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Checklist of the Psylloidea (Hemiptera) of the U. S. Pacific
Northwest
Author(s): Carmen I. Castillo Carrillo, Andrew S. Jensen and William E. Snyder
Source: Proceedings of the Entomological Society of Washington,
118(4):498-509.
Published By: Entomological Society of Washington
DOI: http://dx.doi.org/10.4289/0013-8797.118.4.498
URL: http://www.bioone.org/doi/full/10.4289/0013-8797.118.4.498
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INIAP - Estación Experimental Santa Catalina
PROC. ENTOMOL. SOC. WASH.
118(4), 2016, pp. 498–509
CHECKLIST OF THE PSYLLOIDEA (HEMIPTERA) OF THE U. S.
PACIFIC NORTHWEST
CARMEN I. CASTILLO CARRILLO, ANDREW S. JENSEN, AND WILLIAM E. SNYDER
(CICC, WES) Department of Entomology, 100 Dairy Rd. FSHN building,
Washington State University, Pullman, Washington 99164-1120 U.S.A. (email:
wesnyder@wsu.edu); (ASJ) Northwest Potato Research Consortium, 95873
Goldmohr Ln, Lakeview, OR 97630 U.S.A.
Abstract.—Several members of the Psylloidea vector pathogens of agricultural
crops in the Pacific Northwest (“PNW”) U.S. states of Idaho, Montana, Oregon and
Washington. These crop-production concerns have renewed interest in the bio-
diversity of this superfamily in the region. We compiled a checklist of this group by
examining published records and collections. This revealed 118 species of psyllids
from 23 genera; 29 of these species had not been previously reported in the PNW.
Our species list provides a useful starting point for entomologists investigating the
ecology of emerging insect-transmitted plant pathogens in the region.
Key Words: psyllid, species, biodiversity, Idaho, Montana, Oregon, Washington
DOI: 10.4289/0013-8797.118.4.498
The Pacific Northwest (“PNW”) U.S.
states of Idaho, Montana, Oregon and
Washington span diverse eco-geographic
zones, ranging from relatively cool
and wet conditions east of the Cascade
and within the Rocky Mountains, to
relatively hot and dry conditions at
lower elevations in the rain shadows of
these mountain ranges (Raymond et al.
2014). In the last century the region has
been heavily altered by agriculture
(Robbins and Wolf 1994, Butler et al.
2004). The PNW leads the United States
in production of apples, potatoes, and
wheat, while contributing significantly to
national production of other important
commodities (USDA 2014, 2015, WSDA
2015, OSDA 2015). Much of this agri-
cultural production is centered in the
irrigated, arid Columbia Basin of east-
ern Washington and Oregon, and in
valleys along the Snake River in southern
Idaho (USDA 2014, 2015, WSDA 2015,
OSDA 2015).
Pear (Pyrus communis L.) and potato
(Solanum tuberosum L.) crops in the
PNW have been harmed by plant path-
ogens vectored by the pear psylla,
Cacopsylla pyricola (Foerster), and the
potato psyllid, Bactericera cockerelli
(
Sulc), respectively. The pear psylla
harms pears by vectoring the phyto-
plasma Candidatus Phytoplasma pyri
(Jensen et al. 1964, Seemu¨ller and
Schneider 2004), which causes ‘pear
decline’ (Batiste and Bulla 1980).
Likewise, the potato psyllid vectors
the bacterium Candidatus Liberibacter
solanacearum, responsible for “zebra
chip” disease of potatoes that degrades
tuber quality in storage (Munyaneza et al.
2007; Buchman et al. 2011, Munyaneza
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2012). While B. cockerelli is believed to
be the only insect vector of the zebra
chip bacterium, limited familiarity with
psyllid biodiversity has restricted rigorous
verification of this assumption (Crosslin
et al. 2010, Haapalainen 2014). The
zebra chip bacterium has been found
in other Bactericera species, including
B. tremblayi (Wagner) and B. nigricornis
(Foerster) in Spain (Teresani et al. 2015),
and in Trioza apicalis Foerster in Finland
(Munyaneza et al. 2010). Uncertainty
about the true community of psyllid spe-
cies responsible for zebra chip transmission
has heightened interest in PNW psyllid
biodiversity, both within and outside of
agricultural fields (e.g., Munyaneza 2010,
Munyaneza andHenne 2012,Murphy et al.
2014).
Here, we compile a species list for
Psylloidea in the U.S. Pacific Northwest.
We focus on this region because of its
shared ecography and importance in
U.S. production of a similar suite of
agricultural commodities (USDA 2014,
2015, WSDA 2015, OSDA 2015). We
build upon the valuable earlier contri-
butions by Hodkinson (1988), who
provided a checklist of Nearctic Psy-
lloidea, by Percy et al. (2012) who
provided a checklist of psyllids in Cal-
ifornia (just south of our focal area), by
Ouvrard (2015) who built an extensive
online database of Psylloidea (“Psyl’list”;
http://www.hemiptera-databases.org/
psyllist/), and other published re-
ports as well as examinations of museum
specimens.
MATERIALS AND METHODS
We compiled our species list through
two main steps. First, we examined
existing published species lists for the
region (i.e., Hodkinson 1988, Percy et al.
2012, Ouvrard 2015), to construct an
initial list of known species in the region.
Next, we examined Psylloidea in col-
lections of the following key regional
institutions: the W.F. Barr Museum of
University of Idaho (UIDA), the Oregon
State Arthropod Collection of Oregon
State University (OSAC), the Oregon State
Department of Agriculture (OSDA),
the M.T. James Museum of Washington
State University (WSU), and the USDA
Yakima Agricultural Research Labo-
ratory in Washington (YARL). We did
not evaluate for accuracy the identifi-
cations in the museum collections, an
undertaking that was outside the scope
of our project. We did, however, record
the person who determined each spec-
imen when a determination label was
present; these were as follows: UIDA:
P.W. Oman, L.M. Russel, D.D. Jensen,
H.C. Manis, L. Bauer; WSU: L.M.
Russel, Dodge; OSU: L.M. Russel, S.
Rondon, P.W. Oman, Journet, L.G.
Gentner; OSDA: E.R. Hoebeke, J.J.
Vlach, A.L. Westcott, J.D. Joung, V.
Roth, K. Goeden. Additional species
names were extracted from the collec-
tion list of, and two reports from,
the Washington State Department of
Agriculture (“WSDA”; Johansen and
Brannon 1955, Beers et al. 1993) and
from the online psyllid list posted
by the Smithsonian National Museum
of Natural History (USNM) (http://
collections.nmnh.si.edu/search/ento/).
Updates of the scientific names were
obtained fromOuvrard (2015), a regularly-
updated world Psylloidea database.
Most psyllid species have relatively
narrow host-plant ranges (Hodkinson
and White 1979), and while this in-
formation is not presented here it is
available from Ouvrard (2015).
RESULTS
Our compiled list contains 118 spe-
cies of psyllids, belonging to 5 families
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and 23 genera (Table 1). In total, our list
presents 29 new species-records across
our four focal U.S. States (13/46 species
are new records for Idaho; 1/28 species are
new records forMontana; 30/76 species are
new records for Oregon; and 13/58 species
are new records for Washington).
Checklist.—Species of this list are
classified according to Burckhardt and
Ouvrard (2012). In the list we present the
States in which the psyllids were col-
lected (with underline type indicating
a new record of that species in that State).
Family Aphalaridae
Subfamily Aphalarinae
Genus Aphalara Foerster, 1848
calthae/polygoni complex. Idaho (UIDA),
Oregon (OSDA). The specific identity of
psyllids in this group cannot reliably be de-
termined, and A. calthae is thought not to
exist in North America; consequently it is
best to refer to these specimens as A. calthae/
polygoni complex (D. Burckhardt in litt.).
curta Caldwell, 1937. Idaho (UIDA),
Washington (USNM).
loca Caldwell, 1937. Idaho (UIDA),
Oregon (Psyl’list), Washington (Psyl’list,
USNM).
nubifera Patch, 1912. Washington (Psyl’list).
rumicis Mally, 1894. Idaho (Psyl’list, UIDA),
Oregon (OSDA, Psyl’list), Washington
(Psyl’list).
At OSDA, this species has a question
mark about the identification.
simila Caldwell, 1937. Idaho (Psyl’list),
Oregon (Psyl’list, USNM), Washington
(Psyl’list).
Genus Craspedolepta Enderlein, 1921
americana Klimaszewski, 1979. Washington
(Psyl’list).
angustipennis (Crawford, 1911). Idaho
(OSAC, Psyl’list), Montana (OSAC,
Psyl’list), Oregon (OSAC, Psyl’list),
Washington (OSAC, Psyl’list).
anomola (Crawford, 1914). Oregon (OSAC,
Psyl’list), Washington (OSAC, Psyl’list).
canadensis Journet and Vickery, 1979.
Oregon (OSAC).
constricta (Caldwell, 1936). Idaho (Psyl’list),
Oregon (OSAC).
flavida (Caldwell, 1938). Oregon (OSAC,
Psyl’list).
fumida (Caldwell, 1938). Montana (Psyl’list).
furcata (Caldwell, 1936). Montana (OSAC,
Psyl’list), Oregon (OSAC, Psyl’list).
maculimagna Journet and Vickery, 1979.
Oregon (OSAC, Psyl’list).
magna Journet and Vickery, 1979. Idaho
(Psyl’list), Montana (Psyl’list).
minutissima (Crawford, 1911). Idaho (OSAC,
Psyl’list, UIDA), Oregon (OSAC, OSDA,
Psyl’list).
Table 1. Families and genera found in the PNW
region. Classification based on Burckhardt and Ouvrard
(2012).
Family Subfamily Genus Species
Aphalaridae Aphalarinae Aphalara 6
Craspedolepta 22
Pachypsyllinae Pachypsylla 3
Spondyliaspidinae Cryptoneossa 1
Ctenarytaina 1
Glycaspis 1
Calophyidae Calophyinae Calophya 6
Liviidae Euphyllurinae Psyllopsis 1
Neophyllura 3
Liviinae Livia 2
Psyllidae Psyllinae Arytaina 1
Arytainilla 1
Cacopsylla 30
Ceanothia 1
Euglyptoneura 2
Nyctiphalerus 6
Psylla 6
Purshivora 1
Triozidae Metatriozidinae Bactericera 10
Baeoalitriozus 1
Heterotrioza 1
Phylloplecta 3
Trioza 9
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nebulosa (Zetterstedt, 1828). Washington
(Psyl’list).
oregonensis Journet and Vickery, 1979. Oregon
(OSAC, Psyl’list).
parvula Journet and Vickery, 1979. Oregon
(OSAC, Psyl’list).
pinicola (Crawford, 1914). Idaho (OSAC,
Psyl’list, UIDA), Oregon (OSAC, Psyl’list).
pulchella (Crawford, 1911). Oregon (OSDA).
russellaeKlimaszewski, 1977. Idaho (Psyl’list),
Oregon (OSDA, Psyl’list), Washington
(Psyl’list).
schwarzi (Ashmead, 1904). Washington
(Psyl’list).
smithsonia (Klimaszewski, 1979). Montana
(Psyl’list).
vancouverensis (Klyver, 1931). Idaho (Psyl’list),
Montana (Psyl’list), Oregon (OSAC, Psyl’list),
Washington (Psyl’list, USNM).
veaziei (Patch, 1911). Oregon (OSAC, Psyl’list),
Washington (Psyl’list).
vulgaris (Journet and Vickery, 1979). Montana
(OSDA, Psyl’list).
Subfamily Pachypsyllinae
Genus Pachypsylla Riley, 1885
celtidisgemma Riley, 1885. Idaho (UIDA).
celtidismamma (Fletcher, 1883). Idaho
(Psyl’list, UIDA), Oregon (OSDA),
Washington (WSU).
venusta (Osten-Sacken, 1861). Idaho
(Psyl’list, UIDA), Washington (Psyl’list,
UIDA, WSU).
Subfamily Spondyliaspidinae
Genus Cryptoneossa Taylor, 1990
triangula Taylor, 1990. Oregon (OSDA).
Genus Ctenarytaina Ferris and
Klyver, 1932
eucalypti (Maskell, 1890). Oregon (OSDA).
Genus Glycaspis Taylor, 1960
brimblecombei Moore, 1964. Oregon
(OSDA).
Family Calophyidae
Subfamily Calophyinae
Genus Calophya Lo¨w, 1879
aurea Tuthill, 1942. Montana (Psyl’list).
dubia Crawford, 1914. Montana (Psyl’list).
flavida Schwarz, 1904. Oregon (OSAC).
nigripennis Riley, 1885. Washington
(Psyl’list).
triozomima Schwarz, 1904. Idaho (Psyl’list).
washingtonia (Klyver, 1931). Washington
(Psyl’list).
Family Liviidae
Subfamily Euphyllurinae
Genus Psyllopsis Lo¨w, 1879
fraxinicola (Foerster, 1848). Idaho (Psyl’list),
Oregon (OSDA).
Genus Neophyllura Loginova, 1973
arbuti (Schwarz, 1904). Oregon (Psyl’list).
arctostaphyli (Schwarz, 1904). Montana
(Psyl’list), Oregon (OSAC, OSDA, Psyl’list),
Washington (Psyl’list).
separata (Tuthill, 1943). Oregon (Psyl’list,
USNM).
Subfamily Liviinae
Genus Livia Latreille, 1802
caricis Crawford, 1914. Idaho (Psyl’list),
Oregon (Psyl’list).
vernaliforma Caldwell, 1940. Montana
(Psyl’list).
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Family Psyllidae
Subfamily Psyllinae
Genus Arytaina Foerster, 1848
genistae (Latreille, 1804). Oregon (OSDA,
Psyl’list), Washington (Psyl’list, YARL).
Genus Arytainilla Loginova, 1972
spartiophila (Foerster, 1848). Oregon (OSDA,
Psyl’list), Washington (Psyl’list, YARL).
Genus Cacopsylla Ossiannilsson, 1970
acuminata (Jensen, 1956). Oregon (YARL).
alba (Crawford, 1914). Idaho (Psyl’list),
Washington (Psyl’list).
americana (Crawford, 1914). Idaho
(Psyl’list), Oregon (OSDA, Psyl’list),
Washington (Psyl’list, WSDA, YARL).
breviata (Patch, 1912). Washington (UIDA).
brevistigmata (Patch, 1912). Idaho (UIDA),
Oregon (YARL).
YARL’s note “Questions remain about
this identification.”
confusa (Tuthill, 1943). Oregon (OSAC,
UIDA).
coryli (Patch, 1912). Idaho (Psyl’list, UIDA),
Oregon (Psyl’list, UIDA), Washington
(Psyl’list, YARL).
curta (Tuthill, 1943). Oregon (Psyl’list).
difficilis (Tuthill, 1943). Idaho (Psyl’list,
UIDA), Montana (Psyl’list, USNM).
fatsiae (Jensen, 1957). Washington (WSDA).
fibulata (Crawford, 1914). Idaho (Psyl’list),
Oregon (Psyl’list, UIDA).
hirsuta (Tuthill, 1938). Idaho (Psyl’list),
Montana (Psyl’list), Oregon (OSDA,
Psyl’list, UIDA, USNM, YARL), Washington
(Psyl’list, YARL).
insignita (Tuthill, 1943). Oregon (OSAC,
UIDA).
latiforceps (Tuthill, 1943). Oregon (USNM),
Washington (Psyl’list).
magna (Crawford, 1914). Montana (OSAC),
Oregon (OSAC, UIDA, YARL).
YARL’s note “Questions remain about
this identification.”
magnicauda (Crawford, 1914). Montana
(Psyl’list).
manisi (Tuthill, 1943). Idaho (Psyl’list,
UIDA, USNM).
media (Tuthill, 1943). Idaho (UIDA), Oregon
(OSAC, YARL).
YARL’s note “Questions remain about
this identification.”
minor (Crawford, 1914). Oregon (OSAC,
Psyl’list, USNM), Washington (Psyl’list,
WSU).
minuta (Crawford, 1914). Idaho (Psyl’list),
Oregon (UIDA), Washington (UIDA).
nordica (Jensen, 1951). Oregon (Psyl’list),
Washington (Psyl’list, UIDA, USNM).
omani (Tuthill, 1943). Oregon (UIDA,
YARL).
parallela (Crawford, 1914). Washington
(Psyl’list).
pararibesiae (Jensen, 1956). Washington
(Psyl’list, YARL).
peregrina (Foerster, 1848). Oregon (OSDA,
Psyl’list), Washington (Psyl’list, YARL).
pyricola (Foerster, 1848). Idaho (UIDA),
Washington (OSDA, Psyl’list, UIDA,
WSDA, WSU, YARL).
ribesiae (Crawford, 1911). Idaho (Psyl’list,
UIDA), Oregon (Psyl’list, YARL),Washington
(YARL).
sinuata (Crawford, 1914). Idaho (Psyl’list).
striata (Patch, 1911). Washington (Psyl’-
list).
usitata (Tuthill, 1943). Montana (Psyl’list,
USNM).
Genus Ceanothia Heslop–Harrison,
1961
ceanothi (Crawford, 1914). Montana
(Psyl’list), Oregon (OSDA), Washington
(Psyl’list).
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Genus Euglyptoneura
Heslop–Harrison, 1961
fuscipennis (Crawford, 1914). Idaho (UIDA,
WSU), Oregon (OSDA, Psyl’list).
robusta (Crawford, 1914). Idaho (Psyl’list,
USNM), Montana (Psyl’list, USNM), Ore-
gon (Psyl’list, UIDA), Washington (Psyl’list).
Genus Nyctiphalerus Bliven, 1955.
adustus (Tuthill, 1937). Oregon (YARL).
cercocarpi (Jensen, 1957). Oregon (YARL).
idahoensis Jensen, 1946. Idaho (Psyl’list,
UIDA, USNM).
rugipennis (Crawford, 1914). Oregon (Psyl’-
list, OSDA).
tantillus (Tuthill, 1937). Idaho (UIDA).
vermiculosus (Crawford, 1914). Idaho
(Psyl’list, UIDA), Montana (Psyl’list),
Oregon (Psyl’list).
Genus Psylla Geoffroy, 1762
alni (Linne, 1758). Idaho (Psyl’list), Oregon
(OSDA, Psyl’list), Washington (Psyl’list).
buxi (Linne, 1758). Oregon (OSAC, OSDA),
Washington (WSDA).
caudata Crawford, 1914. Idaho (Psyl’list,
UIDA).
floccosa Patch, 1909. Idaho (UIDA), Montana
(Psyl’list), Oregon (OSDA, Psyl’list,
UIDA), Washington (Psyl’list, WSU).
sanguinea (Provancher, 1872). Idaho (UIDA),
Washington (Psyl’list).
viridescens (Provancher, 1872). Idaho (UIDA,
WSU), Oregon (OSDA, Psyl’list), Wash-
ington (Psyl’list, WSU).
Genus Purshivora Heslop–Harrison,
1961
pubescens (Crawford, 1914). Idaho (Psyl’list),
Oregon (OSAC, Psyl’list, UIDA, USNM,
YARL), Washington (YARL).
Family Triozidae
Genus Bactericera Puton, 1876
arbolensis (Crawford, 1910). Montana
(Psyl’list).
cockerelli (
Sulc, 1909). Idaho (Psyl’list,
UIDA), Montana (Psyl’list), Oregon (OSAC,
Psyl’list, YARL), Washington (Psyl’list,
WSU).
incerta (Tuthill, 1943). Oregon (Psyl’list,
USNM), Washington (Psyl’list, USNM).
lobata (Crawford, 1914). Oregon (OSDA).
maculipennis (Crawford, 1910). Oregon
(OSDA), Washington (WSU, YARL).
minuta (Crawford, 1910). Idaho (Psyl’list,
UIDA), Montana (Psyl’list), Oregon (OSDA,
Psyl’list, YARL), Washington (Psyl’list,
WSU).
pletschi (Tuthill, 1944). Montana (Psyl’list,
USNM).
pulla (Tuthill, 1939). Oregon (Psyl’list,
USNM), Washington (USNM).
rubra (Tuthill, 1939). Oregon (Psyl’list).
salicivora (Reuter, 1876). Idaho (UIDA),
Oregon (Psyl’list), Washington (Psyl’list,
WSDA, WSU). In UIDA, WSDA and
WSU, this species is present as Bactericera
maura (Foerster, 1848); see Hodkinson,
1988.
Genus Baeoalitriozus Li, 2011
diospyri (Ashmead, 1881). Montana
(Psyl’list).
Genus Heterotrioza Dobreanu and
Manolache, 1960
chenopodii (Reuter, 1876). Oregon (YARL),
Washington (YARL).
Genus Phylloplecta Riley, 1884
occidentalis (Tuthill, 1939). Washington
(Psyl’list, USNM).
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rubicola (Tuthill, 1943). Oregon (OSDA),
Washington (Psyl’list, USNM).
tripunctata (Fitch, 1851). Washington
(Psyl’list).
Genus Trioza Foerster, 1848
alacris Flor, 1861. Oregon (OSDA), Washington
(WSDA).
albifrons Crawford, 1910. Idaho (Psyl’list),
Montana (Psyl’list), Oregon (Psyl’list),
Washington (Psyl’list, WSU, YARL).
eugeniae Froggatt, 1901. Oregon (OSDA).
inversa Tuthill, 1939. Oregon (OSDA).
mira Tuthill, 1943. Washington (Psyl’list,
WSU).
obtusa Patch, 1911. Washington (UIDA).
quadripunctata Crawford, 1910. Montana
(Psyl’list, USNM).
robusta Tuthill, 1944. Montana (Psyl’list).
sulcata Crawford, 1910. Oregon (Psyl’list).
DISCUSSION
The recent Psylloidea is composed of
eight families: Aphalaridae, Calophyidae,
Carsidaridae, Homotomidae, Liviidae,
Phacopteronidae, Psyllidae, and Tri-
ozidae (Burckhardt and Ouvrard 2012).
Previous work (Ouvrard 2015) and our
list show that five of these families
are represented in the Pacific Northwest:
Aphalaridae, Calophyidae, Liviidae, Psy-
llidae, and Triozidae. Our list updates the
psyllid species known in the PNW with
the addition of 29 new reports.
The most species-rich genera are Ca-
copsylla (30 species), Craspedolepta (22
species) and Bactericera (10 species).
Species of Cacopsylla and Bactericera
are known as vectors of economically-
important plant pathogens; the following
10 species were found in the PNW and
are listed as pests by Percy (2005): B.
cockerelli (pest of potato as mentioned
above), P. buxi (minor pest in ornamen-
tals), C. pyricola (common pear psyllid,
mentioned above), C. triangula, C. eu-
calypti and G. brimblecombei (pests in
Eucalyptus spp.; see also Brennan et al.
[1999]), L. alacris (pest in bay trees,
Laurus spp.), and reported as minor
pests are B. diospyri (on wild persim-
mon, Diospyros virginiana, in the U.S.)
and T. eugeniae (on myrtaceous trees
such as Syzygium spp. in Australia).
Also present in the PNW is the broom
psyllid, Arytainilla spartiophila, which
in the 1950s was released in the U.S. as
a biological control agent against the
invasive shrub Scotch broom (Cytisus
scoparius) (Syrett et al. 1999); we now
know that any harm to the shrub results
primarily from infection by Candida-
tus Liberibacter europaeus, a bacte-
rium vectored by this psyllid species
(Thompson et al. 2013).
The PNW has periodically seen
emergence of new psyllid-vectored
pathogens of agricultural importance
(Horton 1999, Munyaneza et al. 2007).
Eruption of these pathogens is often
followed by a frantic search for the
causative pathogen and then its insect
vector (e.g., Munyaneza et al. 2007,
Munyaneza et al. 2010). At times, rela-
tive ignorance of regional psyllid
biodiversity, population dynamics,
and phenology has slowed initial
development of effective pathogen-
vector suppression plans (Munyaneza
2010, 2012). Our checklist offers re-
searchers in agriculture and ecology
a picture of psyllid diversity in the PNW
that we hope will inform efforts to
identify possible psyllid vectors of plant
pathogens and put research on particu-
lar species into proper context of the
range of psyllid diversity in the region.
Our list may aid first steps to address the
development of integrated pest man-
agement strategies for any future
emerging pathogens known, or thought,
to be psyllid-vectored. Finally, we hope
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our work will support the education and
development of future Psylloidea spe-
cialists in North America.
We suggest that future work be con-
ducted to confirm museum specimen
identifications, and note the need for
thorough field surveys of psyllid pop-
ulations, diversity, and host plant bi-
ology across seasons and habitats, in the
exceptionally-diverse range of biogeo-
graphic zones in the Pacific Northwest.
ACKNOWLEDGMENTS
Checklist construction was supported
by grants from the Northwest Potato
Research Consortium and the Wash-
ington State Commission on Pesticide
Registration. The authors were supported
during manuscript preparation by the
National Institute of Food and Agricul-
ture, U.S. Department of Agriculture,
under Specialty Crop Research Initiative
award number 2015-09273. C. Castillo
Carrillo was supported by a graduate re-
search fellowship from Ecuador’s Na-
tional Institute of Agriculture Research
(INIAP) and the National Secretary of
Higher Education, Science, Technology
and Innovation (SENESCYT). Special
thanks to those who provided access to
the collections that they curate and sup-
port: James “Ding” Johnson and Frank
Merickel with the Barr Entomological
Museum at the University of Idaho;
Christopher Marshall with the Oregon
State Arthropod Collection at Oregon
State University; William Turner (Emer-
itus professor) and Richard Zack with the
M. T. James Collection at Washington
State University; Joshua Vlach at the
Oregon State Department of Agricul-
ture Plant Division; Debbie Creel at
the USDA-ARS Systematic Entomology
Laboratory; Eric LaGasa (retired scien-
tist), Chris Looney, and Angela Yoder
from theWashington State Department of
Agriculture; and David Horton and
Tamera Lewis from Yakima Agri-
cultural Research Laboratory, USDA
Fruit and Vegetable Insect Research
Unit in Washington. The manuscript
was greatly improved by the reviews of
Daniel Burckhardt, Natural History
Museum Basel, and one anonymous
reviewer, including valuable taxo-
nomic corrections from the former.
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