). In the western United States, populations of orange hawkweed were first reported from coastal Washington and Oregon (Peck, 1941; Abrams and Ferris , 1960). It has also been reported from the l ower mainland of British Columbia (Guppy, 1976) where it infests pastures, old fields, and roadsides. H. aurantiacum is now also recorded in Idaho, Montana, and Wyoming (USDA– NRCS

, 2008 ). It is more widespread than other hawkweed species primarily because it is planted as an ornamental and often escapes cultivation.

Hieracium pilosella (mouse ear hawkweed) is widespread in the eastern

U nited States and Canada. In the west, it occurs in western Washington, western Oregon, Alaska, and British Columbia.

Hieracium flagellare (whiplash hawkweed) occurs primarily in the

northeastern U nited States and eastern Canada, but also occurs in British Columbia and Wyoming.

Hieracium floribundum (king devil hawkweed) w as first reported in northern New York in 1879 (Voss and Böhlke, 1978) and 1900 respectively (Kennedy 1902). Recently, infestations of H. floribundum have been reported in western Washington (WSNWCB, 1997), Northern Idaho (L. Wilson, unpubl. data.), and Montana (USDA– NRCS , 2008), and is reportedly expanding in range.

Hieracium piloselloides has been recorded in the western S tates of Montana and Washington (USDA– NRCS , 2008), and is also reportedly expanding in range. Suspected hybridization between H. caespitosum

, H. floribundum, and H. piloselloides

, and the potential threat of these species in western habitats is the rationale for including these species as target weeds in the biological control program.

Hiera cium caespitosum (meadow hawkweed) was likely introduced into the United States in 1828 (Britton and Brown, 1970), and is now commonly found from Québec and Ontario (Frankton and Mulligan, 1970)11 southward to Georgia and Tennessee (Rickett, 1973). The firs t record of H. caespitosum in the western United States was in Pend Oreille County, Washington, in 1969 (Marion Ownbey Herbarium, Wash. State Univ.). H. caespitosum is now recorded in the western States of Idaho, Montana, Oregon, Washington, and Wyoming, and is widespread throughout most of

southern British Columbia and Alberta.



### C

. Plants R elated to I nvasive H awkweeds and T heir

D istribution

Plants related t axonomically to invasive hawkweeds would be the most likely to be attacked by the proposed biologi cal control organism A.

subterminalis. Plants related to the target hawkweeds are discussed below.

There are many species of Hieracium native to North America (Wilson

et al. , 1 997). Distribution of native Hieracium species ranges from the

east coast t o west coast, from sea level to over 9 ,000 ft. in elevation, and from northern to southern latitudes. As with the European hawkweeds, the number of native species has long been disputed. In order to understand species relationships in Hieracium , a brief outline of the generic divisions is provided.

There are generally considered to be about 36 species of Hieracium belonging to the three subgenera in America north of Mexico (Strother, 2006). A review of the regional flora revealed that about 25 species are native to the United States and Canada and fall into two subgenera:

Hieracium (3 species) and Chionoracium (22 species). Subgenus

Hieracium occurs chiefly throughout the boreal regions of North America and Eurasia, and is represented in North America by H. umbellatum L., H.

robinsonii Zahn, and H. canadense Michx. (considered part of

Chionoracium by Gaskin and Wilson, 2007).

Subgenus Chionoracium is restricted to the New World and contains most

of our native taxa. Conflicting characters used to di stinguish species have generated considerable debate regarding classification. For example, Guppy (1978) suggested that H. scouleri Hook., H. albertinum Farr., and H. cynoglossoides L. are too closely correlated in characters to be distinguished into sepa rate species and

, instead , form a complex of species. Guppy (1978) also suggested that H. umbellatum L. and its close relative, H. canadense Michx., may be two forms of the same species. Deardorff (1977) added H. longiberbe Howell and H. nudicale Heller to
the H. scouleri complex. Kartesz and Meacham (1999) provided a comprehensive synthesis of Hieracium spp. in America north of Mexico in which they list 108 species, subspecies, and hybrids, clearly giving evidence of the myriad of described taxa in the genus.12 Ten additional hawkweed species found in the United States and Canada are introduced from Europe and belong to two subgenera. Four species belong to the subgenus Hieracium : H. argillaceum Jordan group, H. groenlandicum Arv. -Touv., H. lachenalii K. C. Gmel., H. murorum L., and

H. sabaudum L. Plants in this subgenus, except for H. lachenalii, have not become weedy in North America. The remaining six species belong to the subgenus Pilosella and include the species targeted for biological control by A. subterminalis

.

S pecies that were used in testing the specificity of A. subterminalis to
hawkweeds are lis te d in appendix 1.



## IV

. Environmental Consequences

A. No A ction

Most invasive hawkweeds are perennial, creeping stoloniferous plants that

can c rowd out competing vegetation. They are not drought tolerant and cannot withstand prolonged periods of drying. However, they are very cold tolerant and easily persist at higher elevation, making the upland habitats through the Rocky Mountains and oth er w estern mo untain ranges susceptible to invasion. In northern Idaho and British Columbia, habitats are open ponderosa pine and interior cedar-hemlock. The re are no known medicinal or herbal uses for the invasive hawkweeds, nor do they have any

known social or recreational uses.

Invasive hawkweeds do affect threatened, endangered, and sensitive species because of their ability to form dense monocultures. One such example is encroachment by mouse-ear hawkweed of habitat occupied by the threatened golden paintbrush, Castilleja levisecta , in Thurston County, Washington (USFWS , 1997). These monocultures compete for soil moisture and nutrients, thereby posing risks to indigenous species. As hawkweed monocultures invade species-rich range and mountain habitats, ecosystem functions and ecological relationships are affected. No known studies specifically address the effects of hawkweed on nutrient cycling and disturbance regimes ; however, hawkweed’s ability to dominate a community suggests that these species do affect habitat function. Hawkweeds generally negatively impact plant and animal diversity , and there is serious concern about the loss of native plant biodiversity in

infested areas. Ecological losses in plant and animal diversity can be enormous but cannot be economically calculated. Additional expenditures

result from control costs. Hawkweeds are tenacious invaders and , once established, quickly develop into a patch that continues to expand until it

covers the site with a solid mat of rosettes. Fora ge species in pastures and abandoned farmland are choked out by the advancing front of hawkweed.

Hawkweeds also t hreaten lawns and gardens. Hawkweeds have been 1. Impact of
S pread of

Invasive

Hawkweeds13 reported to have allelopathic effects (inhibition of growth in one species of plants by chemic als produced by another species ) on neighboring vegetation (Dawes and Maravolo , 1973; Makepeace , 1976).

Meadow hawkweed causes severe allergenic reactions. Reactions in people closely working with meadow hawkweed include minor skin rashes, sneezing, cong estion, and difficulty in breathing. Similar effects have not been reported in animals. It is not known what portion of the plant causes this allergenic reaction , although latex and other chemicals of closely related Asteraceae plants can cause dermatiti s (Dawe s et al.

, 19 96).

Invasive hawkweeds have not been placed on the Federal Noxious Weed List but have been listed as noxious in Idaho, Montana, Washington, and

British Columbia. Regulations vary by S tate/ Province but generally involve actions such as restricted importation and/or quarantine and prevention, containment, eradication

, or other control measures.

The continued use of chemical herbicides, and mechanical and biological

controls at current levels would be a result if the “no action” alter native is

chosen. These environmental consequences may occur even with the implementation of the biological control alternative, depending on the efficacy of A. subterminalis to reduce invasive hawkweed populations in

the Continental United States.

a. Chemical C ontrol

The use of herbicides, while effective, is limited to relatively accessible sites and control is only temporary. Broadcast applications of herbicides could have adverse impacts on nontarget vegetation if not carefully applied.

b. Me chanical C ontrol

Disturbance of hawkweeds by machinery spreads the weeds across the field. Local disturbances caused by grazing livestock, ungulates, and rodents also enhance the rate of spread of hawkweed. In lawns, mowing

does not kill invasive hawkwe eds because the low lying stolons and rhizomes are missed by the mower blades. Although mowing prevents seed production by removing flowering stems, repeated mowing encourages faster vegetative spread.

c. Biological C ontrol

No organisms for the biolo gical control of hawkweeds have been released

in North America. Gall formation by A. subterminalis diverts plant nutrients from the normal growth of other plant tissues. Under certain 2. Impact from

U se of O ther
C ontrol

M ethods14 conditions, galls may stress the plant, reducing competitive ability, seed production, and long distance spread of the weed.



### B. Issue P ermit s for E nvironmental R elease of **A.**
**subterminalis**

Host specificity of A. subterminalis to the invasive hawkweeds Hieracium pilosella, H. aurantiacum, H. floribundum, and H. flagellare has been demonstrated through scientific literature, field observations, and host specificity testing. If an insect species only attacks one or a few closely

related plant species, the insect is considered to be very host- specific

. Host specificity is an essential trait for a biological control organism proposed for environmental release.

a. Scientific L iterature

In previous literature, A. subterminalis has only been reported to induce galls on H. pilosella and has not been recorded from any other spec ies (Syrett et al., 1 998).

b. Field O bservations

In the field, A. subterminalis has not been observed or reported on any species other than H. pilosella (Syre tt et al., 1998).

c. Host S pecificity T esting

Host specificity tests a re test s to determin e how many plant species A.

subterminalis attacks/eats, and whether nontarget species may be at risk

. In host specificity testing (Syrett et al., 1998; 2001; Grosskopf et al., 2001,

Littlefield et al., 2008), gall formation by A. subterminals only occurred on invasive hawkweed species targeted for biological control. No native hawkweeds or plant species of closely related genera were attacked

.

(1) Site of Q uarantine and F ield S tudies

Laboratory tests were conducted at the CABI -Europe Station in Delémont, Switzerland, at the insect containment facility located at the Canterbury Agriculture and Science Centre, Lincoln, New Zealand, and in quarantine

facilities at Montana State University ( MSU ), Bozeman, Montana. Open

field releases began in 1999 in N ew Zealand.

(2) Test P lant L ist

The list of plant species used for host specificity testing of A.

subterminalis is shown in appendix 1. The strategy used for selecting 1. Impact of **A.**
**subterminalis** on Nontarget

P lants15 plants for testing is based on the phylogenetic approach, where closely related species are theorized to be at greater risk of attack than are distantly related species (Wapshere, 1974).

The initial host specificity testing of A. subterminalis was conducted for the field release of the wasp in New Zealand. Plants utilized for testing were selected from the following categories (Syret t et al. , 1998):

• representatives of the plant family Asteraceae (the family to which

hawkweeds belong);

• plant species on which the gall wasp has been recorded in the field;

• cultivated plant speci es and representatives of New Zealand native plant

species (various non

- Asteraceae families) that have not previously been exposed to the wasp; and

• plant species that are hosts of close relatives of A. subterminalis

, that is,

hosts of other species of Aulacidea

.

Host specificity testing at the CABI -Europe Station, Delémont, and at the Canterbury Agriculture and Science Centre, Lincoln, New Zealand, consisted of eight species within the genus Hieracium , including five from the subgenus Pilosella and three from the subgenus Hieracium

. Thirteen additional species from the tribe Lactuceae were tested, including native New Zealand plant species that would not previously have been encountered by the wasp ( Knightia excelsa

, Raoulia hookeri , Kirkianella
novae-zelandiae

, Celmisia lyalli

, Helichrysum bellidioides

, Gaultheria depressa , and Discaria toumatou ). A further 11 species within the family Asteraceae were tested, including economically significant cultivated plant species. Species from 21 other families containing important cultivated and native plants were also included, making a total test plant list of

48 species.

Plants for testing the host range of A. subterminalis were selected from seven possible categories. Test categories consisted of the follo wing:

CATEGORY 1: Genetic types of Hieracium (varieties, races, forms, genotypes, apomicts, etc.).

CATEGORY 2: North American species in the same genus as Hieracium

, divided by subgenera, including economically and environmentally important plants.

CATEGORY 3: North American species in other genera in the Asteraceae family, divided by subtribe, tribe, and subfamily, including economically and environmentally important plants.16 CATEGORY 4: Threatened and endangered species in the Asteraceae family, divided by subgenus, genus, subfamily, and tribe.

CATEGORY 5: North American species in other families in the Asterales order that have some phylogenetic, morphological, or biochemical relationship to the target weed, including economically and environment ally important plants.

CATEGORY 6: North American species in other orders that have some morphological or biochemical relationship to the target weed, including economically and environmentally important plants.

CATEGORY 7: Any plant on which the biolo gical control agent or its close relatives (within the same genus) have been previously recorded to feed and/or reproduce.

( See appendix 2 for a more detailed description of plants selected in each

category. )

Host specificity testing of A. subterminalis at MSU and CABI -Europe included as many native North American representatives of the various taxa as possible

. Aulacidea subterminalis was tested on 65 plant species of which 35 are species native to North America (appendix 1)

.

(3) Discussion of Host Specificity Testing

New Zealand Tests —In overseas testing, Syrett et al. (1998 and 2001) and Grosskopf et al. (2001) reported gall formation on H. pilosella and H.

aurantiacum but n ot on other tested plant species , while Grosskopf and Senhadji Navarro (2004) reported gall formation on H. pilosella but not on

H. aurantiacum or other teste d plant species.

North American Tests —During the North American host specificity test, galls were only induced on four exotic species: H. pilosella (its native host), H. a urantiacum, H. flagellare, and H. floribundum (appendix 4). No native or other test plants were infested

.

( See a ppendices 3 and 4 for host specificity study results. )

To assess the impact of A. subterminalis on H. pilosella in New Zealand, an experim ent was set up to examine the effect of the gall wasp on plant growth under stress-free conditions in a shade house trial with potted plants (Klöppe l et al., 2003). Plants with galls showed a reduction in stolon length of 75 percent . They had slightly lo wer total dry matter and reduced root weight. In a glasshouse experiment, the impact of water stress, nutrient stress, and plant competition on growth of H. pilosella and 2. Impact of **A.**
**subterminalis**

on Hawkweeds17 performance of the gall wasp were measured. The number and mean diameter of gall cl usters were not significantly different between treatment and control plants . However, galled plants produced more, but shorter,

stolons in all stress treatments and stolons that were more branched in

nutrient- and water

- stressed plants, than ungalled plants. Galling by A.

subterminalis is likely to reduce vegetative reproduction of H. pilosella whether or not plants are stressed, indicating that the wasp may be a successful biocontrol agent (Klöppel et al. , 20 03).

Researchers expect that A. subterminalis will be able to build up substantial populations in North America and, thereby , reduce aboveground biomass and seed output of established invasive hawkweed patches (Littlefiel d et al., 20 08)

. It is difficult to predict the long

- term impact on

invasive hawkweed infestations; however, reductions in the above-ground growth rate and seed output may lead to reduced competitive ability of the weed and reduced patch expansion and long- distance dispersal of invasive

hawkweed seeds and establishment of new infes tations (Littlefield et al., 2008).

Once a biological control agent such as A. subterminalis is released into

the environment and becomes established, there is a slight possibility th at
it could move from the target plant s (invasive hawkweeds) to attack nontarget plants. Host shifts by introduced weed biological control agents to unrelated plants are rare (Pemberton, 2000). Native species that are closely related to the target species are the most likely to be attacked (Louda et al., 2003). If other pl ant species were to be attacked by A.

subterminalis , the resulting effects could be environmental impacts that may not be easily reversed. Biological control agents such as A.

subterminalis generally spread without intervention by man . In principle,

ther efore, release of this biological control agent at even one site must be considered equivalent to release over the entire area in which potential

hosts occur , and in which the climate is suitable for reproduction and

survival.

In addition, th is agent may not be successful in reducing hawkweed

pop ulations in the continental United States. Worldwide, biological weed control programs have had an overall success rate of 33 percent; success rates have been considerably higher for p rograms in individual countri es (Culliney, 2005). Actual impacts on hawkweeds by A. subterminalis will not be known until after release occurs and post- releas e monitoring has been conducted. It is expected that A. subterminalis will reduce invasive hawkweed populations by reducing above-ground biomass and seed output of established invasive hawkweed patches

.

A. subterminalis is a wasp but it does not sting or produce venom. It is a plant-feeding wasp which poses no risk to humans or other animals

. 3. Unce rtainties

R egarding the

E nviron

mental

R elease of **A.**

**subterminalis**

4 . Human H ealth18 “Cumulative impacts are defin ed as the impact on the environment which results from the incremental impact of the action when added to other past, present and reasonably foreseeable future actions regardless of what agencies or person undertakes such o ther actions” (40 CFR 1508.7).

Many Federal and State agencies, as well as private entities, conduct programs to manage hawkweeds, as well as other invasive weeds

. Chemical and mechanical methods, as described previously in this document, are u sed in a wide range of habitats. Some of t hese control programs in Idaho, Washington, Montana, Oregon, and Alaska are listed

below.

Idaho

— Idaho Department of Lands Kootenai County Weed Board Palouse Clearwater Cooperative Weed Management Area Panhandle Lakes Cooperative Weed Management Area Potlach Corporation

U.S. D epartment of Interior (USDI), Bureau of Land Management Rimrock Hawkweed Cooperative Selkirk Cooperative Weed Management Area Benewah County Weed Board Pend Oreille County Weed Board

Washington

— Washington State Weed Board Stevens County Weed Board King County Weed Board

Montana

— Gallatin National Forest

Lolo National Forest

Bitterroot National Forest

Flathead National Forest

Kootenai National Forest

Glacier National Park

U.S. Fish and Wildlife Service

– Lost Trail Refuge

USDI , Bureau of Land Management Lincoln County Weed Department Flathead County Weed Department Flathead County Extension Ravalli County Weed Department Sanders County Extension Missoula County Weed Department Mineral County Weed Department Lake County We ed Department Glacier County Weed Department 5. Cumulative Impacts19 Montana **cont’d.**

— Granite County Weed Department Powell County Weed Department Gallatin County Weed District

Judith Basin Weed District Big Sky Weed Management Area

Montana Department of Agriculture

Montana Noxious Weed Trust Fund Montana Department of N atural R esources and Conservation

Confederated Salish Kootenai Tribe Plum Creek Timber Company

Oregon

— Oregon Department of Agriculture Union County Weed Control Deschute County Weed Control Wallowa County Weed Control Asotin County Weed Control

East Multnomah Soil and Water Conservation District

Wallowa Whitman National Forest

Alaska

— Kodiak National Wildlife Refuge

Homer Soil and Water Conservation District

Kodiak Garden Club Koniag Native Corporation University of Alaska Fairbanks Cooperative Extension Service Upper Susitna Soil and Water Conservation District

Juneau Invasive Plants Action

Release of A. subterminalis is not expected to have any negative cumulative impacts in the continental United States because of its host specificity to invasive hawkweeds

. Effective biological control of invasive

hawkweeds will have beneficial effects for weed management programs, and may result in a long-term, non-damaging method to assist in the

control of hawkweeds.

Section 7 of the Endangered Species A ct (ESA) and ESA’s implementing regulations require Federal agencies to ensure that their actions are not likely to jeopardize the continued existence of federally listed threatened

and endangered species or result in th e destruction or adverse modification

of critical habitat.

T he adverse effects of hawkweed invasion on threatened and endangered species is not fully known ; however, it can be assumed that invasive hawkweeds do affect threatened, endangered, and sensitive species because of their ability to form dense monocultures. One such example is 6. Endangered Species Act20 encroachment by mouse- ear hawkweed into habitat occupied by the threatened golden paintbrush, Castilleja levisecta , in Thurston County, Washington (USFWS , 1997). These ha wkweed monocultures compete for soil moisture and nutrients, thereby posing risks to native plant species. As hawkweed monocultures invade species -rich range and mountain habitats, it is thought that ecosystem functions and ecological relationships

are affected (Littlefiel d et al., 200 8)

.

Host specificity has been demonstrated by A. subterminalis . Fro m the literature, field observations, and host specificity testing, it has been

de termined that A. subterminalis is specific to species in the genus

Hiera cium , subgenus Pilosella (Littlefi eld et al., 2008). Ther e are no
Hieracium species federally listed as threatened or endangered. No federally listed animal species are known to use invasive hawkweeds.

For these reasons , APHIS has determined that envi ronmental release of A.

subterminalis will have no effect , or a potentially beneficial effect

, depending on the efficacy of the organism, on threatened and endangered species.



## V . Other Issues

Consistent with Executive Order (EO) 12 898, “Federal Actions to Address Environmental Justice in Minority Populations and Low-income Populations,” APHIS considered the potential for disproportionately high and adverse human health or environmental effects on any minority populations a nd low-income populations. There are no adverse

environmental or human health effects from the field release of A.

subterminalis and will not have disproportionate adverse effects to any minority or low-income populations.

Consistent with EO 13045, “Protection of Children From Environmental Health Risks and Safety Risks,” APHIS considered the potential for disproportionately high and adverse environmental health and safety risks to children. No circumstances that would trigger the need for special

environmental reviews are involved in implementing the preferred alternative. Therefore, it is expected that no disproportionate effects on

children are a nticipated as a consequence of the field release of A.

subterminalis

.

EO 13175, “Consultation and Coordination with Indian Tribal Governments,” was issued to ensure that there would be “meaningful consultation and collaboration with tribal officials in the development of Federal policies that have tribal implications....”21 APHIS is consulting and collaborating with Indi an tribal officials to ensure that they are well -informed and represented in policy and program decisions that may impact their agricultural interests in accordance with

EO 13175.22 

## VI. Agencies, Organizations, and
Individuals Consulted

The Technical Advisory Group for the Biological Control Agents of Weeds (TAG) recommended the release of A. subterminalis on August 10,

2009

. TAG members that reviewed the release petition ( Littlefiel d et al.,

2008 ) included rep resentatives from APHIS, Cooperative State Research, Education, and Extension Service, Forest Service, Environmental Protection Agency, U.S. Army Corps of Engineers, the National Plant Board, and representatives from Canada

.

This EA was prepared and revi ewed by APHIS. The addresses of participating APHIS units, cooperators, and consultants (as applicable)

follow.

U.S. Department of Agriculture Animal and Plant Health Inspection Service Policy and Program Development

Environmental Services

4700 River Ro ad, Unit 149 Riverdale, MD 20737

U.S. Department of Agriculture Animal and Plant Health Inspection Service Plant Protection and Quarantine Pest Permitting 4700 River Road, Unit 133 Riverdale, MD 20737

Montana State University Department of Land resour ces and Environmental Sciences

P.O. Box 173120 Bozeman, MT 59717–302023 

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Voss, E.G., and Böhlke , M.W., 1978. The status of certain hawkweeds ( Hi eracium subgenus Pilosella ) in Michigan. Mich. Bot. 17(2): 35– 47.

Wapshere, A.J ., 1974. A strategy for evaluating the safety of organisms for biological weed control. Annals of Applied Biology. 77: 201–211.

Wapshere, A. J. , 1983. Problems in the use of plant biochemistry for establishing the safety of biological control agents for weeds: The

Chondrilla and Echium / Heliotropium cases. Entomophaga. 28 : 287–294.

Washington State Noxious Weed Control Board, 1997. WSR 97 –24–051

Permanent Rules Noxious Weed Control Board.

Weber, W.A. , 1990. Colorado flora: e astern slope. Univ. Press of Colorado, Niwot.

Wilson, L.M., Fehrer, J., Bräutigam, S., and Grosskopf, G., 2006 . A new invasive hawkweed, Hieracium glomeratum (Lactuceae, Asteraceae), in

the Pacif ic Northwest. Can adian Journal of Bot any. 84: 133–142.

Wilson, L. M., McCaffrey, J.P., Quimby, Jr., P.C., and Birdsall, J.L., 1997. Hawkweeds in the Northwestern United States. Rangelands . 19(4): 18– 23.

WSNWCB

― S ee Washington State Noxious Weed Control Board

.

Zomlefer, W.B., 1994. Guide to f lowering p lant families. Univ. of North Carolina Press, Chapel Hill, NC

.29 Appendix 1. Plant species tested during A. subterminalis host specificity trials (excluding NZ tests). Asteraceae species are listed by family, subfamily, tribe, and subtribe with species listed alphabetically within tribes. Common names, S tate distributions, growth habits, durations, and U .S. nativities are from the USDA

– NRCS Plants Database (2008) and Canadian from Flora of Nort h America North of Mexico (1993 ) ( f rom Littlefield et al., 2008).

Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S. - State Distribution

Canada - Province Distribution Growth
Habit Duration U.S.

Nativit y Test Category 1

Asteraceae
Cichorioideae

Lactuceae

Microseridinae Agoseris glauca pale agoseris AK, AZ, CA, CO, IA, ID, MI, MN, MT, ND, NE, NM, NV, OR, SD, UT, WA, WY

AB, BC, MB, NT, ON, SK, YT Forb/herb
Shrub
Subshrub Perennial Native 3b

Asteraceae
C ichorioideae
Lactuceae

Microseridinae Agoseris grandiflora bigflower agoseris CA, ID, MT, NV, OR, UT, WA

BC Forb/herb Perennial Native 3b

Asteraceae
Cichorioideae

Lactuceae
Catananchinae Catananche caerulea Cupid’s Dart Cultivated Forb/herb Perennial Cu ltivated 3b
Ornamental

Asteraceae
Cichorioideae

Lactuceae unassigned Cichorium intybus chicory AL, AR, AZ, CA, CO, CT, DC, DE, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY

AB, BC, MB, NB, NF, NS, ON, PE, PQ, SK Forb/herb Biennial
Perennial Introduced 3b
Economic

Asteraceae
Cichorioideae

Lactuceae Crepidinae Crepis atribarba slender

hawksbeard CO, ID, MT, NE, NV, OR, UT, WA, WY

AB, BC, S K Forb/herb Perennial Native 3b

Asteraceae
Cichorioideae

Lactuceae Crepidinae Crepis biennis hawksbeard MI, NY, OH, PA, VA, VT

NF Forb/herb Biennial

Perennial Introduced 3b

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium albiflorum white hawkwe ed AK, CA, CO, ID, MT, NV, OR, SD, UT, WA, WI,

WY

AB, BC, NT, PQ, SK Forb/herb Perennial Native 2

Asteraceae Hieracium argutum southern CA Forb/herb Perennial Native 230 Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S.

- State Distribution

Canada - Province Distribution Growth
Habit Duration U

. S

. Nativit y Test Category 1

Cichorioideae

Lactuceae
Hieraciinae hawkweed

Asteraceae
Cichorioideae

Lactuceae
Hie raciinae Hieracium aurantiacum orange hawkweed AK, AR, CA, CO, CT, FL, GA, IA, ID, IL, IN, MA, MD, ME, MI, MN, MT, NC, NH, NJ, NY, OH, OR, PA, RI, SD, TN, VA, VA, VT, WA, WI, WV, WY

AB, BC, LB, MB, NB, NF, NS, ON, PE, PQ, SK Forb/herb Perennial Introduced 1

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium bolanderi Bolander’s

hawkweed CA, OR Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium caespitosum meadow
hawkweed CT, DC, GA, ID, IL, IN, KY, MA, MD, ME, MI

, MN, MT, NC, NH, NJ, NY, OH, OR, PA, RI, SC, TN, VA, VT, WA, WI, WV, WY

BC, MB, NB, NF, NS, ON, PE, PQ Forb/herb Perennial Introduced 1

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium canadense Canadian

hawkweed MI, ME

AB, BC, LB, MB, NB, N F, NS, NT, ON, PE, PQ, SK, YT Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium carneum Huachuca
hawkweed AZ, NM, TX Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium fendleri ye llow hawkweed AZ, CO, NM, TX, VT, WY Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium flagellare large
mouseeared CT, DE, IN, MA, ME, MI, NJ, NH, NY, OH, PA, VA, VT, WY

BC, NB, NS, PE, PQ Forb/herb Perennial Introdu ced 2

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium floribundum king devil CT, ID, MA, MD, ME, MI, MN, MT, NH, NJ, NY, OH, RI, VA, VT, WA, WV, WY

BC Forb/herb Perennial Introduced 131 Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S.

- State Distribution

Canada - Province Distribution Growth
Habit Duration U

. S

. Nativit y Test Category 1

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium glo meratum queen

- devil WA, ID

BC Forb/herb Perennial Introduced 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium gracile slender

hawkweed AK, CA, CO, ID, MT, NM, OR, UT, WA, WY

AB, BC, NT, YT Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae Hieracium greenei Greene’s

hawkweed CA, OR Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium gronovii queen devil AL, AR, CT, DC, DE, FL, GA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MO, MS, NC, NJ, NY, OH, OK, P A, RI, SC, TN, TX, VA, WY

ON Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium horridum prickly hawkweed CA, NV, OR Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium longiberbe longbeard

hawkweed OR, WA Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium longipilum hairy hawkweed AR, IA, IL, IN, KS, KY, LA, MI, MN, MO, NE, OH, OK, TN, TX, WI

ON Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium parryi Scouler’s woollyweed CA, ID, MT, NV, OR, WA, WY Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium pilosella mouseear hawkweed CT, GA, MA, ME, MI, MN, NC, NH, NJ, NY, OH, OR, PA, R I, TN, VA, VT, WA

BC, NB, NF, NS, ON, PQ Forb/herb Perennial Introduced 132 Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S.

- State Distribution

Canada - Province Distribution Growth
Habit Duration U

. S

. Nativit y Test Category 1

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium piloselloides yellow devil

hawkweed CT, DE, GA, IA, IL, IN, MA, ME, MI, MN, MT, NC, NH, NJ, NY, OH, PA, RI, VA, VT, WA

BC, NB, NF, NS, ON, PE, PQ Forb/herb Perennial Introduced 1

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium scabrum rough hawkweed AR, CT, DC, DE, GA, IA, IL, IN, KY, MA, MD, ME, MI, MN, MO, NC, NH, NJ, NY, OH, OK, PA, RI, SC, TN, VA, VT, WI

NB, NS , ON, PE, PQ Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium scouleri var. albertinum western
hawkweed ID, MT, OR, WA, WY

BC Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium scou leri var. cynoglossoides houndstongue
hawkweed CA, ID, OR, WA

BC Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium scouleri var. scouleri Scouler’s woollyweed CA, ID, MT, NV, OR, WA, WY Forb/herb Perennial Native 2

As teraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium stoloniflorum Not Present in U nited S tates or Canada 2

Asteraceae
Cichorioideae

Lactuceae

Hieraciinae Hieracium umbellatum narrowleaf

hawkweed AK, CO, IA, ID, IL, IN, MI, MN, MO, MT, ND, NE, NH, OR, SD, WA, WI, WV, WY

AB, BC, LB, MB, NB, NF, NS, NT, ON, PE, PQ, SK, YT Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae
Hieraciinae Hieracium venosum rattlesnakeweed AL, CT, DC, DE, FL, GA, IN, KY, MA, MD, ME, MI, MO, MS, NC, NH, NJ, NY , OH, PA, RI, SC, TN, VA, VT, WY

ON Forb/herb Perennial Native 2

Asteraceae
Cichorioideae

Lactuceae Hypochaeris radicata hairy catsear AK, AL, AR, CA, CO, CT, DC, DE, FL, GA, HI, ID, IL, IN, KY, LA, MA, MD, ME, MI, MO, MS, MT, NC, NH, NM, NJ, NV, NY, OH, OR, PA, Forb/herb Perennial Introduced 3b33 Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S.

- State Distribution

Canada - Province Distribution Growth
Habit Duration U

. S

. Nativit y Test Category 1

Hypochaeridinae RI, SC, TN, TX, UT, VA, VT, WA, WI, WV

BC, LB, NF

Asteraceae
Cichorioideae

Lactuceae
Microseridinae Krigia biflora twoflower
dwarfdandelion AL, AR, AZ, CO, CT, DE, GA, IA, IL, IN, KS, KY, MA, MD, MI, MN, MO, MS, NC, NJ, NM, NY, OH, OK, PA, TN, VA, WI, WV

ON Forb/herb Perennial Native 3b

Asteraceae
Cichorioideae

Lactuceae

Lactucinae Lactuca sativa garden lettuce AL, CA, DC, DE, ID, IL, IN, MA, ME, MI, MO, ND, NM, NY, OH, OK, OR, PA, WA, WV,

Puert o Rico, Virgin Islands Forb/herb Annual

Biennial
Perennial Introduced 3b

Economic

Asteraceae
Cichorioideae

Lactuceae

Lactucinae Lactuca serriola prickly lettuce AL, AR, AZ, CA, CO, CT, DC, DE, FL, GA, HI, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, M O, MS, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY

AB, BC, MB, NB, NS, ON, PE, PQ, SK Forb/herb Annual

Biennial Introduced 3b

Asteraceae
Cichorioideae

Lactuceae

Lactucinae Lactuca virosa bitter let tuce AL, CA, DC Forb/herb Annual

Biennial Introduced 3b

Asteraceae
Cichorioideae

Lactuceae Stephanomeriinae Lygodesmia juncea rush skeletonplant AR, AZ, CO, IA, ID, IN, KS, MN, MO, MT, ND, NE, NM, NV, OK, OR, SD, TX, UT, WA

AB, BC, MB, SK Forb/herb Peren nial Native 3b

Asteraceae
Cichorioideae

Lactuceae

Microseridinae Microseris nutans nodding

microseris CA, CO, ID, MT, NV, OR, SD, UT, WA, WY

AB, BC Forb/herb Perennial Native 3b

Asteraceae
Cichorioideae

Lactuceae

Lactucinae Prenanthes racemosa purple

ra ttlesnakeroot CO, IA, IL, IN, KY, ME, MI, MN, MO, MT, ND, NE, NJ, NY, OH, PA, SD, WI, WY

AB, BC, MB, NB, NF, NS, ON, PE, PQ, SK Forb/herb Perennial Native 3b

Asteraceae
Cichorioideae

Lactuceae Sonchus asper spiny sowthistle AK, AL, AR, AZ, CA, CO, CT, DC, DE, FL, GA, HI, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, Forb/herb An nual Introduced 3b34 Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S.

- State Distribution

Canada - Province Distribution Growth
Habit Duration U

. S

. Nativit y Test Category 1

Sonchinae NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY, Puerto Rico

AB, BC, LB, MB, NB, NF, NS, ON, PE, PQ, SK,

YT

Asteraceae
Cichorioideae

Lactuceae
Sonchinae Sonchus arvensis field sowthistle AK, CA, CO, CT, DC, DE, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OH, OR, PA, RI, SD, TN, TX, UT, VA, VT, W A, WI, WV,

WY

AB, BC, MB, NB, NF, NS, NT, ON, PE, PQ, SK, YT Forb/herb Perennial Introduced 3b

Asteraceae
Cichorioideae

Lactuceae
Sonchinae Sonchus oleraceus common sowthistle AK, AL, AR, AZ, CA, CO, CT, DC, DE, FL, GA, HI, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY, Puerto Rico, Virgin Islands

AB, BC, MB, NB, NF, NS, ON, PE, PQ, SK,

YT Forb/herb Annual Introduced 3b

Asteraceae
Cichorioideae

Lactuceae Stephanomeriinae Stephanomeria

cichoriacea chicoryleaf

wirelettuce CA Forb/herb
Subshrub Perennial Native 3b

Asteraceae
Cichorioideae

Lactuceae Stephanomeriinae Stephanomeria minor lesser wirelettuce AZ, CA, CO, ID, MT, ND, NM, NV, OR, TX, UT,

W A, WY

BC, SK Forb/herb
Subshrub Perennial Native 3b

Asteraceae
Cichorioideae

Lactuceae Crepidinae Taraxacum officinale common dandelion AK, AL, AR, AZ, CA, CO, CT, DC, DE, FL, GA,HI, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE , NH, NJ, NM, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY, Puerto
Rico

AB, BC, LB, MB, NB, NF, NS, NT, NU, ON, PE, Forb/herb Perennial Native
Introduced 3b35 Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S.

- State Distribution

Canada - Province Distribution Growth
Habit Duration U

. S

. Nativit y Test Category 1

PQ, SK, YT

Asteraceae
Cichorioideae
Cardueae
Centaureinae Acroptilon re pens hardheads AR, AZ, CA, CO, IA, ID, IL, IN, KS, KY, MI, MN, MO, MT, ND, NE, NM, NV, OH, OK, OR, SD, TX, UT, WA, WI, WY

AB, BC, MB, ON, SK Forb/herb Perennial Introduced 3c

Asteraceae
Cichorioideae
Cardueae
Carduinae Cirsium arvense Canada thistle AK, AL, AR, AZ, CA, CO, CT, DC, DE, IA, ID, IL, IN, KS, KY, MA, MD, ME, MI, MN, MO, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OH, OR, PA, RI, SD, TN, UT, VA, VT, WA, WI, WV, WY

AB, BC, MB, NB, NF, NS, NT, ON, PE, PQ, SK, YT Forb/herb Perennial Introduced 3c

Aster aceae Cichorioideae
Cardueae
Carduinae Cynara scolymus globe artichoke AZ, CA Forb/herb Perennial Introduced 3c

Asteraceae
Asteroideae
Anthemideae

Artemisiinae Artemisia dracunculus tarragon AK, AZ, CA, CO, IA, ID, IL, KS, MA, MN, MO, MT, ND, NE, NM, NV, NY, OK, OR, SD, TX, UT, WA, WI, WY

AB, BC, MB, SK, YT Forb/herb Perennial Native 3d
Economic

Asteraceae
Asteroideae

Anthemideae Chrysantheminae Chrysanthemum
cinerariifolium pyrethrum Cultivated Forb/herb Perennial Cultivated 3d

Asteraceae
Asteroideae Eupatorieae Eupatoriinae Eupatoriadelphus (Eupatorium)

maculatum spotted trumpetweed CT,GA, IA, IL, IN, KY, MA, MD, ME, MI, MN, NC, NH, NJ, NY, OH, PA, RI, TN, VA, VT, WI,

WY

MB, NB, NS, ON, PE, PQ Forb/herb Perennial Native 3d

Asteraceae

Asteroideae
He lenieae

Pectidinae Tagetes erecta Aztec marigold AR, CA, CT, FL, KY, LA, MA, MD, MO, NC, NY, OH, OK, PA, SC, UT, VA, Puerto Rico, Virgin Islands Forb/herb Annual Introduced 3d
Ornamental

Asteraceae

Asteroideae
Heliantheae Dahlia sp. dahlia Cultivated Forb/herb
Subshrub Perennial Cultivated 3d
Ornamental36 Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S.

- State Distribution

Canada - Province Distribution Growth
Habit Duration U

. S

. Nativit y Test Category 1

Coreopsidinae

Asteraceae
Asteroideae

Heliantheae
Helianthinae Helianthus annuus common sunflower AK, AL, AR, AZ, CA, CO, CT, DC, DE, FL, GA, HI, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY, Puerto Rico

AB, BC, MB, NB, NF, NS, NT, ON, PE, PQ, SK Forb/herb Annual Native 3d
Ornamental

Economic

Asteraceae

Asteroideae
Vernonieae

Vernoniinae Stokesia laevis Stokes’ Aster AL, FL, GA, LA, MS, NC, SC Forb/herb Perennial Native 3d
Ornamental

Asteraceae Gnaphalioideae Gnaphalieae
Cassiniinae Antennaria dioica stoloniferous pussytoes AK Forb/herb Perennial Native 3d

Asteraceae Gnaphalioideae Gnaphalieae Gnaphaliinae Gnaphalium audax Not Present in U nited S tates or Canada

Liliaceae Allium cepa garden onion AR, CA, CT, IL, IN, KS, KY, LA, MA, ME, MI, MO, MS, MT, NC, NH, NY, OH, OR, PA, SC, TX, VT, WI Forb/herb Perennial Introduced 5
Economic

Asclep iadaceae Asclepias syriaca common milkweed AL, AR, CT, DC, DE, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NH, NE, NJ, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, VA, VT, WI, WV Forb/herb Perennial Native 5

Chenopodiaceae Beta vulgaris common beet AL, CA, CT, MA, ME, MI, MO, MT, NC, NH, NY, OR, PA, SC, TX, UT, VA, WV Forb/herb Annual

Biennial Introduced 5
Economic

Lamiaceae Monarda fistulosa wild bergamot AL, AR, AZ, CO, CT, DC, DE, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, M O, MS, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY Forb/herb
Subshrub Perennial Native 5

Papaveraceae Papaver nudicale Iceland poppy Cultivated

AK Forb/herb Biennial
Perennial Cultivated 5
Ornamen tal37 Family Subfamily

Tribe
Subtribe Scientific Name Common Name U.S.

- State Distribution

Canada - Province Distribution Growth
Habit Duration U

. S

. Nativit y Test Category 1

YT
Apiaceae Petroselinum crispum parsley AR, CA, CT, FL, GA, HI, IA, ID, KS, LA, MA, MD, MI, MS, MT, NC, NV, NY, OH, PA, RI, TX, SC, UT, WA, Puerto Rico Forb/herb Annual
Biennial Introduced 5
Economic

Fabaceae Pisum sativum garden pea CA, CT, FL, IL, KS , KY, LA, MA, ME, MI, MO, MS, NC, NH, NY, OK, OR, PA, SC, UT, VA,

WA Forb/herb

Vine Annual Introduced 5
Economic

Fabaceae Vicia faba horsebean CA, CT, DC, MA, MD, ME, MT, NY, OR, PA, VA, VT, WA Forb/herb

Vine Annual Introduced 5
Economic

1 Test Categories: 1. Genetic types of target weed; 2. Species of the same (or closely related) genus; 3. Species in the same family as the target weed (3a. Plants in same subtribe; 3b. Plants in same tribe; 3c. Plants in same subfamily; and 3d. Plants in same family); 4. Threatened and endangered species in the same family; and 5. Species in other families.38 Appendix 2. Description of plants chosen for each category for host specificity testing (f rom Littlefield et al., 2008)

CATEGORY 1: Genetic types of **Hieracium** ( varieties, races, forms, genotypes, apomicts, etc.).

The natural host of A. subterminalis is mouse- ear hawkweed, H. pilosella . Although specific genotypes of H. pilosella were not tested, plants from Europe, New Zealand, and the United States (Washington ) were tested. In general , a high proportion of these plant collections supported gall development (54 to 97 percent), although infestation levels probably varied due to testing conditions. Since galls were also induced in other species within the subgenus

Pilosella

, it is thought that A. subterminalis is not highly specialized or limited to a particular H. pilosella genotype. Host specificity tests conducted for New Zealand indicated that orange hawkweed, H. aurantiacum , also supported gall development and the emergence of viable adults. Orange hawkweed is more widespread and abundant in the western United

States and Canada and

, therefore

, a more important target weed. North American testing also indicated that this plant host is suitable, but perhaps less so compared to H. pilosella

. Infestation levels varied from 13 to 67 percent. Although H. pilosella appears to be genetically diverse (Trewick et al. , 2004), H. aurantiacum has a very limited genetic diversity, especially for North American populations (Loomis, 2007). Therefore, Littlefield

et al. (2008) speculate that differences observed in infestations levels were due more to environmental conditions than genotypic differences in H. aurantiacum

.

CATEGORY 2: North American species in the same genus as **Hieracium** , divided by subgenera, including economically and environmentally important plants.

2.a. Species in the same subgenus ( **Pilosella** ):

This subgenus is not represented in the native flora of North America

- all members of this subgenus (or genus) are European. For North American testing H. caespitosum

, H. glomeratum

, H. piloselloides

, H. stoloniflorum

, H. flagellare

, and H. floribundum were included. Hieracium praealtum was also tested in New Zealand but was not attacked. Galls were induced only on H. flagellare and H. floribundum . Gaskin and Wilson (2007) indicated two distinct species grouping within the Pilosella

. Hieracium flagellare is more closely

related to both H. pilosella and H. aurantiacum

. Hieracium floribundum however, is mo re closely related to the group containing H. caespitosum

, H. glomeratum , and H. piloselloides

. Although H. floribundum was attacked in tests conducted at MSU, it was not infested in

CABI tests. Hieracium floribundum is thought to be a cross between H. c aespitosum and H.

lactucella , although back crosses with other Hieraciums (e.g. H. aurantiacum ) sometimes occur (Krahulec et al. , 2008). H ieracium stoloniflorum is another cross ( H. aurantiacum x H. pilosella ) (Krahulec et al. , 2008), although based upon host utilization of H. pilosella and

H. aurantiacum in the host specificity tests H. stoloniflorum was expected to be a potential host, but no gall induction was observed.39 2.b. Species in the subgenus **Hieracium** :

From the subgenus Hieracium , the North American representative to this subgenus was tested, H. umbellatum , and no gall development occurred. Although H. canadense has previously been placed in this subgenus, recent work by Gaskin and Wilson (2007) placed this species in the subgenus Chionoracium . Other representatives ( H. lepidulum

, H.

murorum , and H. sabaudum ) to this subgenus were tested by CABI and in New Zealand, but not attacked by A. subterminalis

.

The subgenus Hieracium is poorly represented in North America. Hieracium umbellatum is Holarctic in distribution and is thought to be a native. Other subgenus Hieracium species in

North America include H. argillaceum

, H. murorum

, H. groenlandicum , and H. sabaudum ; all of which are introduced. However, these species are generally not considered noxious

weeds.

Based on host specificity testing , it is unlikely that any plant in this subgenus would be utilized as a host by A. subterminalis since none are stoloniferous (produce stolons). A.

subterminalis attacks and causes galls only in stol ons and not other plant parts.

2.c. Species in the subgenus **Chionoracium** :

Subgenus Chionoracium is represented throughout North America. Sixteen species representative of the geographic range of native species were tested

. From the Northwest

and Inte rmountain region, H. gracile, H. longiberbe

, H. parryi, and H. scouleri var. albertinum and scouleri, and the white flowered H. albiflorum were tested . From southern Oregon and northern California, H. bolanderi

, H. horridum , and H. greenei were used. Fro m California, H. argutum was tested. From the Southwest, H. carneum and H. fendleri were tested , and from the East and Midwest, H. longipilum

, H. gronovii

, H scabrum , and H.

venosum were tested

. Hieracium canadense was also tested and its distributions e xtend over

much of the northern United States and southern Canada. None of these species were infested by A

. subterminalis and none of the hawkweed species associated with the subgenus

Chionoracium produces stolons in which the gall wasp induces galls. It is highly unlikely

that A

. subterminalis would utilize these plants as hosts.

CATEGORY 3: North American species in other genera in the Asteraceae family, divided by subtribe, tribe, and subfamily, including economically and environmentally important pl ants.

The Asteraceae is the largest plant family and is divided into 3 subfamilies, 17 tribes, and numerous subtribes. In developing our list of plants to be tested, we followed the reclassification of the Asteraceae by Bremer (1994).40 3.a. Species in the same subtribe (Hieraciinae) as **Hieracium** :

Other genera in the Hieraciinae subtribe include Andryala, Arnoseris, Hispidella, Hololeion,

and Tolpis

. No species from these genera were tested since they are not native to North America, nor do they contain any economically important plants in North America (Bremer, 1994). Only two species, Arnoseris minima and Tolpis barbata are reported to occur in the

U nited States and Canada (Flora of North America North of Mexico, 2008; USDA

- NRCS

Plants Database 2008 ).

3.b. Species in different subtribes of the same tribe (Lactuceae) as **Hieracium** :

The Lactuceae is one of the best known tribes of the Asteraceae and is well supported as a distinct monophyletic group. Subtribal and generic classifications are more defined in this tribe than in most other tribes (Bremer , 1994). The Lactuceae are set apart from other Asteraceae by a ligulate capitula, milky latex, and absence of oil ducts (except in Scolymus ), although none of these characteristics are restricted to the Lactuceae alone (Mañez et. al ., 1994). Besides the Hieraciinae, there are five Lactuceae subtribes that contain species native to North America (Crepidinae, Lactucinae, Malcothrinae, Microseridinae, and Stephanomeriinae); four subtribes with introduce d species (Catananchinae, Hypochaeridinae = Leontodontinae, Scorzonerinae, and Sonchinae); and two introduced genera ( Cichorium and Scolymus ) that have not been assigned to a subtribe (Bremer

, 1994).

Two genera in the Crepidinae subtribe ( Crepis and Taraxacum ) contain species native to North America (Bremer , 1994). Crepis is an important genus to test because of the number of native and afew introduced species in the United States (Flora of North America North of Mexico 2008, USDANRCS , 2008). The two C repis species we tested were Crepis atribarba (native) and Crepis biennis (introduced). Taraxacum is another important genus to test because of the presence of native species in Western North America (USDA

- NCRS , 2008), some of which are threatened, endangered, or sensitive (see Category 4). The c osmopolitan species Taraxacum officinale was tested , and in New

Zealand

, T. magellanicum was also tested.

Within the Lactucinae, two genera contain species native to North America: Lactuca and Prenanthes (Bremer , 1994; USDA

- NRCS , 2008). Lactuca sativa (lettuce) was included because of its economic importance. In addition, Lactuca serriola and L. virosa were tested since they are widespread

weeds. Prenanthes sagittata was tested since it is a native.

The Micros eridinae contains the following genera native to North America: Agoseris

, Krigia

, Microseris

, Nothocalais

, Phalacroseris

, Pyrrhopappus

, Stebbinsoseris , and Uropappus (Bremer

, 1994; USDA

- NCRS , 2008). Four native species: Microseris nutans

, Agoseris glauca

, A

. grandifolia , and Krigia biflora were included in tests. In New Zealand, Microseris scapigera was used.

The Stephanomeriinae contains the following genera native to North America: Chaetadelpha

, Lygodesmia

, Prenanthella

, Rafinesquia

, Shinnersoseris , and Stephanomeria (Bremer , 1994; USDA

-41 NCRS , 2008). T hree native western species were tested : Lygodesmia juncea

, Stephanomeria cichoriacea, and S. minor (= S. tenuifolia ).

The Malcothrinae contains the following genera native to North America: Anisocoma

, Atrichoseris

, Calycoseris

, Glyptopleura

, Malacothrix , and Pinaropappus (Bremer , 1994; Flora of North America North of Mexico, 2008; USDA

- NCRS , 2008). Most are native to the Southwestern United States,

except Glyptopleura (western annuals) and Malacothrix (western annuals and perennials). No species from this subtribe were tested because sources for seeds and/or plant material were identified

.

Of the four subtribes with introduced species but no natives, Catanache caerula , an introduced

ornamental from th e Catananchinae was included. For the subtribe Sonchinae: Sonchus arvensis

, S.

asper , and S. oleraceus were tested; plus S. kirkii

, which was also tested in New Zealand. No species from the Hypochaeridinae, or Scorzonerinae were included, as these subtribes are not native to North America and contain no economically important introduced species (Bremer, 1994; USDA

NCRS , 2008). Among the introduced genera unassigned to a subtribe, Cichorium intybus (chicory)

was used in both New Zealand and North American tests becaus e of its economic importance.

In summary, 26 species belonging to other subtribes within Lactuceae were tested during North American and New Zealand host specificity tests. None were utilized by A. subterminalis as a host.

3.c. Species in different tribes in the same subfamily (Cichorioideae):

According to Bremer (1994), the subfamily Cichorioideae contains three tribes with natives in North America (Cardueae, Mutisieae, and Vernonieae) plus a tribe with some introduced ornamentals (Arc toteae). The Mutisieae and the Vernonieae are believed to be closely related to the Lactuceae (Mañez et al.

, 1994; Tomb , 1977). Like the Lactuceae, several species of Mutisieae and Stokesia have ligulate corollas (Tomb , 1977). Stokesia laevis , a native ornamental in the Veronieae was used for testing. From the Cardueae, we used two weedy species that can reproduce asexually by lateral roots, Acroptilon repens (Russian knapweed) and Cirisium arvense (Canada thistle), as well as Cynara scolymu s (globe

art ichoke). Both Carthamus tinctorius (safflower) and globe artichoke were tested from this subfamily for the New Zealand tests. Galls were not induced by A. subterminalis on any of these species.

3.d. Species in different subfamilies in the same family ( Asteraceae):

Besides the Cichorioideae, there are two additional subfamilies (Barnadesioideae and Asteroideae) in the Asteraceae family (Bremer , 1994).

The Barnadesioideae comprise nine genera and are exclusively South American. Because

there are no s pecies native to North America (Bremer , 1994), no Barnadesioideae were

included in testing.42 The Asteroideae include the majority of the Asteraceae and comprise eight tribes with species native to the United States (Anthemideae, Astereae, Eupatorieae, Gnaphalieae, Helenieae, Heliantheae, Plucheeae, and Senecioneae), two additional tribes with ornamental species (Calenduleae and Inuleae), and eight genera unassigned to a tribe (Bremer , 1994). Although acetylenes are relatively rare in the Lactuceae, those which are present belong to a special group which may link the Lactuceae chemotaxonomically with the Astereae and the Anthemideae (Mabry and Bohlmann, 1977). In combined North American and New Zealand tests, 12 species were used representing the eight subtribes of the Asteraceae. Galls were not induced on any of these species.

Category 3 Summary : From testing these 42 species in all of Category 3, Littlefield et al. (2008) obtained sufficient evidence to conclude that the feeding range of this gall w asp is restricted to the genus Hieracium , subgenus Pilosella , and that species in other subfamilies of the Asteraceae family are not at significant risk.

C ATEGORY 4: Threatened and endangered species in the Asteraceae family, divided by subgenus, genus, subfamily, and tribe.

4.a. Species in the same subgenus ( **Pilosella** ):

There are no Hieracium species belonging to subgenus Pilosella listed as threatened, endangered, or sensitive.

4.b. Species in the same genus as **Hieracium** :

While there are no threa tened or endangered species of Hieracium in the United States (or Canada), several states identify one or more Hiearcium species as being restricted in

distribution.

The following is a list of S tates that list one or more Hiearcium species on their Spec ial Plants List (or equivalent).

Pennsylvania http://www.dcnr.state.pa.us/forestry/pndi/fullplants.asp lists H. kalmii and H.

trailii

.

Ohio http://www.dnr.state.oh.us/odnr/dnap/heritage/plantlst.html lists H. canadense and H. longipilum

.

Utah http://www.nr.state.ut.us/dwr/dwr.htm lists H. fendleri.

In Oregon, the Oregon Natural Heritage Program http://www.heritage.tnc.org/nhp/us/or/vdl.htm lists H. bolanderi

, H. greenei , and H. horridum.43 Tennessee http://www.state.tn.us/environment/nh/tnplants.html lists H. longipilum , and H.

scabrum

.

Michigan http://www.dnr.state.mi.us/wildlife/heritage/ lists H. longip ilum.

N ative Hieracium species were selected from across the geographic range of the genus throughout North America. All of the Hieracium species native to North America belong the subgenus Chionoracium, none of which were infested by A. subterminalis

.

4 .c. Species in the same subtribe (Hieraciinae) as **Hieracium** :

There are no threatened, endangered, or sensitive species within the same subtribe as

Hieracium.

4.d. Species in different subtribes of the same tribe (Lactuceae) as **Hieracium** :

Four of the Lactuceaae subtribes (besides Hieraciinae) have threatened, endangered, or sensitive species in the continental United States (USFWS , 1993; USFWS , 1996). Because obtaining plant material of threatened, endangered, and sensitive species can be difficult and can further decimate populations, listed species were not tested

. Instead , a more common member from each genus was selected as a surrogate

.

Two Crepidinae species, Taraxacum californicum Munz & Johnston, proposed endangered,

and T. officinale ssp. ce ratophorum (Ledeb.) Schinz ex. Thellung (syn. T. carneocoloratum ), sensitive, are listed in the continental Uni ted States ( http://www.fws.gov/r9endspp/endspp.html ). T he more common species Taraxacum officinale was tested (listed in Category 3.b.). The Lactucinae has two species of Prenanthes

listed as sensitive [ P. barbata (Torr. & Gray) Milstead and P. boottii (DC.) Gray]. The more common native species, Prenanthes sagittata

, was substituted (liste d in Category 3.b.). The Microseridinae has two species of Microseris listed as sensitive ( M. decipens Chambers and

M. howellii Gray). T he more common native species, Microseris nutans

, was tested (listed in Category 3.b.). The Stephanomeriinae has one federally endangered Stephanomeria ( S.

malheurensis Gottlieb), one sensitive Stephanomeria ( S. blairii Munz. & Johnston) and one

sensitive Lygodesmia species ( L. doloresensis S. Tomb). I nstead Stephanomeria minor (= S. tenuifolia ), S. cichoriacea

, and Lyg odesmia juncea were tested (listed in Category 3.b.)

4.e. Species in different tribes in the same subfamily (Cichorioideae):

The Cardueae has several Cirsium species listed as threatened, endangered, or sensitive in the

continental United States ( http://www.fws.gov/r9endspp/endspp.html ): Cirsium fontinale var. fontinale ( fountain thistle), Cirsium fontinale var. obispoense (Chorro Creek bog thistle), Cirsium hydrophilum var. hydrophilum (Suisun thi stle), Cirsium loncholepis (La Graciosa thistle), and Cirsium pitcheri (Pitcher's thistle). All but the last species is federally endangered, while C. pitcheri is threatened. Because obtaining achenes and/or plant material of threatened, endangered, and sensitive species can be difficult and can further decimate44 populations, these rare species were not tested

. Cirsium arvense

, a more common member from this genus which also reproduces asexually by lateral roots was sel e cted

.

4.f. Species in different subfamilies in the same family (Asteraceae):

At this level, no representatives from genera with threatened, endangered, or sensitive

individuals were tested . With over 20 genera with federally listed threatened or endangered species and over 70 addition al genera with sensitive species in the continental United States, the number of species we would need to test is prohibitively large. Testing was conducted on representatives to the various subtribes of Asteraceae (Category 3.d). It was evident from

MSU testing and testing conducted for New Zealand that A. subterminalis was restricted in its host range and plants outside the genus Hieracium subgenus Pilosella are not at risk of attack by this agent.

CATEGORY 5: North American species in other families in the Asterales order that have some phylogenetic, morphological, or biochemical relationship to the target weed, including economically and environmentally important plants.

Neither Cronquist nor Dahlgren list any families other than the Asteraceae in t he Asterales order; however, Thorne lists the Calyceraceae as belonging in the Asterales. Cronquist places the Calyceraceae in the Calycerales while Dahlgren places it in the Dipsacales. Since two out of three systems list the Calyceraceae in an order other than Asterales, this family is

discussed in Category 6.

CATEGORY 6:
North American species in other orders that have some morphological or biochemical relationship to the target weed, including economically and environmentally important plants.

6.a. Species in other orders that are phylogenetically related to the Asteraceae:

The Asteraceae form such a well

- defined group that they are sometimes considered systematically isolated. The most closely related families are generally considered to be the

C alyceraceae, Campanulaceae sensu lato , and Goodeniaceae (Bremer , 1994). The Calyceraceae contain six genera with about 60 species in southern South America (Bremer

, 1994). One species, Acicarpha tribuloides , is listed as present in North America (Kartesz

and Kartesz , 1980). The Campanulaceae sensu lato contain about 85 genera and more than 2, 200 species (Bremer , 1994). There are about 23 genera and 290 species in the United States and Canada (Zomlefer , 1994). The Campanulaceae have been shown to be chemotaxonomically linked to the Asteraceae (Mabry and Bohlmann, 1977). The Goodeniaceae contain 12 genera and 400 predominately Australian species (Bremer, 1994). One genus, Scaevola , has 2 species listed as present in the central and s outhern United State s (USDA

- NRCS , 2008). Due to the apparent specificity of A. subterminalis , plants within

these orders were not tested (Littlefield et al.

, 2008).45 6.b. Species in other orders that are biochemically similar to the Asteraceae:

Species in the Lactuceae are characterized, in part, by their milky latex. However, more than 12,500 species in 900 genera and 20 families have been identified worldwide that produce latex (Metcalfe , 1967). Over 300 species in the United States have been shown to contain latex (Buc hanan, 1978). The following four latex-producing species from different families

were selected for testing: Allium cepa

, Asclepias syriaca

, Monarda fistulosa L., and Papavar

nudicaule L. All are native or introduced to the w estern United States. None of these latex

producing species were attacked by A. subterminalis (Littlefield et al.

, 2008).

The Lactuceae also differ phytochemically in that their sesquiterpene lactones are predominately guaianolides, whereas the other Asteraceae subfamilies contain pl ants with considerable portions of other sesquiterpene lactones (Wapshere , 1983). Unlike most other Asteraceae, the Lactuceae possess either no or a low level of polyacetylene compounds (Sorensen , 1977). The Apiaceae (= Umbellifereae) have been shown to be chemotaxonomically linked to the Asteraceae (Mabry and Bohlmann, 1977). Petroselinum crispum (parsley), which is economically important and chemically similar to the Asteraceae

, was not infested during the New Zealand tests (Littlefield et al.

, 2008).

6.c. Selected cultivated species in other orders:

No cultivated species in other plant orders were tested, although several species were tested for the New Zealand project

.

CATEGORY 7: Any plant on which the biological control agent or its close relati ves (within the same genus) have been previously recorded to feed and/or reproduce.

Cynipid wasps of the genus Aulacidea generally have a narrow host range, confined to the family Asteraceae. A number of species utilized by other Aulacidea spp. wasps, in cluding Acroptilon

, Hieracium, Hypochoeris

, Lactuca

, Prenanthes

, Senecio

, Solidago, Sonchus , and Tragopogon, were tested. Galls were only induced on several species belonging to the

Hieracium subgenus Pilosella (Littlefield et al.

, 2008).

Conclusions

B ased upon test results from M ontana S tate U niversity , as well as testing of A. subterminalis for importation into New Zealand, host utilization by A. subterminalis is confined to a few closely related hawkweed species all of which are Old World species. Native North American hawkweeds are precluded from attack in t hat they do not produce stolons, the

location where the gall wasp induces galls and completes its development. All native North American hawkweeds are phylogentically distant and distinct from the exotic, and often weedy, hawkweeds of the Pillosella group (Littlefield et al.

, 2008).46 Appendix 3

. Galling observed in multiple

- choice and larval development tests with A.

subterminalis (combined results for New Zealand host specificity tests 1995 to 19 97; taken from Syrett et al.

, 1998).

Test Plant Species Total No. of Plants

Offered Attacked

Asteraceae

Lactuceae

Subgenus Pilosella

Hieracium pilosella L. EUR 76 74

H. pilosella L. NZ 14 10

H. aurantiacum L. 4 2

H. caespitosum Dumort EUR 6 0

H. caespitosum Dumort USA 5 0

H. praealtum Gochnat 3 0

H.x stoloniferum Waldst. et Kit 3 0

Subgenus Hieracium 0

H. lepidulum (Stenstroem) Omang 6 0

H. murorum L. 7 0

H. sabaudum L. 8 0

Representatives of other Subtribes of Lactuce ae
Cichorium intybus L. 7 0

Embergia grandifolia (Kirk) Boules 4 0

Hypochoeris radicata L 8 0

Lactuca sativa L. 4 0

Microseris scapigera (Solex A.Cunn.) Sch.Bip. 5 0

Picris hieracioides L. 7 0

Sonchus kirkii Hamlin 6 0

Sonchus oleraceus L. 6 0

T

araxacum officinale L. 5 0

Taraxacum magellanicum Schutz- Bip. 2 0

Tragopogon porrifolius L. 6 0

Anthemidae

Artemesia dracunculus L. 1 0

Chrysanthemum cinerariifolium (Treviri) Vis. 7 0

Astereae

Celmisia sp. 3 0

Olearia avicenniifolia (Raul.) Ho ok f. 4 0

Cardueae

Carthamus tinctorius L. 7 0

Cynara scolymus L. 6 0

Heliantheae

Helianthus annuus L. 6 047 Test Plant Species Total No. of Plants

Offered Attacked

Senecioneae

Brachyglottis monroi Hook. f. 2 0

Other Plant Families

Api aceae: Patroselinum crispum (Miller) A.W. Hill 8 0

Brassicaceae: Brassica oleracea L. 7 0

Cannaceae: Canna edulis Ker- Gawl. 9 0

Caryophyllaceae: Dianithus barbatus L. 7 0

Cyperaceae: Carex testacea Sol ex Booth Hook. f. 5 0

Fabaceae: Trifolium repe ns L. 7 0

Iridaceae: Gladiolus communis L. 8 0

Lamiaceae: Mentha sp. 8 0

Liliaeae: Allium cepa L. 8 0

Malvaceae: Althea rosea L. 5 0

Myrtaceae: Leptospermum scoparium J.R. et G. Forst. 3 0

Oleaceae: Olea europaea L. 2 0

Poaceae: Poa colensoi Hook. f

. 7 0

Poaceae: Agrostis tenuis Sibth. 5 0

Poaceae: Festuca novae

- zelandiae (Hack.), Ckn 7 0

Polyganaceae: Rumex acetosella L. 7 0

Rutaceae: Citrus sp. 8 0

Scrophulariaceae: Antirrhinum majus L. 2 0

Solanaceae: Lycopersicon esculentum Miller 8 0

Thea ceae: Camellia japonica L. 8 0

Urticaceae: Urtica dioica L. 7 048 Appendix 4. No

- choice oviposition and larval development tests with A.

subterminalis conducted from 2000 to 2006 at CABI

- Europe and Montana State University (Littlefield et al., 2008 ).

Test P lant 1 No. Reps Reps. Infested Mean No. Gall Clusters/Rep (+ SE) No. %

ASTERACEAE

Tribe: Lactuceae

Subtribe: Hieraciinae Subgenus: Pilosella

Hieracium aurantiacum ID 15 2 13.3 0.13 (0.09)

H. aurantiacum MT 1 0 0 0

H. aura ntiacum NZ 6 4 66.7 1.33 (0.49)

H. aurantiacum ID, MT 44 17 38.6 0.89 (0.21)

H. caespitosum ID 33 0 0 0

H. caespitosum ID 3 0 0 0

H. flagellare NH 16 8 50.0 1.62 (0.79)

H. floribundum US 9 0 0 0

H. floribundum 17 9 52.9 1.24 (0.48)

H. glomeratum BC 2 0 0 0

H. glomeratum WA 6 0 0 0

H. glomeratum ID 7 0 0 0

H. pilosella EUR 28 23 82.1 2.68 (0.42)

H. pilosella US 13 11 84.6 1.92 (0.62)

H. pilosella WA 46 25 54.3 1.35 (0.29)

H. piloselloides US 6 0 0 0

H. piloselloides 11 0 0 0

H. stoloniflorum Eur 9 0 0 0

Subtribe: Hieraciinae Subgenus: Hieracium

H. umbellatum 13 0 0 0

Subtribe: Hieraciinae Subgenus: Chionoracium

Hieracium. albiflorum 2 0 0 0

H. albiflorum 18 0 0 0

H. argutum 3 0 0 0

H. argutum 18 0 0 049 Test Plant 1 No. Reps Reps. Infested Mean No. Gall Clusters/Rep ( + SE) No. %

H. bolanderi 15 0 0 0

H. canadense 7 0 0 0

H. canadense 15 0 0 0

H. carneum 1 0 0 0

H. carneum 15 0 0 0

H. fendler i 3 0 0 0

H. fendleri 10 0 0 0

H. gracile 7 0 0 0

H. greenei 14 0 0 0

H. gronovii 3 0 0 0

H. gronovii 17 0 0 0

H. horridum 3 0 0 0

H. longiberbe 8 0 0 0

H. longipilum 15 0 0 0

H. parryi 11 0 0 0

H. scabrum 15 0 0 0

H. scouleri var. albertinum 6 0 0 0

H. scouleri var. albertinum 11 0 0 0

H. scouleri var

. cynoglossoides 17 0 0 0

H. venosum 5 0 0 0

Other Subtribes of Lactuceae

Subtribe: Catananchinae

Catananche caerula 6 0 0 0

Subtribe: Crepidinae

Crepis atribarba 10 0 0 0

Crepis biennis 15 0 0 0

Taraxacum officinale 6 0 0 0

Subtribe: Hypochaeridinae

Hypochoeris radicata 6 0 0 0

Subtribe: Lactucinae

Lactuca sativa 9 0 0 050 Test Plant 1 No. Reps Reps. Infested Mean No. Gall Clusters/Rep ( + SE) No. %

Lactuca serriola 3 0 0 0

Lactuca virosa 6 0 0 0

Prenanthes racemosa 4 0 0 0

Subtribe: Microseridinae

Agroseris glauca 16 0 0 0

Agoseris grandiflora 5 0 0 0

Agroseris grandifolia 3 0 0 0

Krigia biflora 3 0 0 0

Krigia biflora 13 0 0 0

Microseris nutans 11 0 0 0

Subtribe: Sonchi nae

Sonchus arvensis 6 0 0 0

Sonchus asper 8 0 0 0

Sonchus oleraceus 6 0 0 0

Subtribe: Stephanomeriinae

Lygodesmia juncea 14 0 0 0

Stephanomeria cichoriacea 14 0 0 0

Stephanomeria minor 5 0 0 0

Subtribe: unassigned

Cichorium int ybus 10 0 0 0

Other Tribes of Asteraceae

Tribe: Anthemideae

Artemisia dracunculus 6 0 0 0

Chrysanthemum cinerariifolium 9 0 0 0

Tribe: Cardueae

Acroptilon repens 8 0 0 0

Cirsium arvense 10 0 0 0

Cynara scolymus 9 0 0 0

Tribe: Eup atorieae

Eupatorium maculatum 9 0 0 051 Test Plant 1 No. Reps Reps. Infested Mean No. Gall Clusters/Rep ( + SE) No. %

Tribe: Gnaphalieae

Antennaria dioica 9 0 0 0

Gnaphalium audax 9 0 0 0

Tribe: Helenieae

Tagetes erecta 9 0 0 0

Tribe: Heliantheae

Dahlia sp. 3 0 0 0

Helianthus annuus 9 0 0 0

Tribe: Vernonieae

Stokesia laevis 5 0 0 0

OTHER FAMILIES

Apiaceae : Petroselinum crispum

(parsley) 3 0 0 0

Asclepiadacea : Asclepias syriaca

(common milkweed) 7 0 0 0

Chenopo diaceae : Beta vulgaris (beet) 9 0 0 0

Fabaceae : Pisum sativum (garden

pea) 7 0 0 0

Fabaceae : Vicia faba (broad bean) 5 0 0 0

Lamiaceae : Monarda fistulosa (wild

bergamot) 6 0 0 0

Liliaceae : Allium cepa (onion) 8 0 0 0

Papavaracea : Papaver nudicaule

(Is landic poppy) 3 0 0 0

1 Plant species highlighted in blue indicate those tested at Montana State University.