Research Article
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Research Article
Sawflies (Hymenoptera, Symphyta) newly recorded from Washington State
expand article infoChris Looney, David R. Smith§, Sharon J. Collman|, David W. Langor, Merrill A. Peterson#
‡ Washington State Department of Agriculture, Olympia, United States of America
§ Smithsonian Institution, Washington, United States of America
| WSU Snohomish County Extension, Everett, United States of America
¶ Natural Resources Canada, Edmonton, Canada
# Western Washington University, Bellinghamu, United States of America
Open Access

Abstract

Examination of museum specimens, unpublished collection data, and field surveys conducted between 2010 and 2014 resulted in records for 22 species of sawflies new to Washington State, seven of which are likely to be pest problems in ornamental landscapes. These data highlight the continued range expansion of exotic species across North America. These new records also indicate that our collective knowledge of Pacific Northwest arthropod biodiversity and biogeography is underdeveloped, even for a relatively well known and species-poor group of insects. Notable gaps in the knowledge of Washington State’s Symphyta remain for the Olympic Peninsula, the Cascade Mountain Range, and the arid interior of the state. Washington’s shrub-steppe appears to be particularly poorly surveyed for sawflies.

Keywords

Exotic species, range expansion, state record, museum data

Introduction

Sawflies (Hymenoptera, Symphyta) comprise 14 families worldwide, with 12 of these and about 1,000 described species known from North America (Smith 1979a, Taeger et al. 2010). Eleven families and approximately 180 named species are currently recorded from Washington State (Smith 1979a, Gibson 1980, Goulet 1986, Smith 1989, Goulet 1996, Looney et al. 2012, Schiff et al. 2012). Most sawfly species are herbivores, including leaf- and stem-mining species, chewing defoliators, wood-borers, and leaf-tying defoliators. Species in the family Orussidae are external parasitoids of wood-boring insects (Powell and Turner 1975, Deyrup 1984), and adult Tenthredo (Tenthredinidae) are commonly observed feeding upon other arthropods (Pasteels and Gregoire 1984). Some sawflies are important forestry, horticultural and agricultural pests.

The 2009 discovery of the introduced alder-feeding sawfly Monsoma pulveratum (Retzius, 1783) in the Pacific Northwest (Looney et al. 2012) provided impetus to conduct a broad sawfly survey in Washington State from 2010 through 2012. Specimens collected during that survey revealed that the Pacific Northwest range of many sawfly species is incompletely delineated, and that relatively few contributions have been made towards understanding Pacific Northwest Symphyta during recent decades. Subsequent to the two year survey, we collected data from other researchers, museum specimens, and further serendipitous discoveries. Here we report 22 species not previously known from Washington or documented only in gray literature, expanding known ranges in some cases and filling gaps in others.

Methods

New sawfly records were compiled from many sources, including regional entomological collections and recent field surveys. More than 3,500 identified and unidentified sawflies in entomology collections at the Evergreen State College, the College of Idaho, Oregon State University, the University of Idaho, Washington State University, and Western Washington University were examined for species of interest.

Field surveys from 2010 through 2012 employed sticky traps and Malaise traps (Fig. 1), with subsequent targeted and opportunistic hand collections made through 2015. Double-sided yellow, green, or white sticky traps with hot-applied adhesive were placed in various woody host plants across Washington in 2010–2012. Host material surveyed included alder, poplar, hawthorn, mountain ash, cherry, pear, apple, elderberry, and various conifers, with approximately 105 sites surveyed in Washington. Nine Malaise traps were installed in Washington west of the Cascade mountain range. Three traps were installed near ports of entry, two along the Columbia River, two in mixed-use forest stands, and two in residential areas. One Malaise trap was installed in a prairie remnant surrounded by agricultural fields in eastern Washington. Traps were installed in April, 2012, and maintained through September 2012. Opportunistic hand collections of larvae and adults were made throughout 2010–2015 as part of this study. We also present novel data for leaf-mining sawflies in Washington collected during a survey conducted in 2006 (see also Digweed et al. 2009). Lastly, some of the species recorded here were first detected by citizens reporting new pest problems (e.g., Neodiprion sertifer).

Figure 1.

Locations of yellow sticky traps and Malaise traps deployed in Washington State in 2011.

Collection data were compiled for each species, and narratives were composed that briefly describe each species’ natural history and other details. Species names follow Taeger et al. (2010). We have chosen to not alter two combinations in the Nematinae suggested by new work by Prous et al. (2014). Taxonomic changes for the North American fauna resulting from their research have not yet been made, and are best approached via a thorough examination of North American Nematinae rather than piecemeal in papers such as this. Probable combinations are noted in the narrative accompanying each species. There are numerous common names ascribed to many of the species reported on here, since so many of these species are pestiferous and conspicuous. We do not provide those names, but North American common names can be found at the websites for the Entomological Society of Canada (esc-sec.ca) and the Entomological Society of America (entsoc.org). Voucher specimens are deposited at the Northern Forestry Centre Arthropod Collection in Edmonton, Alberta (NFRC), the Washington State Department of Agriculture Collection (WSDAC), Western Washington University (WWUC), the Evergreen State College (TESC), and the National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM).

Results

Twenty-two species not previously documented in Washington State in peer-reviewed literature were detected in these field and museum surveys, primarily in western Washington (Table 1). One species was collected as by-catch in a survey for other pests, and seven species were first detected due to citizen complaints. Five species were first detected by examining unidentified material in museum collections, and the remainder were collected in sawfly surveys or general collecting. A map of collection localities for more than 1,200 sawfly specimens indicates that most collecting has been near developed towns and cities, or along major highways (Fig. 2). Remote and rugged areas are under-sampled, and very few collections have been made in the arid interior shrub-steppe.

Figure 2.

Map of specimen localities (black circles) for over 1,200 specimens collected in Washington State, including museum data and field data generated during this project. Vegetation zones are simplified from Washington’s GAP Analysis project (Cassidy et al. 1997).

Collection information for 22 sawfly species newly reported from Washington State.

County Lat N Long W Date Specimens Collection Method Collector
Xiphydria prolongata
King 47.5374 122.3040 7 Aug 2012 1♀ Japanese beetle trap D. Kitchen
Neodiprion sertifer
Whatcom 48.7659 122.4518 30 May 2008, em. ~ 20 Sep 2008 6♀, 1♂ Rrd. ex Pinus mugo L. Haines
Whatcom 48.7632 122.4505 May 2012 Mul. larvae Obs. on Pinus sylvestris C. Looney
Whatcom 48.7612 122.4482 May 2012, em. Aug 2012 2♂, 1♀ Rrd. ex Pinus mugo C. Looney
Whatcom 48.7412 122.4745 25 Jul-4 Oct 2012 Multiple ♂ Wing trap w/ Neodiprion lure C. Looney
Diprion similis
Mason 47.1978 123.0995 26 Jul 2012, em. ~2 Aug 2012 3♂, 2♀ Rrd. ex Pinus sylestris C. Looney
Thurston 47.0802 123.0203 4 Sep 2012 1 larva Obs. on Pinus contorta C. Looney
C. Fate
Thurston 47.0799 123.0203 4 Sep 2012 3 larvae Obs. on Pinus monticola C. Looney
C. Fate
Thurston 47.1056 123.0009 4 Sep 2012 2 larvae Obs. on Pinus monticola C. Looney
C. Fate
Thurston 47.0902 123.0471 4 Sep 2012 1♂,
5 larvae
Wing trap w/ D. pini lure,
Hand coll. on Pinus monticola
C. Looney
C. Fate
Thurston 47.0540 122.9254 10 Sep 2012 1♀, 1 larva Obs. on Pinus monticola C. Looney
A. Pelegrin
Whatcom 48.7412 122.4748 Aug 2012, em. 4 Jun 2013 1♂ Rrd. ex Pinus monticola C. Looney
M. Peterson
Gilpinia hercyniae
Whatcom 48.7412 122.4745 31 Jul-6 Aug 2008;
9 Jul 2011, em Aug 2011;
8 Aug 2012
1♀;
1♀;
1♀
Hand coll.;
Rrd. ex. Picea abies;
White sticky trap, Picea abies
M. Peterson
Cladius grandis
Thurston 47.0734 122.9767 9-16 May 1997 1♀ Malaise trap J. Longino
Cladius gregarious
Okanogan 48.4204 119.7115 16 Sep 2010, em. spring 2011 1♀, 1♂ Rrd. ex Populus tremuloides G. Kohler
Pristiphora geniculata
King 47.7295 122.3045 25 Jul 2009 Mul. larvae Obs. on Sorbus aucuparia S. Collman
King 47.7295 122.3045 10 Aug 2010 Mul. adults Obs. on Sorbus aucuparia S. Collman
King 47.6808 122.1106 12 Jul 2011 Mul. larvae Obs. on Sorbus aucuparia A. Clarke
King 47.4857 121.7674 7 Jul 2011, em. 11-22 Aug 2011 15♀ Rrd. ex Sorbus aucuparia K. Ripley
King 47.7724 122.3270 7 Aug 2012 Mul. larvae Obs. on Sorbus aucuparia S. Collman
King 47.4503 122.4908 6 June 2015 Mul. larvae Obs. on Sorbus aucuparia K. Ripley
King 47.4483 122.4836 6 June 2015 Mul. larvae Obs. on Crataegus douglasii K. Ripley
Snohomish 47.8626 121.8165 8 Aug 2009 Mul. larvae Obs. on Sorbus aucuparia S. Collman
Snohomish 47.9571 122.2318 1 Jul 2011 Mul. larvae Obs. on Sorbus aucuparia A. Jordan
Snohomish 47.8788 122.2240 9 Aug 2011 Mul. larvae Obs. on Sorbus aucuparia S. Collman
Snohomish 47.8788 122.2240 18 Jun 2012 Mul. larvae Obs. on Sorbus aucuparia S. Collman
Whatcom 48.7084 122.4433 11 Jul 2011 Mul. larvae Obs. on Sorbus aucuparia D. Wallesz
Whatcom 48.7427 122.4350 7 Jul 2013, em. 19-23 Aug 2013 7♀, 3♂ Rrd. ex Crataegus douglasii T. Cahill
Pristiphora rufipes
King 47.7295 122.3045 17 Sep 2010;
5 May 2012
Mul. larvae;
Mul. larvae
Hand coll. on Aquilegia sp. S. Collman
Snohomish 47.8656 121.9876 3 Sep 2013 Mul. larvae Hand coll. on Aquilegia sp. S. Collman
Snohomish 47.8788 122.2240 7 Sep 2013;
6 Nov 2014
Mul. larvae;
Mul. larvae
Hand coll. on Aquilegia sp. S. Collman
Thurston 47.0562 122.9250 24 Mar 2014 2♀, 6♂ Hand coll. on Aquilegia sp. C. Looney
Craesus alniastri
Island 47.9590 122.3607 7-26 Jul 2011 1♀ Yellow sticky trap, Alnus rubra K. Ripley
King 47.3809 122.2348 22 Sep 1976 1♀ Rrd. ex Alnus rubra D. Rhoades
Kitsap 47.4400 122.9365 22 Jun-1 Jul 2011 1♀ Yellow sticky trap, Alnus rubra K. Ripley
Pierce 47.2500 122.3502 17 Jun-16 Aug 2010 1♀ Green sticky trap, Alnus rubra D. Kitchen
Skagit 48.4204 122.4142 13 Sep 2010 1♀ Yellow sticky trap, Alnus rubra D. Maclean
Skamania 45.5763 122.1917 20 May-23 Jun 2011 1♀ Yellow sticky trap, Alnus rubra K. Sheehan
Skamania 45.6260 122.0241 15 Jun 2011 1♀ Yellow sticky trap, Alnus rubra T. Murray
Skamania 45.5870 122.1595 2-23 Jun 2011 2♀ Yellow sticky trap, Alnus rubra K. Sheehan
Skamania 45.7105 121.7801 30 Jun 2011 5♀ Yellow sticky trap, Alnus rubra T. Murray
Skamania 45.5763 122.1917 30 Aug-6 Oct 2011 1♀ Yellow sticky trap, Alnus rubra K. Sheehan
Snohomish 48.1987 122.1251 16 Aug-29 Aug 2011 2♀ Japanese beetle trap R. Taylor
Whatcom 48.9970 122.2635 14-26 Jun 2012 2♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9940 122.6876 26 Jul-20 Aug 2012 1♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 49.0019 122.7547 26 Jul 2012 1♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 49.0013 122.7493 26 Jul 2012 1♀ Malaise trap C. Looney
Whatcom 48.9988 122.2684 26 Jul-20 Aug 2012 4♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.7479 122.4343 1-21 Aug 2012 1♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9131 122.5741 8-20 Aug 2012 10♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9915 122.5294 8-20 Aug 2012 6♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9388 122.4443 8-20 Aug 2012 7♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9636 122.3675 8-20 Aug 2012 4♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9965 122.2632 8-20 Aug 2012 2♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9970 122.2635 8-20 Aug 2012 6♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9940 122.6878 20-30 Aug 2012 4♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9933 122.5874 20-30 Aug 2012 4♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9131 122.5741 20-30 Aug 2012 3♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9915 122.5294 20-30 Aug 2012 4♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9619 122.5091 20-30 Aug 2012 1♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9357 122.4817 20-30 Aug 2012 1♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9388 122.4443 20-29 Aug 2012 5♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9637 122.3675 20-30 Aug 2012 5♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9988 122.2684 20-30 Aug 2012 4♀ Yellow sticky trap, Alnus rubra D. Maclean
Nematus lipovskyi
King 47.7295 122.3046 12 Jun 2008 Mul. larvae Hand coll. on Rhododendron mollis S. Collman
King 47.7632 122.3147 9 Jun 2008 Mul. larvae Hand coll. on Rhododendron occidentalis S. Collman
King 47.6367 122.2966 12 Jun 2011 Mul. larvae Hand coll. on Rhododendron occidentalis S. Collman
King 47.6569 122.2899 Jun 2013 Mul. larvae Hand coll. on Rhododendron sp. S. Collman
King 47.7632 122.3147 30 May 2014 Mul. larvae Hand coll. on Rhododendron occidentalis C. Looney
A. Pelegrin
Thurston 47.0393 122.7985 26 Apr 1996 1♀ Hand coll. B. Dightman
Thurston 47.0329 122.8992 28 May 2014 Mul. larvae Obs. on Rhododendron sp. C. Looney
Thurston 47.0384 122.8984 2 Jun 2014;
14 Apr 2015
Mul. larvae;
10♀
Hand coll. on Rhododendron sp. C. Looney
Heterarthrus nemoratus
Whatcom 48.8053 121.8936 22 Jul 1967 1♀ unknown unknown
King 47.5085 122.3095 1 May 2007 1 sex unk. Photograph, bugguide C. Moorehead
Heterarthrus vagans
Whatcom 48.9353 122.4817 2 Sep 2011 1 pupa Hand coll. on Alnus rubra W. Hellman
Whatcom 48.9939 122.6830 2 Sep 2011 1 pupa Hand coll. on Alnus rubra W. Hellman
Whatcom 48.9623 122.5085 2 Sep 2011 1 pupa Hand coll. on Alnus rubra W. Hellman
Whatcom 48.9636 122.3675 14-26 Jun 2012 3♂, 1♀ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9381 122.4428 26 Jun-12 Jul 2012 4♂ Yellow sticky trap, Alnus rubra D. Maclean
Whatcom 48.9636 122.3675 8-20 Aug 2012 1♂ Yellow sticky trap, Alnus rubra D. Maclean
Metallus lanceolatus
Thurston 47.0734 122.9767 15-22 Aug 1997 1♀ Malaise trap J. Longino
Thurston 47.0802 122.9749 Jun-Aug 2012 Mul. larvae Mines in Geum macrophyllum C. Looney
King 47.6538 122.1098 Jul 2014 Mul. larvae Mines in Geum macrophyllum C. Looney
King 47.5583 122.2503 22 Aug 2015 Mul. larvae Mines in Geum macrophyllum C. Looney
Fenusella nana
Grays Harbor 47.0565 123.2739 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Grays Harbor 46.9826 123.6043 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Grays Harbor 46.9755 123.8670 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
King 47.9820 122.1947 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
King 47.7449 122.3424 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
King 47.5067 122.2900 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
King 47.5285 121.8760 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
King 47.6579 122.1105 5 May 2013 2♀ Hand coll. on Betula sp. C. Looney
King 47.4886 121.7946 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Pierce 47.2505 122.2896 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Skagit 48.4046 122.3315 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Skagit 48.4838 121.5991 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Skagit 48.5270 121.4420 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Snohomish 47.8562 121.6960 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Whatcom 48.9640 122.4625 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Whatcom 48.7949 122.4833 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Snohomish 48.2003 122.1266 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Snohomish 48.2462 121.6066 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Profenusa thomsoni
King 47.5285 122.8760 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
King 47.4886 121.7946 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Skagit 48.4046 122.3315 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Skagit 48.5270 121.4420 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Snohomish 47.8562 121.6960 2006 Mul. larvae Hand coll. on Betula sp. D. Langor
Whatcom 48.8053 121.8936 15 Jul 1967 1♀ Unknown Unknown
Profenusa inspirata
Yakima 46.7157 120.8633 7 Jun 2015 Mul. larvae Hand coll./obs. on Quercus garryana C. Looney
Yakima 46.7441 120.7884 8 Jun 2015 Mul. larvae Hand coll./obs. on Quercus garryana C. Looney
Skamania 45.7182 121.4746 24 Sep 2015 Mul. larvae Hand coll./obs. on Quercus garryana C. Looney
T. Murray
Lewis 46.6451 123.0198 2 Oct 2015 Mul. larvae Hand coll./obs. on Quercus garryana C. Looney
M. Freeman
Fenusa ulmi
King 47.6379 122.2961 20 Apr-9 May Mul. ♀ Hand coll. near Ulmus carpinifolia C. Scannell
Lewis 46.5523 122.8126 20 Jul-11 Aug 2011 3♀ Yellow sticky trap, Alnus rubra D. Kitchen
San Juan 48.7017 122.9136 12 Jun 2015 Mul. larvae Obs. on Ulmus sp. C. Looney
Thurston 47.0415 122.8617 Jun 2012 Mul. larvae Obs. on Ulmus sp. C. Looney
Thurston 47.0379 122.8991 11 Jun 2014 Mul. larvae Hand coll. on Ulmus sp. E. Spurrier
Thurston 46.8716 122.9116 5-13 May 2010 1♀ Yellow sticky trap, Alnus rubra E. LaGasa
Halidamia affinis
Whatcom 48.9061 122.4991 6 Jun 1989 2♀ Hand coll. E. LaGasa
Ferry 48.6091 118.138 6-27 Jun 2011 1♀ Yellow sticky trap, Alnus rubra M. Johnson
San Juan 48.5514 123.0781 1 Apr 2010 1♀ Yellow sticky trap, Alnus rubra T. Hanson
Clallam 48.0851 124.2636 29 Jun-19 Jul 2011 1♀ Yellow sticky trap, Alnus rubra G. Kohler
Jefferson 47.9227 122.8156 18 May-7 Jun 2011 1 sex unk. Yellow sticky trap, Alnus rubra G. Kohler
King 47.4558 122.4529 5 May 2015 2 sex unk. Hand coll C. Looney
King 47.4473 122.4599 3-13 May 2010 1♀ Yellow sticky trap, Alnus rubra K. Ripley
Kitsap 47.44 122.9365 26 May-4 Jun 2011 1 sex unk. Yellow sticky trap, Alnus rubra K. Ripley
Kitsap 47.4325 122.6126 26 May-4 Jun 2011 1 sex unk. Yellow sticky trap, Alnus rubra K. Ripley
King 47.3766 122.2418 12 Apr 2010 1♀ Sweep net J. Cena
Pierce 47.2607 122.3513 2-16 May 2012;
16 May-7 Jun 2012
3 sex unk.;
1 sex unk.
Malaise trap C. Looney
Thurston 47.0783 122.9732 18 May 2011 1♀ Sweep net C. Looney
Thurston 47.0734 122.9767 9-16 May 1997 3 sex unk. Malaise trap J. Longino
Thurston 47.0231 122.9089 8 Jun-1 Aug 2011 1 sex unk. Emerald Ash Borer trap D. Kitchen
Thurston 47.0026 123.0002 24 May 2011 1 sex unk. Yellow sticky trap, Alnus rubra C. Looney
Grays Harbor 46.9738 123.2945 5-27 May 2010 2♀ Yellow sticky trap, Alnus rubra G. Kohler
Thurston 46.8716 122.9116 10-27 May 2011;
2-14 Jun 2011
2 sex unk.;
3 sex unk.
Green sticky trap, Alnus rubra E. LaGasa
Thurston 46.8207 123.1162 27 May-14 Jun 2010 8 sex unk. Yellow sticky trap, Alnus rubra K. Ripley
Pacific 46.5204 123.887 4-18 May 2011 2 sex unk. Yellow sticky trap, Alnus rubra D. Kitchen
Lewis 46.4497 122.7989 18 May-1 Jun 2011 5 sex unk. Yellow sticky trap, Alnus rubra G. Kohler
Cowlitz 46.1103 122.8945 24 May-1 Aug 2011 1 sex unk. Emerald Ash Borer trap D. Kitchen
Clark 45.8623 122.7467 18 May-1 Jun 2011;
1-15 Jun 2011
8 sex unk.;
7 sex unk.
Yellow sticky trap, Alnus rubra G. Kohler
Skamania 45.8473 121.4122 29 Jun 2012 1♀ Malaise trap J. Markgraf
Skamania 45.8042 121.9348 19 May-23 Jun 2011 1 sex unk. Yellow sticky trap, Alnus rubra K. Sheehan
Clark 45.8004 122.6811 4 May-8 Jun 2011 1 sex unk. Yellow sticky trap, Alnus rubra K. Sheehan
Clark 45.7997 122.6818 31 Mar-4 May 2011;
4 May-8 Jun 2011
1 sex unk.;
6 sex unk.
Yellow sticky trap, Alnus rubra K. Sheehan
Skamania 45.7106 121.6395 16 Jun 2011 1 sex unk. Yellow sticky trap, Alnus rubra T. Murray
Skamania 45.7105 121.7801 16-30 Jun 2011 1 sex unk. Yellow sticky trap, Alnus rubra T. Murray
Skamania 45.6257 122.0241 31 May 2011 3 sex unk. Yellow sticky trap, Alnus rubra T. Murray
Skamania 45.6142 122.115 19 May-23 Jun 2011 1 sex unk. Yellow sticky trap, Alnus rubra K. Sheehan
Clark 45.6053 122.5459 23 Mar 2010 1♀ Hand collected A. Karankou
Skamania 45.587 122.1595 2-23 Jun 2011 2♀ Yellow sticky trap, Alnus rubra K. Sheehan
Skamania 45.5763 122.1917 19 Apr-19 May 2011 2 sex unk. Yellow sticky trap, Alnus rubra K. Sheehan
Monophadnus pallescens
Chelan 47.3120 120.2811 15 May 1999 1♀ Unknown D. Knutson
Chelan 47.2802 120.1865 4 May 2003 1♀ Unknown R. MacLean
Grays Harbor 46.9954 123.5951 28 Apr 2010 1♀ Hand coll. C. Looney
King 47.3637 122.1202 20 Apr 1985 1♀ Unknown P.E. Kalina
San Juan 48.4924 122.8944 10 May 1987 1♀ Unknown D. Overdorff
Thurston 46.8716 122.9116 2-13 May 2012 2♀ Malaise trap E. LaGasa
Thurston 47.0734 122.9767 18-25 Apr 1997;
25 Apr-2 May 1997;
2-9 May 1997;
9-16 May 1997
1♀;
1♀;
3♀;
1♀
Malaise trap J. Longino
Whatcom 48.4534 122.2918 15 May 1986 1♀ Unknown W. R. Buce
Whatcom 48.9465 122.4521 2 Jun 1967 2♀ Unknown Unknown
Whatcom 48.7502 122.4750 28 May 1975 1♀ Unknown F. Robertson
Whatcom 48.9974 122.7278 18 Apr-11 May 2012 1♀ Yellow sticky trap, Alnus rubra W. Hellman
Eupareophora parca
King 47.6538 122.1098 5 May 2015 Mul. larvae Hand coll. on Fraxinus sp. C. Looney
Monostegia abdominalis
King 47.5732 121.8856 26 Jun 2013 Mul. larvae Obs. on Lysimachia vulgaris K. Wal
King 47.6284 121.9334 Jun 2013 Mul. larvae Obs. on Lysimachia vulgaris K. Wal
King 47.5565 122.0735 11 Jun 2014, em. 13 Jul 2014 6♀ Reared ex. Lysimachia vulgaris K. Wal
King 47.5701 122.0948 11 Jun 2014 Mul. larvae Obs. on Lysimachia vulgaris K. Wal
King 47.6532 122.1070 30 May 2014, em. 6 Jul 2014 3♀ Reared ex. Lysimachia vulgaris C. Looney
A. Pelegrin
Macrophya puntumalbum
Whatcom 48.7596 122.4882 20 May 1977 1♀ Hand coll. D. Manley

Xiphydriidae

Xiphydria prolongata (Geoffroy, 1785)

Xiphydria prolongata is a European species that is a wood-borer in small limbs of deciduous tree genera, including Salix L., Quercus L., and numerous Betulaceae (Smith 1983). Larvae generally bore in decaying wood, and are not pestiferous. This western European species was first documented in North America in Michigan and New Jersey in the early 1980s, bringing the known Xiphydria species in North America to nine (Smith 1983). Mudge et al. (2001) recorded the first west coast specimen from Multnomah County, Oregon, in 1989. The single specimen from Washington was found in a trap for Japanese beetle, near Boeing Field in King County (Fig. 3). The specimen is housed at WSDAC.

Figure 3.

Localities of Xiphydriidae and Diprionidae newly detected in Washington State.

Diprionidae

Neodiprion sertifer (Geoffroy, 1785)

A member of Ross’s (1955) sertifer species group, this species is the only Neodiprion native to Europe, where it is a forestry and nursery pest (Day and Leather 1997). Its primary hosts are Pinus resinosa Aiton and P. sylvestris L., although it will feed on most Pinus species and can be a significant source of damage in pine plantations during outbreak years (Alford 2012). Introduced to New Jersey, USA, in 1925 (Schaffner 1939), its more visible impacts have included feeding damage on Christmas trees, making them less valuable or unmarketable, and defoliation of older needles on ornamental and landscape plants (Wilson 1971). Economic damage in North America has been recorded on P. strobus L., P. sylvestris, and P. mugo Turra, well as several native North American Pinus spp. (Schaffner 1943, Craighead 1950, Benjamin et al. 1955, Griffiths 1959, Baker 1972). Since its introduction, the species has spread westward at least to North Dakota (Van Driesche et al. 2012) and Saskatchewan, Canada (Langor, unpublished data).

Neodiprion sertifer was detected in Washington State in 2008, when citizens in Bellingham, Washington, alerted pest control professionals to several defoliating outbreaks on ornamental pines. Larvae were observed again in 2012 on P. sylvestris and P. mugo in Bellingham, and adult males of Neodiprion sp. were collected in pheromone traps in the city that year. Although the latter were likely N. sertifer, male Neodiprion are not readily identifiable to species using external or genitalic morphology, or by mitochondrial DNA sequences (Linnen and Farrel 2012). Despite extensive surveys in 2010–2013, N. sertifer populations have to date only been found in Washington State within the Bellingham city limits (Fig. 3). The records from Washington State could represent a newer and separate introduction event in the Pacific Northwest, although transport on nursery stock seems to be the most likely introduction pathway. Specimens are housed at WSDAC.

Diprion similis (Hartig, 1836)

Taeger et al. (2010) list 13 world species of Diprion. Most are Asian, but D. similis is one of two Diprion species native to Europe. Large populations of this solitary feeder occasionally occur in production forestry, often in association with D. pini L., but it is typically not a serious pest (Taeger et al. 1998). The only Diprion recorded from North America (Taeger et al. 2010), D. similis was discovered in New Haven, Connecticut, in 1914, presumably introduced on imported European nursery stock or associated packing materials (Britton 1915). Diprion similis feeds upon multiple pine species, with marked oviposition preference for P. strobus observed in North America (Tsao and Hodson 1956). The species can potentially defoliate entire trees when populations are high; however, this appears to be rare in North America, perhaps due to control by weather events and introduced parasitoids (Wilson 1966, Van Driesche et al. 1996). It is known to occur from the northeastern states westward to the Great Lakes region, and south to North Carolina.

Specimens of D. similis were collected in 2012 when adult females emerged from P. sylvestris boughs collected in Shelton, Washington. Following this detection, yellow card traps were deployed in the south Puget Sound area and Whatcom County. Subsequent visual surveys for larvae were conducted in western Washington. The distinctive larvae are readily recognized, and were found at eight sites in three western Washington counties (Fig. 3). Larvae were most commonly found feeding on P. monticola Douglas ex D. Don, but also on P. sylvestris and P. contorta Douglas ex Loudon. A single male specimen was captured in a Dipion pini pheromone-baited trap, probably by chance since the lure is not known to be attractive across species (O. Anderbrant, in litt.). Voucher specimens are deposited at WSDAC and WWUC.

Gilpinia hercyniae (Hartig, 1837)

Gilpinia comprises 37 described species native to Europe and Asia. Gilpinia hercyniae is a solitary spruce feeder, first detected in Ottawa, Ontario, Canada in 1922 and in New Hampshire in 1929 (Baker 1972). It quickly became a forest pest in the eastern United States and Canada (Balch 1939, Reeks and Barter 1951). Ambitious biological control programs during the 1930s imported and released several parasitoid species throughout the region. Concurrent with this, a nuclear polyhedrosis virus was inadvertently released which resulted in consistent region-wide control (Balch and Bird 1944). The species has not been an active management concern in North America for many decades (Nielon and Morris 1964, Kelleher and Hulme 1984). There are no published records of its occurrence west of Manitoba.

A specimen of G. hercyniae was collected in Bellingham in 2008 (although not identified until 2011) from Picea abies (L.) Karst. in a residential neighborhood. Wide-ranging visual and sticky-trap surveys in northwestern Washington failed to detect it beyond the original site, where more specimens were collected in 2011 and 2012 (Fig. 3). Specimens are housed at WWUC and WSDAC.

Tenthredinidae, Nematinae

Cladius grandis (Serville, 1823)

The earliest North American collections for this Palaearctic species are from Albany, New York, in 1887 (Smith 1974a). The species was presumably introduced separately to the west coast, with records from British Columbia in 1914 (Blackmore 1917). The specimens from British Columbia were first described as a new species, Platycampus victoria MacGillivray, 1920, reared from Populus nigra L. (MacGillivray 1920). In most previous literature, the species is known as Trichiocampus viminalis (Fallén, 1808). The most common host plants are Populus spp., although Salix (Benson 1958, Raizenne 1957) and Alnus P. Mill. (Smith 1974a) are also recorded. It has been reported as a minor pest of Populus L. in eastern North America and British Columbia (Béique 1961, Downes 1925). In Quebec, the species is bivoltine, with adults active in late May and early June, and again in late July through September (Béique 1961). There appears to be only one generation per year in British Columbia (Downes 1925). A single specimen from Washington was collected in a Malaise trap on the Evergreen State College campus in 1997 (Fig. 4), and detected while examining unidentified material in the college’s natural history museum. The specimen is housed at WSDAC.

Cladius gregarius Dyar, 1895

This poplar-feeding species is native to North America, and is known from the northeastern United States and adjacent Canada, west to Michigan and Ontario (Smith 1974a, 1979a). Larvae feed on species of Populus in the spring, and adults can be found in late summer and fall. Two adult specimens were reared from larvae collected on Populus tremuloides Michx. in Okanogan County, Washington, 2010 (Figs 45). A visit to the same site in late summer 2013 failed to find more larvae, although similar feeding damage was seen on trees throughout the area. Specimens are housed at WSDAC.

Figure 4.

Localities of Nematinae (Tenthredinidae) newly detected in Washington State.

Figure 5.

Cladius gregarius larvae on poplar, Okanogan County WA.

Pristiphora geniculata (Hartig, 1840)

Pristiphora is a large, primarily Holarctic genus, although there are several described Neotropical and southern Asian species (Taeger et al. 2010). The genus is most diverse in Europe, with ca. 115 known species. Five European Pristiphora spp. introduced to North America are pestiferous, including P. geniculata. This species was first detected in the United States in Haines Falls, New York, and in Massachusetts in 1926, and now occurs throughout the northeastern states and provinces, and west to Minnesota and Ontario (Schaffner 1936, Smith 1979a). Pristiphora geniculata feeds on Sorbus aucuparia L., S. americana Marshall, S. decora (Sarg.) C.K. Schneid., and the hybrid cultivar Sorbaronia hybrida (Moench) C.K. Schneid. (Forbes and Daviault 1964). Kunneman and Albers (1991) list P. geniculata as a pest of Tilia L., but this seems to be a misreporting of the “elm sawfly”, Cimbex americana Leach, 1817 (Cimbicidae; Dirr 1983, Sinclair et al. 1987). Larvae are voracious feeders and can almost completely defoliate healthy trees; however, mortality of even repeatedly defoliated trees is infrequent (Forbes and Daviult 1964). Release and establishment of the ichneumonid Olesicampe geniculatae Quednau and Lim in the eastern states and provinces has resulted in diminished outbreaks of P. geniculata since the 1980s (Kelleher and Hulme 1984, Quednau 1990).

The species was first detected in Washington State in 2009, and is now common throughout the Puget Sound region (Fig. 4). Most specimens have been found via visual survey of defoliated trees, where larvae were readily apparent and subsequently reared. Several adults were reared from Crataegus douglasii Lindl. (a native hawthorn species) in 2011 and 2012, a new host record for the species (Fig. 6). The record in Washington probably represents a separate introduction event, either from its native range or a translocation from eastern North America. Specimens are housed at WSDAC.

Figure 6.

Pristiphora geniculata larvae on Crataegus douglasii, Whatcom County, WA.

Pristiphora rufipes Serville, 1823

Pristiphora rufipes is native to central Europe, and spread to the United Kingdom in the mid-20th century (Benson 1947). It was first recorded in North America from Ottawa, Canada, in 1963 (MacNay 1964). The species is a significant pest of Aquilegia L. (Alford 2012). The species was previously known from the eastern states and provinces to the midwest. Most North American literature refers to the species by the synonym Pristiphora aquilegiae Snellen van Vollenhoven, 1866. It has been present in Washington since at least 1996, when Seattle-area gardeners began complaining of a new and voracious pest on columbine (Seattle Times 1996). Populations in Washington are currently known from Snohomish, King, and Thurston Counties (Fig. 4). The species has at least three generations per year in western Washington, with larvae present as late as November (Table 1). Specimens are housed at WSDAC.

Craesus alniastri (Scharfenberg, 1805)

Craesus alniastri is an alder-feeding nematine sawfly native to Europe, where it has been well-studied for its distinctive larval feeding aggregations (e.g., Boevé 1991). Buckle (1930) collected a single specimen of C. alniastri on parsnip flowers near Mt. Royal (Montreal), Quebec, Canada, in 1926, and Kirby (1882) lists a single male specimen housed in the Natural History Museum, London, that was collected in Nova Scotia. Smith (1972) does not report any further specimens, and there appears to be no further literature discussing the species in North America. Prous et al. (2014) reassign species in this genus to Nematus.

Numerous specimens of C. alniastri were collected in western Washington while conducting surveys for alder-feeding sawflies throughout the Pacific Northwest in 2009–2011 (Fig. 4). Adult specimens were collected from May through August. Specimens were collected very near the British Columbia and Oregon borders, indicating that the species is likely widespread in the region. An additional female specimen collected near Seattle in 1976 is in the USNM; other voucher specimens are deposited at WSDAC.

Nematus lipovskyi Smith, 1974

This species was described from the eastern United States (Smith 1974b) and is known from Maine to Alabama, and west to Wisconsin. It is recently established in the Czech Republic (Macek and Šípek 2014). Larvae feed on deciduous azaleas in the Pentanthera subgenus and section of Rhododendron (e.g., R. viscosum (L.) Torr., R. molle (Blume) G. Don, R. luteum Sweet, and R. calendulaceum (Michx.) Torr.). Macek and Šípek (2014) also noted feeding damage on R. obtusum (Lindl.) Planch. inflorescences, but proposed that this was opportunistic feeding due to the proximity of more suitable host plants. This species is reassigned to Euura in Prous et al. (2014).

A 1996 specimen from Lacey, Washington, is the earliest record from the western USA. Larvae have been observed in several locations in King, Thurston, and Clark counties (Fig. 4). Adults were collected in Olympia, Washington, in April 2015, and are stored at WSDAC.

Tenthredinidae, Heterarthrinae

Heterarthrus nemoratus (Fallén, 1808)

Heterarthrus is a relatively small genus of leaf-mining sawflies native to Europe and Asia, which generally mine leaves of trees in Aceraceae, Betulaceae, and Salicaceae (Taeger et al. 2010). Two species have been introduced to North America, H. nemoratus and H. vagans (Fallén, 1808), which mine species of Betula L. and Alnus, respectively (Smith 1971, Taeger et al. 1998, Humble 2010). Heterarthrus nemoratus was first collected in North America in Pictou, Nova Scotia, in 1905 (Dowden 1941), and now has a transcontinental distribution in Canada (Digweed et al. 2009). Digweed et al. (2009) review the biology and provide detailed keys to mines, larvae, and adults of this and other birch-feeding Heterarthrinae in North America. The earliest known Washington specimen is a female collected in Whatcom County (the northern-most county in western Washington) in 1967, housed at WWUC. Specimens are also known from the Seattle area (Fig. 7), and the species is likely widespread in western Washington.

Figure 7.

Localities of Heterarthrinae (Tenthredinidae) newly detected in Washington State.

Heterarthrus vagans (Fallén, 1808)

Heterarthrus vagans was recently detected in North America, discovered in British Columbia in 2009 in numerous locations west of the Cascade Range and very close to the Washington State border (Humble 2010). Visual and sticky trap surveys in 2011 and 2012 detected the species at six sites in Whatcom County (Fig. 7). We have not yet found the species south of Whatcom County, but the widespread distribution of alder makes it likely that this species will continue to spread through Washington and into Oregon. This species’ mines are identifiable by the round cocoon formed in the leaves of its host plants; other Heterarthrus spp. pupate outside of leaves (Humble 2010). Specimens are housed at WSDAC.

Metallus lanceolatus (Thomson, 1870)

Metallus lanceolatus (referenced as M. gei (Brishcke, 1883) in much of the European literature) is a leaf-miner of Geum L. The earliest North American specimens of this Palaearctic species were collected in British Columbia in 1933, and multiple specimens were collected in the northeastern USA and Canada in the 1960s (Smith 1971). Smith (1971) described M. bensoni as a new species from the New York and British Columbia specimens; Koch (1989) subsequently recognized M. bensoni and M. gei as junior synonyms of the Palaearctic M. lanceolatus. Metallus lanceolatus forms coalescing blotch-mines in ornamental and wild Geum, and can be a pest of garden plants (Buhr 1941, Hoebeke and Wheeler 2005). It has apparently spread through southern Puget Sound (Fig. 7), where it was found attacking Geum macrophyllum Willd. in cultivated and wild conditions. Specimens have been hand-collected and captured in Malaise traps. The earliest mines in a large patch of G. macrophyllum near Olympia, Washington, were visible by early July in 2012 and 2013, and could be found on nearly every plant by mid-July of both years. Specimens are housed at WSDAC.

Fenusella nana (Klug, 1816)

This Palaearctic species is commonly recorded in the literature as Messa nana (see Taeger and Blank 1998 for nomenclatural discussion). Fenusella nana larvae form coalescing blotch mines in many birch species (e.g., Buhr 1941, Taeger et al. 1998, Digweed et al. 2009). The first North American records for this leafminer are from Maine in 1966 (Smith 1967). It has since spread across Canada and into Washington State (Digweed et al. 2009). Digweed et al. (2009) review its biology and provide detailed keys to mines, larvae, and adults of this and other birch-feeding Heterarthrinae in North America. This species has been collected as far southwest as Hoquiam, Grays Harbor County (Fig. 7). Voucher specimens are housed at the NRFC.

Profenusa thomsoni (Konow, 1886)

The first North American records for this birch leafminer are from Hamden, Connecticut in 1926. It was known from Maine, Ontario, Quebec, and Vermont by the 1960s (Smith 1971), and has since spread across Canada and into Alaska, the Yukon Territory, and Northwest Territories (Digweed and Langor 2004, Digweed et al. 2009, MacQuarrie et al. 2013). It is a significant pest in urban forests (Martin 1960, Drouin and Wong 1984, Snyder et al. 2007). Digweed et al. (2009) review its biology and provide detailed keys to mines, larvae, and adults of this and other birch-feeding Heterarthrinae in North America. The earliest Washington specimen was collected in 1967, in Whatcom County, and was discovered by examining specimens housed at WWUC. Numerous specimens have also been collected south to King County (Fig. 7) and are housed at NRFC.

Profenusa inspirata (MacGillivray, 1909)

Profenusa inspirata, a native North American species, is the only known sawfly leaf-miner of oaks in western North America. It creates blotch mines in the upper surface of oak leaves (Fig. 8), that can coalesce when multiple larvae are present in a leaf. The species was previously known from Nevada, California, and Oregon (Smith 1971). Multiple mines were observed in Garry oak (Quercus garryana Douglas ex Hook.) along the Tieton River, Yakima County, and in Skamania and Lewis counties in 2015 (Fig. 7). Larvae were identified using characters described in Smith (1971). Voucher specimens are at WSDAC.

Figure 8.

Profenusa inspirata mines on Quercus garryana, Yakima County, Washington.

Fenusa ulmi Sundevall, 1847

This Palaearctic elm leafminer was already well established in New York by 1898 (Felt 1898), and was certainly introduced with elms from Europe. It is known to attack several elm species, especially Ulmus glabra Huds. and its hybrids (Slingerland 1905, Liston 1993, Scannell 2000); Scannell (2000) added U. pumilla L. and U. davidiana Planch. to the known elm hosts. Early records of significant damage to U. americana L. seem to be unsubstantiated, with most authors reporting that larvae are unable to do more than initiate feeding (Slingerland 1905, Guries and Smalley 1994, Scannell 2000). Scannell (2000) provided detailed information on the life history and host preferences of this species in the Seattle area. The oldest known west coast specimens were collected in British Columbia in 1947, and are stored in the Canadian National Collection (H. Goulet, pers. comm.). There is anecdotal evidence that the species has been present in Washington since the early 1990s (Scannell 2000), and has since spread south through the Puget Sound area (Fig. 7). Washington State voucher specimens are at WSDAC.

Tenthredinidae, Blennocampinae

Halidamia affinis (Fallén, 1807)

This is an introduced European species that feeds on Galium L (Smith 1969, Taeger et al. 1998). The earliest North America records are 1931 from Cold Spring Harbor, New York, 1933 from Connecticut, and 1934 from New Jersey (Smith 1969). It has since spread to California and the Pacific Northwest. The species is common and was collected as larvae from host plants and readily on yellow sticky traps and in Malaise traps in nearly every county surveyed (Fig. 9). Voucher specimens are at WSDAC.

Figure 9.

Localities of Blennocampinae, Allantinae, and Tenthredininae (Tenthredinidae) newly detected in Washington State.

Monophadnus pallescens (Gmelin, 1790)

A Ranunculus-feeding European introduction, this species has been present in the eastern United States and Canada since the late 1800s and in British Columbia since at least 1919 (Smith 1969). This species is widespread (Fig. 9), and was collected with sticky card traps, Malaise traps, and by hand. The earliest specimens discovered from Washington were collected in Whatcom County in the late 1960s, and were found in the material at WWUC. Numerous species of Ranunculus L. provide suitable hosts in western Washington, including the widespread invasive European weed Ranunculus repens L. Voucher specimens are housed at WWUC and WSDAC.

Eupareophora parca (Cresson, 1880)

The genus Eupareophora contains three species, with only E. parca native to North America. It is known from most of the northeastern and central states, and north and west to Alberta (Williams 2007). The species is not known from the Rocky Mountains, but has been collected from several northern California locations and Oregon (Smith 1969). The single historical specimen from Oregon was collected by Koebele, who collected in and around California in the 1880s. Most collection records for this western disjunct population are from California (Smith 1969), suggesting the species may have been native to eastern North America and spread west via human commerce. Williams (2007) similarly suggests that the species was not present in Alberta until the early 2000s. Several Fraxinus L. species are recorded hosts, as well as Carya illinoinensis (Wangenh.) K. Koch (Smith 1969) and Chionanthus L. (Dyar 1898), although Smith (1969) described the latter association as “dubious”. Larvae are readily recognizable by the bristly appearance caused by numerous thick, bifurcate spines. Williams (2007) presented a very detailed account of larval and adult biology in Alberta. Heavy ash defoliation and shed skins of an unknown sawfly were noticed at a public park on the border of King and Snohomish counties in 2014, but no living animals were observed. A return visit in May 2015 found the easily identifiable larvae feeding on Fraxinus latifolia Benth. throughout the park (Fig. 9). Voucher specimens are at WSDAC.

Tenthredinidae, Allantinae

Monostegia abdominalis (Fabricius, 1798)

Monostegia abdominalis is a European sawfly pestiferous on Glaux L., Lysimachia L. and Anagallis L. (Price 1970, Taeger et al. 1998). It was first collected in North America in Massachusetts in 1899, and described as M. martini MacGillivray, 1908 (Smith 1979b). In late summer of 2013, noxious weed management staff in King County noticed heavy defoliation of populations of Lysimachia vulgaris L., regulated as a noxious weed in Washington State, by unknown sawfly larvae. Larvae were subsequently collected and reared in the lab on a Lysimachia cultivar in 2014. Specimens are known from several Lysimachia infestations in western Washington (Fig. 9). Voucher specimens are at WSDAC.

Tenthredinidae, Tenthredininae

Macrophya punctumalbum (Linnaeus, 1767)

This adventive European species was first recorded in North America from Toronto, Ontario, in 1932, from British Columbia in 1934 (Gibson 1980), and later from New York (Hoebeke and Johnson 1985). Larvae feed on Ligustrum L., Syringa L., and Fraxinus (Gibson 1980). A specimen collected in 1977 in Bellingham, Washington, and housed at WWUC is the only record from Washington State (Fig. 9).

Discussion

In addition to expanding regional knowledge of an ecologically interesting and economically important group of insects, these data highlight the continual intra-continental spread of introduced species. Eighteen of the 22 sawflies reported here represent range expansions for exotic sawflies introduced to North America long ago. Most of these species were first recorded from eastern states or provinces, likely introduced with nursery stock. Five species may have been first or simultaneously introduced on the west coast based on their historical detection data - Cladius grandis, Heterarthrus vagans, Metallus lanceolatus, Monophadnus pallescens, and Macrophya punctumalbum. It is unknown how most of the introduced species became established in the west. Certainly, commerce from eastern North America could explain the distribution of some species. For some pests of common cultivated plants, such as Pristiphora rufipes and Neodiprion sertifer, it seems probable that insects were transported inadvertently with nursery trade or by home gardeners, although natural dispersal by adults can not be discounted.

The movement of other species, such as Halidamia affinis and Monostegia abdominalis, is more mysterious – Galium is not commonly cultivated, and Lysimachia vulgaris is a noxious weed. Halidamia affinis has likely spread of its own accord, expanding through the immense range of its host plant, Galium. Monostegia may have moved with other cultivars of Lysimachia that are commercially sold, although one would expect that such voracious and notable sawfly larvae on garden plants would have been reported. The reports of previously more southern species, such as Profenusa inspirata and Eupareophora parca could indicate northward range expansion concurrent with increasingly moderate winters. Profenusa inspirata in particular seems suggestive of such new expansion, since Garry oak conservation and ecology have long been studied in the Puget Sound region. However, it is certainly possible that P. inspirata has been present but undetected in Washington for decades.

The older specimens recorded here from collections made decades ago emphasize the value of institutional insect collections as repositories of valuable biodiversity information. It is noteworthy that for several of the species discussed herein, the first records for the state were found in the holdings of the insect collections at regional universities, providing evidence that many species had been transported west earlier than was previously known, or were perhaps derived from separate introduction events. Thus, despite the relatively small size of such collections compared to those at land grant universities, these collections fill a valuable role in documenting shifts in regional species composition. As a whole, the data from all museum specimens examined during this research also demonstrate gaps in our regional knowledge of sawflies. Unsurprisingly given remoteness and lack of access, the arid Columbia Basin and rugged mountain ranges in Washington are less frequently collected than other parts of the state (Fig. 2). Undoubtedly, other sawfly species occur in Washington that as yet are undetected and perhaps limited to these under-surveyed habitats. For example, the juniper-feeding genus Susana Rohwer & Middleton, 1932 occurs in states and provinces surrounding Washington but has not been recorded from the state.

Acknowledgements

We thank Kaile Adney, Jennifer Andreas, Rachel Chai, Caitlin Fate, Maggie M. Freeman, Thor Hanson, Warren E. Hellman, Amelia Jordan, Don Kitchen, Glenn Kohler, Andi Kopit, Eric LaGasa, Diane Maclean, Joy Markgraf, Todd Murray, Arlo Pelegrin, Karen Ripley, Alexis Sarah, Kathy Sheehan, Holli Watne, and Sue Welch for field and/or lab assistance. Tim Cahill and multiple anonymous property owners generously allowed access to collecting sites. Frank Merickel (University of Idaho), Chris Marshall (Oregon State University), Jack Longino and Brendan Boudinot (The Evergreen State College), and Richard S. Zack (Washington State University) graciously facilitated research at their respective institutional entomology collections. We thank two anonymous reviewers for comments and observations that significantly improved the readability and professionalism of this article. The data in this report were collected in part with funding from the USFS Forest Health Monitoring Program and USDA-APHIS Cooperative Agreements #11-8550-1505-CA and #11-8550-1502-CA.

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