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Volume 99 ISSN #0071-0733
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COVER: An apterous vivipara of Myzus persicae (Sulzer) feeding on the leaf of Chinese
pe-tsai, Brassica pekinensis (Loureiro) Ruprecht, 70x magnification.

Scanning electron microscopy. Live green peach aphid feeding on Chinese pe-tsai leaf
was transferred to an aluminum tape-rimmed SEM stub. The loaded stub and its holder
were placed in the specimen chamber, viewed and pictured with a Hitachi S-500 SEM at an
accelerating voltage of 15 kv under vacuum.

Remark. Cho-kai Chan took this picture in 1979, and won the competition of insect photo
contest during an ESBC annual general meeting in the early 1980's. The judge was SFU’s
renowned photographer, Mr. Ron Long. This picture was then used as a logo on the

pamphlet for the Agriculture Canada Vancouver Research Station. Cover photo and text by
Cho-kai Chan.

The Journal of the Entomological Society of British Columbia is published annually in
December by the Society

Copyright® 2002 by the Entomological Society of British Columbia
Designed and typeset by Ward Strong and David Holden.
Printed by Reprographics, Simon Fraser University, Burnaby, BC, Canada.

Printed on Recycled Paper.

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J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 1

Journal
of the
Entomological Society
of British Columbia

Volume 99 Issued December 2002 ISSN #0071-0733
Directors of the Entomological Society of British Columbia 2000-2001.............ceceeeeeeeeees 2

Hamilton, K.G.A. Homoptera (Insecta) in Pacific Northwest grasslands. Part 1 — New and
revised taxa of leafhoppers and planthoppers (Cicadellidae and Delphacidae) ............ 3

Hamilton, K.G.A. Homoptera (Insecta) in Pacific Northwest grasslands. Part 2 — Pleistocene
refugia and postglacial dispersal of Cicadellidae, Delphacidae and Caliscelidae ....... 33

Raworth, D.A. and M.C. Robertson. Occurrence of the spider mite predator Stethorus
punctillum (Coleoptera: Coccinellidae) in the Pacific Northwest ................ (NOTE) 81

Furniss, M.M., E.H. Holsten, and M.E. Schultz. Species of Alaska Scolytidae: Distribution,
HO StS ed ING MA SCOT Cale VS Wire aces cle ch x enue Oo ern en gee ae Ee 83

McGregor, R.R., R.P. Prasad, and D.E. Henderson. Searching behaviour of Trichogramma
wasps (Hymenoptera: Trichogrammatidae) on tomato and pepper leaves.................. 93

Kenner, R.D. Stylops shannoni (Stylopidae, Strepsiptera): A New species for Canada, with
WMI 19057 OM VCO SED CCNIE sen at ce, ait tia 1itvanarenaes¥ostsniudeuakse tom teen neem eee ee eee 99

Miller, D.R. Short-range horizontal disruption by verbenone in attraction of mountain pine
beetle (Coleoptera: Scolytidae) to pheromone-baited funnel traps in stands of lodgepole
SEL Saree cree se ner eee eer ene ts Ned eae ce roe As oes th neat deren ea meena (NOTE) 103

Knight, A.L., D. Larson, and B. Christianson. Flight Tunnel and Field Evaluations of Sticky
Traps for Monitoring Codling Moth (Lepidoptera: Tortricidae) in Sex Pheromone-treated
(OE RIT C0 ISN esr Rec ga een len en cn eto ere eee 107

Morewood, W.D. and K.E. Simmonds. a@Pinene and ethanol: Key host volatiles for
Xylotrechus longitarsis (Coleoptera: Cerambycidae) ...............cccccccsccceessseceesssseeeeeens hy

Knight, A.L. A Comparison of Gray Halo-butyl Elastomer and Red Rubber Septa to Monitor
Codling Moth (Lepidoptera: Tortricidae) in Sex Pheromone-Treated Orchards ...... 123

Blades, D.C.A. A new species of Boreus (Mecoptera: Boreidae) from Vancouver Island,

| BVI SS) OA Efe) Wu oa op Cen aera ree rca pease peas get oe In a gee Ree on 133
Erratum: Insect population ecology in British Columbia ...............00.ccccccccscccessceeseeeeeees 14]
Erratum: Behavioral and chemical ecology in British Columbia .................:.:::ceeeeee 143

NOTICE RO GONURIBWTORS 27 ae eee eee we ee Inside Back Cover

2 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

DIRECTORS OF THE ENTOMOLOGICAL SOCIETY OF BRITISH
COLUMBIA FOR 2002-3

President:
Lorraine MacLauchlan
BC Ministry of Forests

President-Elect:
Gayle Anderson
Simon Fraser University

Past-President.
Rob Cannings
Royal BC Museum

Secretary/Treasurer:
Robb Bennett
BC Ministry of Forests, 7380 Puckle Rd. Saanichton BC V8M 1W4

Directors, first term:
Chris Guppy, Ian Wilson

Directors, second term:
Keith Deglow, Rene Alfaro, Tracy Huepplesheuser

Regional Director of National Society:
Terry Shore
Canadian Forest Service, Victoria

Editorial Committee, Journal:
Ward Strong (Editor), Peter Belton,
H.R. MacCarthy (Editor Emeritus), Ken Naumann,
Lorraine MacLauchlan, Dave Raworth

Editor of Web Site:
Bill Riel
Canadian Forest Service, Victoria

Editor, Boreus:
Cris Guppy
Quesnel, BC

Honourary Auditor:

Rob Cannings
University of Victoria

Web Page: http://esbc.harbour.com/

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Homoptera (Insecta) in Pacific Northwest grasslands.
Part 1 — New and revised taxa of leafhoppers
and planthoppers (Cicadellidae and Delphacidae).

K.G. ANDREW HAMILTON

BIODIVERSITY, RESEARCH BRANCH, A.A.F.C.
C.E.F. OTTAWA, ONTARIO, CANADA K1A 0C6

ABSTRACT

Nine new combinations are created when Acanthodelphax LeQuesne (Delphacidae) is
synonymized with Kosswigianella Wagner and the genus Aschedelphax Wilson is reduced
to subgenus in Elachodelphax Vilbaste: Delphacodes analis Crawford and three Palaearctic
species in Acanthodelphax are transferred to Kosswigianella; Aschedelphax hochae Wilson,
Delphacodes coloradensis Beamer, D. indistinctus Crawford, D. bifidus Beamer, and D.
pedaforma Beamer are transferred to Elachodelphax. Seven other new combinations include
Delphacodes atridorsum Beamer, transferred to Caenodelphax Fennah; Delphacodes
kilmani (Van Duzee), transferred to Paraliburnia Jensen-Haarup; Delphacodes venusta
Beamer, transferred to Nothodelphax Fennah; Eurysa magnifrons (Crawford), E. montana
Beamer and £. obesa Beamer, transferred to Eurybregma Scott; plus the leafhopper
Sorhoanus helvinus (Van Duzee) transferred to Lebradea Remane. Twenty new species and
3 new subspecies are described from the Pacific Northwest, including the leafhoppers
Athysanella castor, A. hyperoche, A. lemhi, A. occidentalis megacauda, A. repulsa,
Diplocolenus configuratus bicolor, Mocuellus caprillus anfractus, M. quinquespinus,
Psammotettix diademata, P. nesiotus, Sorhoanus involutus, S. virilis, S. xiphosura,
Stenometopiellus vader and Unoka dramatica, plus the planthoppers Achorotile apicata,
Elachodelphax unita and E. mazama (both compared to E. borealis sp. nov. from east of the
Rocky Mountains), Eurybregma eurytion, Kosswigianella irrutilo, K. wasatchi,
Paraliburnia furcata and P. lecartus. Two new names are created: Athysanella
(Gladionura) hicksi for A. expulsa Blocker and Hicks (preoccupied), and Psammotettix
greenei for P. emarginata Greene (preoccupied). The presence of intermediate morphs
suggestive of hybrids reduce Athysanella ladella Johnson and A. wilburi Ball & Beamer to
subspecific status in A. occidentalis Baker, Diplocolenus nigrior Ross & Hamilton to
subspecies status in D. configuratus (Uhler) and Mocuellus strictus Ross & Hamilton to
subspecies status in M. caprillus Ross & Hamilton. Variation within series over a wide area
reduces Athysanella vativala Blocker to synonymy with A. terebrans Gillette & Baker, and
A. mansa 1s synonymized with A. occidentalis wilburi as a hybrid morph.

INTRODUCTION

Grasslands form a minor part of the rugged landscape of the Pacific Northwest (PNW).
They are restricted to favourable elevations sandwiched between huge areas of coniferous
forest and sagebrush semidesert. The elevation of these grasslands varies with both latitude
and local topography. In the southern Okanagan Valley of British Columbia (latitude 49°) the
following communities are generally found on west-facing slopes at the following elevations:
sagebrush steppe to 350 m; steppe-grassland ecotone, to 700 m; Palouse grassland to 800 m;
grassland-forest ecotone to 1000 m. At Bannock Pass on the Idaho-Montana border (latitude
45°) sagebrush steppe reaches 2000 m and forests begin again just above the pass (2200 m),
compressing grassland and its ecotones into a few hundred metres. South-facing slopes can
raise the elevation of the grassland community as much as 200 m. Conversely, grasslands are

4 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

often absent on steep north-facing slopes.

Much of these grasslands have been converted to agriculture, or confined to small patches
in intermontane valleys, dissected by deep canyons, or separated by mountain ranges. Pacific
Northwest grasslands extend over six states and two provinces. Their main mass runs from
isolated valleys of central British Columbia (BC) south to the Palouse hills of Washington
(WA) and the Snake River plains of Idaho (ID), and down the foothills of the Cascade
Mountains to Oregon (OR). Patches of this grassland may be found on mountains of Colorado
(CO), Utah (UT) and Wyoming (WY). On the eastern slopes of the Rocky Mountains a similar
grassland extends from southwestern Alberta (AB) to southern Montana (MT).

PNW grasslands are home to many grassland-endemic phytophagous insects. The most
numerous species of these endemics belong to the leafhopper family Cicadellidae (Figs. 1-2).
Leafhoppers are diverse, with over 2000 species in North America (Oman 1949). A large
proportion of these are grassland insects (Ross 1970) which include more than 150 species
monophagous on grasses (Hamilton and Whitcomb 1993) and many others oligophagous on
arid-adapted forbs, principally composites such as sagebrush, Artemisia spp., balsamroot,
Balsamorhiza spp. and rabbitbrush, Chrysothamnus spp. (Hamilton 1998a, Hamilton and Zack
1999).

The leafhopper fauna we find today in the PNW has a high percentage of endemic species.
A smaller number of leafhopper species are widely dispersed across many habitat types. These
have little to tell us about historical biogeography. For example, species of Helochara Fitch
(Hamilton 1986, maps 17-18) which are not restricted to prairie plants range far outside native
grasslands.

The purpose of this paper is to present new and revised taxa of PNW endemic leafhoppers
and planthoppers (Fulgoroidea) of the family Delphacidae (Fig. 3). The second part analyses
their biogeography, together with that of planthoppers of the family Caliscelidae (Fig. 4), for
evidence of endemism and postglacial spread. These planthoppers, although fewer in number
and less studied than leafhoppers, show similar dispersal patterns and thus are included in this
analysis.

METHODS

Numerous leafhopper and planthopper species inhabit prairies and semidesert grasslands
adjacent to the PNW, but have no populations in the PNW. They are therefore excluded from
this study. These include the leafhopper genera Dicyphonia Ball, Driotura Osborn & Ball,
Extrusanus Oman, and various genera of the Sonoran subregion. Nevertheless, hundreds of
grassland-endemic leafhopper and planthopper species do inhabit PNW grasslands, permitting
detailed analysis of their known distributions.

The few PNW species of the leafhopper genera Acinopterus Van Duzee, Chlorotettix Van
Duzee, Commellus Osborn & Ball, Endria Oman, Frigartus Oman, Hardya Edwards,
Idiodonus Ball, Lonatura Osborn & Ball, Lystridea Baker, Mesamia Ball, Paluda DeLong,
Pinumius Ribaut, Stragania Stal, Twiningia Ball and Xerophloea Germar are distinctive
(Oman 1949; Beirne 1956) and present no taxonomic problems to the biogeographer.
Revisionary work has been done for the more complex leafhopper genera Amblysellus
Sleesman (Kramer 1971), Athysanella Baker (Blocker and Johnson 1988, 1990a, b, c; Blocker
and Hicks 1992), Attenuipyga Oman (Hamilton 2000), Auridius Oman (Hamilton 1999),
Balclutha Kirkaldy (Hamilton 1983), Carsonus Oman (1938), Ceratagallia Kirkaldy
(Hamilton 1998a), Colladonus Ball (Nielson 1957), Cuerna Melichar (Hamilton 1970),
Diplocolenus Ribaut (Ross and Hamilton 1970a), Draeculacephala Ball (Hamilton 1985),
Elymana DeLong (Chiykowski and Hamilton 1985), Errhomus Oman (Hamilton and Zack
1999), Euscelis Brullé and Evacanthus LePeletier & Serville (Hamilton 1983), Flexamia

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

DeLong (Whitcomb and Hicks 1988), Gyponana Ball (Hamilton 1982), Hebecephalus
DeLong (Hamilton 19985), Hecalus Stal (Hamilton 2000), Laevicephalus DeLong (Ross and
Hamilton 1972a), Latalus DeLong & Sleesman (Ross and Hamilton 19725), Limotettix
Sahlberg (Hamilton 1994), Mocuellus Ribaut (Ross and Hamilton 19705), Norvellina Ball
(Lindsay 1940), Orocastus Oman (Beirne 1956; Ross and Hamilton 19725), Paraphlepsius
Baker (Hamilton 1975), Prairiana Ball (DeLong 1942), Psammotettix Haupt (Greene 1971),
Rosenus Oman (Ross and Hamilton 1975), Scaphytopius Ball (Hepner 1947), Texananus Ball
(Crowder 1952); in Caliscelidae, Bruchomorpha Newman (Doering 1940); and among
Delphacidae, Eurysa Fieber (Beamer 1952), Laccocera Van Duzee (Scudder 1963), and
Pissonotus Van Duzee (Bartlett and Deitz 2000).

The first part of this study comments on the taxonomy of the leafhopper genera
Athysanella (with four new species and one new subspecies), Mocuellus and Sorhoanus Ribaut
(each with three new species), Psammotettix (two new species and one new name),
Stenometopiellus Haupt and Unoka Lawson (each with one new species), and Diplocolenus
(one new subspecies). One Delphacid is newly assigned to Nothodelphax Fennah, and eight
others to four genera that are recorded from the Nearctic for the first time: E/achodelphax
Vilbaste, Eurybregma Scott, Kosswigianella Wagner and Paraliburnia Jensen-Haarup. Eight
new Delphacid species are described in Elachodelphax subgenus Aschedelphax Wilson,
stat.nov. (three new species), Kosswigianella and Paraliburnia (each with two new species),
and Eurybregma (with one new species).

CICADELLIDAE

Cicadellid species are identified mainly by male genital structures (Figs. 5-37), although
proportions of the head and colour patterns are sometimes useful. Many grassland leafhoppers
feed on only one species (monophagous) or one genus of plant (oligophagous). This often
helps resolve species complexes where morphologically similar insects have different
biological requirements (see below). Leafhopper genera mentioned here are identifiable using
keys presented by Oman (1949). The following leafhopper taxa represent two redefined
genera, two new synonymies, two new names, 12 new species and three new subspecies.

Athysanella (Athysanella) terebrans (Gillette & Baker)

(Figs. 5 A-E)

Eutettix terebrans Gillette & Baker, 1895: 102.

Nephotettix terebrans: Van Duzee 1917: 653.

Athysanella terebrans: Osborn 1930: 703 (misidentification of A. fredonia Ball & Beamer).

Athysanella incongrua: Osborn 1930: 703 (nec Baker, 1898).

Athysanella vativala Blocker [in Blocker and Johnson], 1988: 38 (new synonymy).

Diagnosis. This species has been much confused with other taxa. Because the holotype of

terebrans is a female without associated male, its identity can be deduced only from colour,

female genitalia and locality (North Park, Colorado). Lengthy series from northern prairie

localities show that only a few species agree in all three particulars with the type of terebrans.

Athysanella incongrua Baker (TX-CO) is eliminated because it is monophagous on little

bluestem, Schizachyrium scoparium (Michx.) Nash, a grass that does not occur in Colorado.

More southerly species, such as A. /aeta Ball & Beamer (AZ-NM), are highly unlikely to occur

at the type-locality in northern CO. This leaves only three possibilities (with south-to-north

range in brackets):

(1) Athysanella fredonia Ball & Beamer (AZ-WY), monophagous on galleta, Hilaria jamesii
(Torr.) Benth.

(2) A. kadokana Knull (CO-AB), monophagous on alkali grass, Distichlis stricta (Torr.)

Rydb.;

6 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

(3) A. vativala Blocker (UT-AB), monophagous on sand grass, Calamovilfa longifolia (Hook.)
Scribn.

Series of these three species taken on their hosts were examined to ensure that no mixing
of populations had occurred. Only specimens of A. vativala match the figure of the female
pregenital sternite in the type of terebrans, with long, conical lobes separated by a small
median lobe. The lobes of A. kadokana and A. fredonia are low and rounded, and in the latter
the median lobe 1s wide. Therefore the type of terebrans from Colorado is clearly conspecific
with 4. vativala.

The male genitalia of A. terebrans are similar to those of A. incongrua and A. laeta, but
the latter characteristically have an apically enlarged aedeagus. The male characters cited by
Blocker to differentiate “terebrans” from “vativala” are variable within lengthy series, for
example in series collected on sand grass from 40 km S Orion, Alberta, from Colorado
Springs, Colorado, and from Fertile, Minnesota (Figs. SA-E) which represent the extremes of
its distribution on the prairies. These forms therefore represent a single species.

Athysanella (Amphipyga) occidentalis megacauda ssp. nov.
(Figs. 8 A-B)
Etymology: mega, large; cauda, tail, in reference to the aedeagal size.

Adults. Characters as in typical A. occidentalis Baker (Blocker and Johnson 19902: 126, figs.
102-104), but aedeagus 1.4 x as long as in typical subspecies, and more evenly curved, with
finer lateral serrations (Fig. 8A-B).

Types. Holotype male, CANADA. BC- Osoyoos IR 1, 119°29'W 49°04'N, 3 Oct. 1994-11
Apr. 1995 (G.G.E. Scudder) pitfall trap. Paratypes: 20 males, 1 female, 11 nymphs, same data
as holotype; 3 males, 1 female, same data, 119°32'W 49°09'N, 31 May-5 July 1994; 7 males,
8 females, same data, 119°31'W 49°13'N, 2 June-7 July 1994; 1 male, 3 females, same data,
119°32'W 49°13'N, 7 July-4 Aug. 1994. All types No. 22831 in CNCI.

Additional material: 54 males, 55 females, 6 nymphs from CANADA- BC: Chopaka,
Douglas Lake, Hedley, Kamloops, Lac du Boris, Oliver, Osoyoos, Penticton; USA- WA:
Azwell, 9 km SW Havillah, 9 km NE Tonasket, 5 km NE Vantage.

Diagnosis. Populations of A. occidentalis megacauda from central WA to Kamloops, BC
show little if any variation in aedeagal size. However, populations from the north side of the
upper Snake River valley in ID and the Bitterroot valley in adjacent MT show evidence of
introgression, having an aedeagus of intermediate length (0.40 mm) between the typical
subspecies (aedeagal length 0.35 mm, Fig. 7) and the western megacauda (0.50 mm, Fig. 8).

Additional samples of A. occidentalis from ID and CO indicate there are two more
subspecies, discussed below.

Athysanella (Amphipyga) occidentalis ladella Johnson, stat.nov.

(Figs. 6 A-B)

Athysanella ladella Johnson [in Johnson and Blocker], 1979: 383.

Diagnosis. A population on the south side of the upper Snake River valley near the UT border
is intermediate between A. occidentalis megacauda and A. ladella, a taxon described from
southern NM. This population has a broader aedeagal shaft in caudal aspect (Fig. 6A-B) than
that figured by Blocker and Johnson (1990, figs. 58-59) from the type series of /adella. This
taxon therefore becomes a subspecies of A. occidentalis Baker (1898).

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Athysanella (Amphipyga) occidentalis wilburi Ball & Beamer, stat.nov.

Athysanella wilburi Ball & Beamer, 1940: 38.

Athysanella mansa Johnson [in Johnson and Blocker], 1979: 381, syn.nov.

Diagnosis. A. wilburi (described from Kansas) appears to hybridize occasionally with
occidentalis where their ranges meet in CO. The intermediate form mansa Johnson & Blocker
probably represents local hybrids, as all three taxa are known to occur together (e.g., at
Ellicott, CO).

Athysanella (Gladionura) hicksi nom.nov.

A. (Gladionura) expulsa Blocker and Hicks, 1992: 41, preoccupied; nec A. (Amphipyga)
expulsa Blocker [in Blocker and Johnson], 1990: 112.

Etymology: patronym in honour of A.L. Hicks, who first found this species.

Remarks. This Californian species should not be confused with true 4. expulsa from OR,
discussed under A. (Pedumella) repulsa sp.nov.

Athysanella (Gladionura) hyperoche sp.nov.
(Figs. 9 A-B)
Etymology: /yperoche (noun in apposition), summit; process.

Adults. Grey-green (body faded to tan) without definite dark markings, except median lobe
of female pregenital sternite which is bordered with black. Ocellus very close to eye; crown
obtusely pointed, 0.9 x as long as wide in male, 1.0-1.05 x in female, 0.75 x as long as
pronotum in male, 0.67 x in female; hind tibial spur half as long as basitarsomere; female
pregenital sternite as in A. directa Ball & Beamer (Blocker and Johnson 19908, fig. 58), but
central conical lobe longer, distinctly produced beyond low lateral angles; male subgenital
plates large, longer than pygofers, divergent, apices sharp, directed outwards; pygofers
apically rounded, with spine on upper margin at tip, directed caudodorsad, then hooked laterad
(Fig. 9A), its length one-third that of pygofer; styles as in A. concava Ball & Beamer (Blocker
and Johnson 19908, fig. 66), but parallel-margined on apical half almost to narrowed, blunt
tip (Fig. 9B); aedeagus smooth and parallel-margined, as in concava (idem, fig. 65). Length:
male 2.9-3.2 mm, female 4.1-4.3 mm. Width of head: male 0.95-1.0 mm, female 1.05 mm; of
crown between eyes: 0.50-0.55 mm, of pronotum: male 0.9-0.95 mm, female 0.95-1.0 mm.

Types. Holotype male, USA. WY- Laramie Mts. summit [on] I-80 [16 km SE Laramie,
2500m], 24 Aug. 1969 (Ross, Ross & Miller) GL 1161 [heavily pastured range with a variety ,
of close cropped grasses”’]. Paratypes: 2 males, 2 females, same data as holotype. All types No.
228352 in CNCI.

Diagnosis. This species shares the rounded pygofer lobe and erect pygofer process of A. falla
Blocker (Blocker and Johnson 1990, figs. 72-73), but is distinguished by the much longer
pygofer process (Fig. 9A) and style (Fig. 9B). From A. concava it is distinguished by the
shorter, rounded pygofer with long, hooked dorsal process (in A. concava the pygofer is long,
tapered to a short hook, as in Blocker and Johnson 19905, fig. 68).

Biology. The types were taken along with two females of A. robusta Baker and one of A.
obesa Ball & Beamer. These latter species are both specialists on June grass, Koeleria
macrantha (Ledeb.) Schultes (=K. cristata auct.), a grass that is common in heavily grazed,
arid sites. Therefore the host of this new species also may be the same grass.

8 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Athysanella (Pedumella) castor sp.nov.

(Figs. 11B-D, F)

Etymology: castor (noun in apposition), beaver; named for the Beaverhead Mountains that
divide its range.

Adults. Grey-green with abdomen spotted and streaked with black, as in A. obesa (Beirne
1956, fig. 436); pleura and sternites often irregularly darkened; median lobe of female
pregenital sternite contrastingly black. Ocellus very close to eye; crown parabolically
produced, 0.9 x as long as wide, 1.5 x as long as pronotum; hind tibial spur absent; female
pregenital sternite short, transverse or weakly emarginate laterally, median lobe prominent to
large and rounded; male subgenital plates shorter than pygofers, divergent, apices sharp,
directed caudad with small tooth on inner margin just before tip; pygofers tapered to small
apical lobe with angle on lower margin at tip (as in Fig. 10A); ventral connective scarcely
narrowed on apical half; styles in ventral aspect as in A. attenuata Baker (Blocker and Johnson
1990a, fig. 25), but with bladelike tip strongly produced as a flange on inner margin in widest
aspect (Fig. 11F) and beyond this, tip tapered, curved ventrad; aedeagus curved cephalad,
strongly widened apically, spatulate, bearing paired serrate ridges on anterior and caudal
margins (Figs. 11B, D), in caudal aspect 4 x as long as wide. Length: male 3.1-3.6 mm, female
3.8-4.8 mm. Width of head: male 1.1-1.2 mm, female 1.2-1.25 mm; of crown between eyes:
male 0.50-0.55 mm, female 0.55-0.60; of pronotum: male 0.9.5-1.05 mm, female 1.1-1.15 mm.

Types. Holotype male, USA. MT- Badger Pass N of Bannock, 31 May 1992 (K.G.A.
Hamilton) [on Festuca mixed with Agropyron]. Paratypes: 6 males, 16 females, same data; 4
males,,8 females, /D- 8 km NE Carmen, 4 June 1992 (K.G.A. Hamilton); 1 female, JD- 11 km
E Tendoy, 29 May 1995 (K.G.A. Hamilton). All types No. 22833 in CNCI.

Diagnosis. The spatulate aedeagus is distinctive, even when reduced in size by parasitism
(Fig.11C).

Remarks. A. castor belongs to a group of species along with A. attenuata, A. expulsa Blocker
and two new species (described below). All five of these species differ from other Athysanella
in having a terminal gonopore and at least two pairs of serrated ridges on the aedeagus (Figs.
10-14). These are tentatively placed in the subgenus Pedumella (which presently has only two
unrelated species) on the basis of their short pygofer process, tapered subgenital plates with
two apical angles making them appear truncate at tips, and massive apical half of the ventral
connective.

Athysanella (Pedumella) lemhi sp.nov.

(Figs. 10 A-B, D-F)

Etymology: Lemhi (noun in apposition): this species is known only from just below Lemhi
Pass.

Adults. Colour and form as in A. castor, but male sometimes (type) with crescent-shaped
coronal marks; crown more obtuse, 0.8 x as long as wide, 1.25 x as long as pronotum; female
pregenital sternite variable, from narrowly and weakly produced medially, to broadly and
triangularly produced; male subgenital plates shorter than pygofers, divergent, apices sharp,
directed caudad with small tooth on inner margin just before tip; pygofers tapered to small
apical lobe with angle on lower margin at tip (Fig. 10A); styles broadly spatulate in widest
aspect (Fig. 10F); aedeagus curved cephalad, slightly widened apically, bearing paired serrate
ridges on anterior margin and also laterally (Figs. 10B, D), in caudal aspect 3 x as long as

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

wide. Length: male 3.5 mm, female 4.0-4.9 mm. Width of head: male 1.1 mm, female 1.25
mm; of crown between eyes: male 0.5 mm, female 0.55-0.6 mm; of pronotum: male 1.05 mm,
of female 1.2 mm.

Types. Holotype male, USA. /D- 14 km E Tendoy [16 km by road], 29 May 1995 (K.G.A.
Hamilton). Paratypes: 1 female, same data as holotype; 1 male, 8 females, 15 km E Tendoy
[17 km by road], 5 June 1992 (K.G.A. Hamilton). All types No. 22834 in CNCI.

Diagnosis. This species has genitalia similar to those of A. castor, but the aedeagus is longer
and straighter, with the serrate ridge placed laterally rather than along the caudal margin, and
the enlarged style apex is spatulate rather than tapered beyond the flange.

Athysanella (Pedumella) repulsa sp.nov.
(Figs. 14 B, D-G)
Etymology: repu/lsa, repelled; named for its allopatric distribution with respect to A. expulsa.

Adults. Female unknown. Male grey-green mottled with brown, including two crescentric
marks on crown. Ocellus set further than 2 diameters from eye; crown bluntly angled, as long
as wide, 1.3 x as long as pronotum; hind tibial spur absent; male subgenital plates short and
lyriform, as in A. expulsa; pygofers tapered to small, but elongate apical lobe with dark tooth
on lower margin at tip; styles tapered to sharp point in ventral aspect (Fig. 14G), with strongly
produced flange on upper surface which makes style apex spatulate in widest aspect (Fig.
14F); aedeagus strongly curved cephalad, slender, slightly widening towards apex in lateral
aspect, tip obliquely truncate, bearing paired serrate ridges on anterior and caudal margins
(Fig. 14B), gonopore lying in shallow dorsal depression (Fig. 14D) at narrowed tip (Fig. 14E).
Length to tip of wings: male 2.0 mm (estimated overall length 3.5 mm). Width of head 1.2
mm, of crown between eyes 0.50 mm, of pronotum 1.1 mm.

Types. Holotype male, USA. M7- 15 km W Philipsburg [1 mi E junction of Hwy. 348 and
Upper Willow Creek Road], 1 June 1992 (K.G.A. Hamilton); No. 22835 in CNCI.

Diagnosis. This species has a pygofer lobe intermediate in size between those of A. /emhi (Fig.
10A) and A. expulsa (Blocker and Johnson 1990a, fig. 27). The aedeagal shaft is much
narrower in both lateral and caudal aspect than in expu/sa (Fig. 12B), more closely resembling
that of A. attenuata (Fig. 13B), but without lateral flanges. It shares with A. expulsa a style tip
bearing a large flange on the upper surface and aedeagal shaft strongly curved, bearing serrate
margins.

Remarks. Athysanella repulsa and A. expulsa are probably sister species. Each is known from
a very limited geographical area. Their ranges are separated by the width of both the Columbia
basin of OR and the Snake River plain of ID. In this respect they resemble the sister species
Auridius vitellinus Hamilton and A. cosmeticus Hamilton which occur in the same widely
separated localities.

Diplocolenus configuratus bicolor ssp.nov.

(Fig. 15)

Etymology: bicolor, two toned; named for the unusual wing pattern (divided down the
midlength) that occurs in half the population.

Adults. Two colour morphs in nearly equal proportions. Pale morph: uniformly tan, more or

10 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

less mottled with brown as in typical D. configuratus (Uhler) (Beirne 1956, fig. 469);
bicolored morph: head brown with pale coronal margins, body and basal half of tegmina
blackish brown with narrow pale lines, apical half of tegmina contrastingly pale with brown
mottling along veins. Male subgenital plates variable in length and shape of apices, as long as
wide, tapered with tips truncate, or (especially in pale individuals) shorter and apically
divergent as in typical subspecies (Ross and Hamilton 1970a, fig. 3C); pygofers shorter than
subgenital plates, distinctly shorter than high, abruptly tapered to small apical spine at tip,
ventral margin sinuate (Fig. 15); styles evenly tapered, as in typical subspecies (Ross and
Hamilton 1970a, fig. 3B); aedeagus strongly curved in lateral aspect as in D. nigrior Ross and
Hamilton (19725, fig. 3), in caudal aspect widening towards apex, with divergent prongs at
tip as in typical subspecies (Ross and Hamilton 1970a, fig. 3D). Length: male 3.3-3.8 mm,
female 2.5-4.2 mm.

Types. Holotype male, bicolored morph, USA. WY- Wilson [W edge of town adjacent to
Teton Pass grade] 14 June 1992 (K.G.A. Hamilton) [mainly on Festuca]. Paratypes: 19
nymphs, 30 bicolored and 33 pale males, 21 bicolored and 22 pale females, same data, No.
22836 in CNCI.

Diagnosis. The short pygofer with sinuate lower margin is distinctive, as is the bicolored
morph. The genitalia most resemble those of D. nigrior, which is reduced to subspecific status
(see below).

Diplocolenus configuratus nigrior Ross & Hamilton, stat.nov.
Diplocolenus nigrior Ross & Hamilton, 19726:442.

Diagnosis. The variability of the male subgenital plates suggests that the types are descendants
of a hybrid swarm (configuratus x bicolor). If so, all three subspecies of D. configuratus
appear to hybridize where their ranges abut. Strongly truncate subgenital plates (Fig. 16)
indicate intermediates of bicolor x nigrior at 20 km E of Alpine, WY, and of configuratus x
nigrior at Lemhi Pass and Spencer, ID. The aedeagus of all these intermediates resembles that
of subspecies bicolor.

Genus Lebradea Remane, redefined
Lebradea Remane, 1959: 386.Type-species by original designation, L. calamagrostidis
Remane, 1959.

Diagnosis. By redefinition of Sorhoanus (see below), species of Lebradea are excluded from
Sorhoanus (sensu lato of Oman 1949; Beirne 1956; Hamilton and Langor 1987). They are
more slender, and have a distinctive female pregenital sternite with a long, median process
(Beirne 1956, figs. 459, 761). Males are also differentiated in having pygofer setae confined
to the caudal margin (Ossiannilsson 1983, fig. 2852) and in having the connective tip turned
dorsad just before its articulation with the aedeagus, with the sides unconnected across the
apex.

Included species. This small genus consists of the Holarctic L. flavovirens (Gillette & Baker),
the Californian L. helvina (Van Duzee: Thamnotettix), comb.nov. from Sorhoanus, and three
Old World species (Nast 1972).

Mocuellus caprillus anfractus spp. nov.
(Figs. 17 A-B, 20)
Etymology: anfractus, crooked or twisted, in reference to the curved aedeagal shaft compared

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 1]

to that of M. strictus Ross & Hamilton, which is straight.

Adults. Tawny with indefinite pale brown markings; crown longer than midline of pronotum,
as in typical M. caprillus Ross & Hamilton (Beirne 1956, fig. 448 “M. collinus”); genitalia as
in typical M. caprillus (Ross & Hamilton 19706, fig. 3B), but aedeagus in lateral aspect more
sinuate throughout length (Fig. 17A), ventral teeth of similar size (Fig. 17B) and style tip with
long, conical apical process (Fig. 20). Length to tip of pygofer (omitting setae): male 2.9-3.6
mm, female 3.3-4.2 mm.

Types. Holotype male, USA. UT7- 7 km E Laketown, 13 June 1992 (K.G.A. Hamilton).
Paratypes: 2 nymphs, 12 males, 9 females, same data as holotype; 1 male, U7T- 23 km W
Woodruff, 13 June 1992 (K.G.A. Hamilton); 3 males, 2 females, /D- 23 km SW Darlington,
7 June 1992 (K.G.A. Hamilton); 2 nymphs, 7 males, 1 female, /D- 12 km S Hamer, 19 June
1984 (K.G.A. Hamilton); 4 males, 2 females, WY- N of Jackson Hole, 15 Aug. 1971 (H.H.
Ross) GL 1265 [on Poa and Agropyron]. All types No. 22837 in CNCI.

Additional material: 5 females (without associated males) from /D - 20 km N Malad City,
20 km N Rexburg, and 15 km W of Stone.

Diagnosis. This taxon resembles M. strictus, but has a distinctly sinuate aedeagal shaft. From
typical M. caprillus it is most readily distinguished by the conical tip of the style. It is judged
to be a subspecies of the prairie leafhopper M. caprillus based on four males taken along with
two nymphs and 13 females at the summit of Lemhi Pass on the ID/MT border. Their male
style is intermediate in form (Fig. 21).

Remarks. The subspecies appears to be restricted to the upper Snake River drainage basin of
southeastern ID and northeastern UT. It is separated from the typical subspecies in
southeastern UT by Daniels Pass (2500m) 27 km SSE of Heber City between the Uinta
Mountains and the Monte Cristo Range. The typical subspecies southeast of the pass has a
narrower style tip (Fig. 22A) than populations on the plains (Fig. 22B), suggesting character
displacement at a time when the two closely related taxa were adjacent.

Mocuellus caprillus strictus stat.nov.

Mocuellus strictus Ross and Hamilton, 1970b: 174.

Diagnosis. Subspecies strictus was previously considered to be a distinct species from the
prairie-inhabiting M. caprillus, as they both occur in AB; but both taxa hybridize with
subspecies anfractus (q.v.). Presumptive hybrids of anfractusx strictus from the upper Clark
Fork valley in MT have the female pregenital segment with divergent instead of parallel-sided
notches (compare in Ross and Hamilton 19705, figs. 10, 12).

Mocuellus quinquespinus sp.nov.

(Figs. 18 A-C)

Etymology: quinque, five; spinus, spine (noun in apposition); referring to the armature of the
aedeagal tip.

Adults. Tawny with indefinite pale brown markings; crown weakly angled, as long as midline
of pronotum, as in M. larrimeri (DeLong); genitalia as in typical M. caprillus (Ross &
Hamilton 19708, fig. 3B), but aedeagus shorter, in lateral aspect with wider tip and narrower
socle (Fig. 18A), in ventral aspect with bulbous shaft and tip armed with an unpaired ventral
tooth on the lip of the gonopore at the apical third of shaft, plus 2 pairs of divergent lateral

12 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

processes, the longer pair directed caudad either side of rounded, transparent, apical knob (Fig.
18B); style short, tip with small, sharp preapical process (Fig. 18C). Length to tip of pygofer
(omitting setae): male 3.4-3.5 mm, female 3.9 mm.

Types. Holotype male, USA. UT- [SE] Tabiona [0.9 mi S junction hwys. 35 and 208 along
dry tributary of Duchesne R.], 11 June 1992 (K.G.A. Hamilton). Paratypes: 1 male, 2 females,
same data as holotype. All types No. 22838 in CNCI.

Additional material: 2 females from UT- Ouray, 4 Aug. 1986 (R.F. Whitcomb) IPL 002506,
and Zion National Park, 8 Aug. 1986 (R.F. Whitcomb) IPL 002561, both in CNCI. The first
of these is 60 km E of the type-locality, but at a much lower elevation; the latter is in
southwestern UT. Either or both may represent an undescribed species.

Diagnosis. The divergent apical processes are unique. The Palaearctic M. ruthenicus
Emel ’janov has similar male genitalia (Emel’janov 1962, figs. 84-85), but the apical processes
are parallel rather than diverging, the aedeagal shaft is tubular rather than bulbous, and the
unpaired ventral tooth on the lip of the gonopore lies at midlength of the shaft rather than at
the apical third.

Psammotettix diademata sp.nov.

(Figs. 26 A-B)

Etymology: diadema, an ornamented fillet, -ta, adjectival form; in reference to the wing
pattern.

Adults. Grey with dark brown markings; head angled, as in P. dentatus Knull (Beirne 1956,
fig. 453); tegmina 3 x as long as wide, in female marked with a circlet of 6 dark spots, in male
with more extensive markings between veins as in Hebecephalus occidentalis Beamer &
Tuthill (Beirne 1956, fig. 492); aedeagus in lateral aspect evenly curved and very narrow at
base, ballooning to membranous apical half 3 x as wide as narrowest part of shaft (Fig. 26A),
in caudal aspect strongly expanded on apical half beyond gonopore, deeply grooved on meson,
shallowly notched at tip (Fig. 26B). Length: male 4.0 mm, female 4.2 mm.

Types. Holotype male, CANADA. BC- Queen Charlotte Islands (Graham I.), North Beach,
halfway between Masset and Tow Hill 54°02'00"N 131°57'00"W, 19 Aug. 2001 (Allombewrt
& Sylvain) GG-01-052, ENT001-008907. Paratypes: 2 females, same data as holotype, but
ENT001-008897 and ENT001-008920. Holotype and paratype in Royal BC Museum;
paratype No. 22839 in CNCI.

Diagnosis. This species has an angulate head like P. dentatus, from which it may be
distinguished by the unusually bold markings and larger size. It is the most robust of the
Nearctic Psammotettix (other species that are as elongate have narrower tegmina). Its male
genitalia resemble those of P. Jatipex (Sanders & DeLong) (Beirne 1956, fig. 1204 “P.
alienus’’), but the gonopore lies further from the base and the shaft is narrower in lateral aspect.
Other Psammotettix from the Old World that have a shaft with a narrow basal half and flared
apical half have the tip pointed or rounded, not notched.

Psammotettix greenei nom.nov.
P. emarginatus Greene, 1971: 25, preoccupied; nec Sawai Singh 1969: 356.
Etymology: patronym in honour of J.F. Greene, who first named this species.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Remarks. This is a species endemic to Oregon, and should not be confused with true P.
emarginatus from India.

Psammotettix nesiotus sp.nov.
(Figs. 27 A-B)
Etymology: nesiotus (noun in apposition), an islander.

Adults. Tan, head bluntly produced but not angled, tegmina 3 x as long as wide; similar to the
abundant and ubiquitous P. /ividellus (Zetterstedt) (Beirne 1956, figs. 456, 759, 1210), but
aedeagus in lateral aspect evenly curved throughout length (Fig. 27A), in caudal aspect
parallel-margined to tip (Fig. 27B), not narrowed beyond gonopore. Length: male 2.8-3.3 mm,
female 3.0-3.5 mm.

Types. Holotype male, CANADA. BC- Bowser, [Vancouver I.], 20 June 1955 (G.E. Shewell).
Paratypes: 15 males, 36 females, same data as holotype; 9 males, 4 females, Fanny Bay,
V[ancouver] I., 20 July 1976 (K.G.A. Hamilton); 8 males, 5 females, Sea I., Vancouver
[airport] (H.H. Ross) GL 458 on Agrostis palustris. All types No. 21874 in CNCI.

Diagnosis. As in dozens of other species in the genus Psammotettix, species characters are
confined to details of the aedeagus. The evenly curved aedeagus of nesiotus is similar to that
of P. dentatus Knull (Beirne 1956, fig. 1207), but lacks the toothed lateral margins. From the
new-world P. attenuens (DeLong & Davidson) (Beirne 1956, fig. 1208) and the holarctic P.
lividellus (including its possible synonym, P. altimontanus Mitjaev, 1969) it may be
distinguished by the apical half of the aedeagus being a continuation of the shaft, and not a
distinctly flattened process that is straighter than the shaft (Fig. 28A) or recurved. Other
species in the genus do not have the aedeagal shaft arising at right angles from the socle.

Remarks. This species occurs on both sides of the Strait of Georgia. This species will
probably be found in adjacent parts of Washington state.

Genus Sorhoanus Ribaut, redefined

Sorhoanus Ribaut, 1946: 85. Type-species by original designation, Cicada assimilis Fallén,
1806.

Boreotettix Lindberg, 1952: 145, syn.nov. Type-species by original designation: Cosmotettix
serricauda Kontkanen, 1949.

Zelenius Emel’janov, 1966: 129, syn.nov. Type-species by original designation,
Laevicephalus orientalis DeLong & Davidson, 1935.

Diagnosis. A number of Nearctic species (Figs. 23-25) do not fit readily into European generic

concepts that split Sorhoanus into a number of genera based solely on male genitalia (e.g.,

Ossiannilsson 1983). Thus, the genus in its broad sense (Oman 1949 and Beirne 1956,

including species later assigned to Boreotettix Lindberg, Lebradea Remane and Zelenius

Emel ’janov) is preferred. This broad definition was expanded by Hamilton and Langor (1987)

to include Arthaldeus Ribaut and Lemellus Oman. It is here restricted, by exclusion of

Lebradea (q.v.), to those species having a combination of tenth segment undivided medially

(Fig. 23B) and male pygofers with very long, dense brush of macrosetae extending from

midlength. Cosmotettix Ribaut and Hebecephalus DeLong may have such setae, but their tenth

segment is divided or only narrowly connected across the posterior margin.

Sorhoanus involutus sp.nov.
(Figs. 23 A-E)
Etymology: involutus, rolled inwards; in reference to the unique pygofer margins.

14 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Adults. Female unknown. Male unmarked yellow except for pale dashes either side of tip of
right-angled crown, as in S. pascuellus (Fallén) (Beirne 1956, fig. 457), tegmina pale greenish
and male abdomen black; male genitalia with pygofer tapered, dorsal edges inrolled and
densely spiculate (Figs. 23A-B); tenth tergite longer than wide; internal genitalia similar to
those of S. debilis (Uhler), but aedeagus shorter and stout (Fig. 23D), with inconspicuous
dorsolateral spicules on shaft (Fig. 23E) and style tip blunter (Fig.23C). Length: male 3.6 mm.

Types. Holotype male, USA. CO- Marshall, nr Boulder, 12 June 1961 (C.H. Mann), swept
from top of low mesa; No. 22840 in CNCI.

Diagnosis. The inrolled pygofer margin is unique.

Sorhoanus virilis sp.nov.
(Figs. 24A-B, 29A)
Etymology: virilis, of the male; in reference to the large male pygofers.

Adults. Similar to S. debilis (Figs. 25A-C, 30A), with dorsum unmarked yellow, tegmina pale
green, face brown with pale arcs, abdominal tergites of female and rest of body of male black,
but tenth tergite and male pygofers much larger, the latter ending in truncate, coarsely serrate
tips (Fig. 24A); aedeagus sinuate at base, serrate on both dorsal and ventral margins (Fig.
29A), style evenly curved and without serrations (Fig. 24B). Length: male 3.7-3.9 mm, female
4.3-4.6 mm.

Types. Holotype male, USA. OR- Siskiyou, 14 June 1959 (Kelton & Madge). Paratypes: 1
male, 2 females, same data as holotype. All types No. 22841 in CNCI.

Diagnosis. The long tenth tergite is similar to that of S. involutus (Fig. 23B), but these species
are not closely related.

Sorhoanus xiphosura sp.nov.

(Figs. 32A, 33A)

Etymology: xiphos, sword; ura, tail (noun in apposition); in reference to the long spine at the
base of the aedeagus.

Adults. Similar to S. uhleri (Oman) (Beirne 1956, figs. 461, 764, 1214), with dorsum
unmarked yellow, tegmina pale green, usually influscated on apical cells and sometimes
between veins throughout, face brown (at least on margins) with pale arcs, abdominal tergites
of female and rest of body of male black; genitalia as in S. uhleri (Figs. 34-35A), but aedeagal
base sharply pointed, caudal margin not minutely serrate (Fig. 33A), and shaft often longer and
more slender (Fig. 32A). Length: male 3.5-3.9 mm, female 4.0-4.5 mm.

Types. Holotype male, USA. U7- Smithfield, 30 May 1968 (G.F. Knowlton) GL 983-984.
Paratypes: 6 nymphs, 27 males, 8 females, same data as holotype; 26 nymphs, 60 males, 75
females, 22 May-14 August, from CANADA. BC- Creston, Michael, Okanagan lakehead,
Summerland; USA. /D- Arco, Ketcham, 32 km SW Lolo Hot Springs [MT], 20 km N of
Malad City, 12 km SE Moyie Springs, 4 km N of New Meadows; M7- 9 km S of Libby, 18
km W of Ovando, 3 km W Potomoc, Wisdom; OR- Bend, Joseph and 11 km E; UT- Logan;
WA-4kmS Del Rio; WY- 13 km SE of Cooke City [MT]. All types No. 22842 in CNCI.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Diagnosis. Males with short aedeagal shafts resemble those of S. uhleri, but lack the minute
serrations on the shaft, having instead a single sharp point on the anterior angle. Males with
longer aedeagal shafts, as in S. orientalis (DeLong & Davidson) (Fig. 31A), are immediately
distinguishable by the degree to which the aedeagal base is produced. Both S. uh/eri and S.
xiphosura differ from S. orientalis and other species in Sorhoanus in having the caudal third
of the ventral connective fused into a common stem, instead of being a simple loop.

Remarks. Associated nymphs vary from entirely tan to greenish with black venters. This series
may represent a mixture of species.

Stenometopiellus vader sp.nov.

(Fig. 36A)

Etymology: vader (noun in apposition), wanderer; in reference to its disjunct distribution with
respect to S. cookei.

Adults. Characters as in S. cookei (Gillette) (Beirne 1956, figs. 428, 1182), but head markings
usually faint, female pregenital sternite with a slightly narrower median process, either with
excavated sides and rounded tip, or with straight sides converging at approximately 60°, and
aedeagus in lateral aspect most strongly curved near base, shorter by 0.1 mm (Fig. 36A).
Length: male 2.2-2.3 mm, female 2.2-2.5 mm.

Types. Holotype male, USA. /D- 12 km N Leslie, 6 June 1992 (K.G.A. Hamilton) [ca. 2000
m ]. Paratypes: 1 male, 6 females, same data as holotype. All types No. 22843 in CNCI.

Diagnosis. The short, U-shaped aedeagus distinguishes this species from its congener (Fig.
37A). The female pregenital sternite has a narrower process than that of most specimens of S.
cookei (which usually has a broadly rounded or blunt-tipped process with sides converging at
approximately 90°).

Remarks. Some females not associated with males cannot be distinguished with certainty. For
this reason, two females taken 8 km W of Anaconda, MT are not included in the type series
of S. vader although they are probably conspecific.

Unoka dramatica sp.nov.

(Fig. 1)
Etymology: dramatica, dramatic; in reference to its bold markings.

Adults. Characters as in U. gillettei Metcalf (Beirne 1956, figs. 505, 801, 1259), but more
darkly patterned (Fig. 1): head including face and abdominal tergites black or blackish-rufous,
as dark as pronotal and tegminal bands, face unmarked or with faintly indicated paler lines on
frontal arcs, and gena sometimes marked with brown lines bordering clypellus or with genal
lobes embrowned; legs deep crimson to black; scutellum with black band at base sometimes
extending across half its length; tegminal bands broad, uninterrupted, at least as wide at costa
as distance between them.

Types. Holotype male, CANADA. BC- Osoyoos L., Osoyoos IR 1, 49°04'N 119°29'W, 5-30
May 1994 (G.G.E. Scudder) pitfall trap P 3-2 [in] Purshia assoc. AN BGxh. Paratypes: 14
males, 8 females, 2 nymphs, same data as holotype, but various traps; 2 males, same locality,
6-31 May; 5 males, 13 females, same data, 30 May-4 July; 1 male, same data, 3 Oct. 1994 -
11 Apr. 1995; 7 males, 5 females, Inkaneep, Osoyoos IR 1, 49°09'N 119°32'W, 31 May-5 July

16 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

1994; 1 male, Mud L., Osoyoos IR 1, 49°13'N 119°31'W, 2 June-7 July 1994. Holotype and
29 paratypes No. 22844 in CNCI; 29 paratypes in University of British Columbia, Vancouver.

Additional material: 30 males, 15 females from CANADA. BC- Mt. Kobau 560 m, Oliver,
Osoyoos, Vaseau Ck., Venables Vly. [50° 36'30" N 121° 20'0" W], 30 May, 3 and 23 June,
16 July, 14-17 Aug. and (female only) 2 Sept. Possibly two broods (May - 4 July, 16 July - 2
Sept.)

Diagnosis. This species is closely related to U. gillettei, but has more extensive dark markings
and a darker head. The latter species usually has a rufous head, which in the darkest form has
paler brown areas across the clypellus; the pronotal band as well as the abdominal tergites may
also be paler than the tegminal bands; the tegminal bands are narrower than the white spaces
between them (Fig. 2), and the median band is often broken into spots or obsolete between the
costas.

Biology. According to G.G.E. Scudder (pers. comm.) the type-locality has much sand
dropseed, Sporobolus cryptandrus (Torrey) A. Gray. This is probably its host, since the related
U. gillettei is a specialist on this grass.

Remarks. This species is known from two widely separated localities in southern BC: the
southern Okanagan Valley and the Fraser Valley south of Ashcroft.

DELPHACIDAE

Delphacidae (Figs. 3, 38-60), in contrast to leafhoppers, exhibit considerable interspecific
variation in head, body and leg proportions; their colour patterns are generally conservative
within most genera or subgenera. The characteristic Delphacid “calcar” or hinged process at
the apex of the hind tibia is an important but unused generic character. The higher
classification of Nearctic Deiphacidae is presently under revision, and keys to the Canadian
fauna will become available in the near future. The following Delphacid taxa represent five
redefined genera with one new synonymy, one genus reduced to subgenus, four new
combinations, and nine new species.

Achorotile apicata sp.nov.
(Figs. 38A-C)
Etymology: apicata, apical; in reference to the spiny tip of the aedeagal shaft.

Adults. Colour and form as in A. acuta Scudder (1963, figs. 1 a-c), but male anal process
shorter, more strongly hooked (Fig. 38B); aedeagus straight nearly to tip, strongly constricted
beyond midlength, apex with multiple rows of spicules, apical gonopore small and round (Figs.
38A, C). Length: brachypterous male 2.5 mm, female 2.5-2.9 mm; macropterous form
unknown; width of crown between eyes 0.3 mm, of male pronotum 0.8 mm, of female 0.9 mm,
slightly wider than head; length of face above clypellus 0.5-0.6 mm, of antenna 0.4 mm, with
2™ antennal segment subequal to calcar; medial length of fore tibia 0.6 mm, of hind tibia 0.7-
0.8 mm, of hind tarsus 0.7 mm.

Types. Holotype male, USA. UT- Summit Co., 58 km S of Evanston (WY), 12 June 1992
(K.G.A. Hamilton). Paratypes, 2 males, 18 females, same data as holotype. All types No.
22845 in CNCI.

Diagnosis. This species is similar to 4. acuta, but the male has longer legs (fore tibia 0.5 mm

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

and hind tarsus 0.6 mm in 4. acuta); the most striking genital character is the aedeagus with
a distinctly shorter, more spiculate tip. Other species of Achorotile Fieber have the tip of the
aedeagus bent ventrad or with a ventral process near midlength.

Genus Caenodelphax Fennah, redefined
Caenodelphax Fennah, 1965:96. Type-species by original designation: Liburnia teapae
Fowler, 1905.

Diagnosis. This genus with a tropical genotype and 10 Nearctic species (currently placed in
Delphacodes Fieber) have males which combines a narrow crown and a black dorsum with
contrastingly pale antennae. Their calcars are small and knife-shaped, as in the transcontinental
pest species Delphacodes campestris (Van Duzee) and other wide-headed genera including
Eurybregma and Kosswigianella. Other Delphacid genera with narrow crowns have large,
foliaceous calcars. In some species of Nearctic Caenodelphax the face is unusually broad,
convex and shining; these species may be sexually dimorphic, with females pale tan without
contrasting antennae.

Included species. Included species. The only species that occurs in the PNW, Caenodelphax
atridorsum (Beamer), comb. nov. from Delphacodes, is a dimorphic species with a wide face
like that of Caenodelphax nigriscutellatus (Beamer) (see Bouchard et al., 2002). Other species
in the genus occur in eastern and central North America and will be discussed in a later paper.

Genus Elachodelphax Vilbaste, redefined

Elachodelphax Vilbaste, 1965: 14. Type-species by original designation: Liburnia metcalfi
Kusnezov, 1929.

Aschedelphax Wilson, 1992: 89, stat.nov. (subgenus). Type-species by original designation:
A. hochae Wilson, 1992.

Diagnosis. This Holarctic genus (with 1 Palaearctic species) is defined by the dark gena

contrasting with the pale face, a character otherwise only known from tropical Delphacidae

of the genus Sogatodes which have a much narrower crown. The gena is most strongly

darkened in subgenus Aschedelphax. The styles are also usually short; but in other characters

(e.g., foliaceous calcar shape) this genus resembles Javasella Fennah.

Included species. Five new combinations are created by this generic definition:
Elachodelphax (s.s.) indistinctus (Crawford), E. (Aschedelphax) bifida (Beamer) and
Elachodelphax (Aschedelphax) pedaforma (Beamer), all from Delphacodes, plus E. (A.)
coloradensis (Beamer) and E. (A.) hochae (Wilson), both from Aschedelphax.

Elachodelphax (Aschedelphax) borealis sp.nov.
(Figs. 40A, 51A)
Etymology: borealis, of the north wind.

Adults. Colour and form as in A. hochae Wilson (1992, figs. 62-66), but male pygofers
smaller, not wider than preceding segment, tips bearing only one short hook (Fig. 51 A) instead
of two; styles when viewed edgewise narrow (as in Fig. 53A), but in widest aspect truncate
with sharp, outwardly directed angles; aedeagus broad to midlength, then abruptly narrowed
to half its width, straight nearly to rounded tip (Fig. 40A). Length: brachypterous male 2.3-2.9
mm, female 2.8-3.5 mm; macropterous male 4.3 mm, female 4.4-4.9 mm; width of crown
between eyes 0.2 in male, 0.3 mm in female, of pronotum 0.8-0.9 mm, 0.1 mm wider than
head; length of face above clypellus 0.6-0.7 mm, of antenna 0.3-0.4 mm, with 2"4 antennal
segment much shorter than calcar, entire antenna not more than 0.05 mm longer than calcar;

18 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

apical 2 segments of rostrum 0.35-0.4 mm; medial length of fore tibia 0.9 mm in male, 0.8 mm
in female, of hind tibia 1.1 mm in male, 1.2 mm in female, of hind tarsus 0.9 mm in male, 0.8
mm in female.

Types. Holotype brachypterous male, USA. NH- Oakes Gulf, Mt. Washington 4700-5000"
[1400-1500 m], 8 Aug. 1954 (Becker, Munroe & Mason). Paratypes, 3 brachypterous males,
same data; 9 brachypterous males and 4 brachypterous females, same data, but 1 Aug.; 4
brachypterous males and 2 brachypterous males, same data, but 9 Aug.; 2 brachypterous males
and 1 brachypterous female, Mt. Washington, 14 Aug. 1958 (J.R. Vockeroth); 1 brachypterous
male, Mt. Washington 5000', 13 Aug. 1951 (G.S. Walley); 34 brachypterous males, 8
brachypterous and 8 macropterous females, NY- Whiteface Mt. 4600-4872', 19 July 1962 (J.R.
Vockeroth); CANADA. AB- 1 brachypterous male each, Elkwater, 9 June 1956 (O. Peck) and
Grande Prairie, 11 June 1951 (A.R. Brooks); MF (Labrador) - 1 brachypterous male and 6
brachypterous females, Goose Bay, 10-19 July 1978 (W.E. Beckel); 2 brachypterous males,
3 brachypterous females, Nutak, 26 July 1954 (E.E. Sterns); OC- 1 macropterous male,
Forestville, 9 July 1950 (R. de Ruette); 2 brachypterous males, Mt. Lyall 1500' [450 mJ, 25
July 1933 (W.J. Brown); 2 brachypterous males, 3 brachypterous females, Parke Reserve,
Kam. Co., 9 July 1957 (G.E. Shewell). All types No. 22846 in CNCI.

Additional material. Eleven brachypterous females (without associated males): 21 May-19
Sept., from CANADA. NWT- Bathurst Inlet; ON- Minnitaki; SK-Saskatoon.

Diagnosis. The styles are more obviously short and truncate than in any other species in the
genus. This common and widespread Nearctic species appears to be related to E. bifida (from
Arizona), but the styles are not bifid and the aedeagus is longer in proportion to its width, and
does not have the tips turned dorsad.

Elachodelphax (Aschedelphax) mazama sp.nov.
(Figs. 41A, 52A-B)
Etymology: Mazama (noun in apposition), a settlement northwest of the type-locality.

Adults. Colour and form as in 4. hochae Wilson (1992, figs. 62-66), but male pygofers
intermediate in size between that species and A. borealis (Fig. 52A); styles in widest aspect
with fingerlike processes directed caudolaterad; aedeagus broad on basal third, then abruptly
narrowed to half its width, straight nearly to rounded tip (Fig. 41A). Length: brachypterous
male 2.6 mm, female 2.8-3.2 mm; macropterous form unknown; width of crown between eyes
0.3 mm, of pronotum 0.9-1.0 mm, 0.1 mm wider than head; length of face above clypellus 0.6
mm, of antenna 0.35 mm, with 2"4 antennal segment much shorter than calcar, entire antenna
0.05 mm longer than calcar; apical 2 segments of rostrum 0.35-0.4 mm; medial length of fore
tibia 0.9 mm in male, 0.8 mm in female, of hind tibia 1.1 mm, of hind tarsus 0.9 mm in both
SEXES.

Types. Holotype male, USA. WA- [4 km NW] Winthrop, [ca. 18 km SE Mazama], 10 June
1984 (K.G.A. Hamilton). Paratypes: 1 male, 4 females, same data as holotype. All types No.
22847 in CNCI.

Diagnosis. Although the series from WA is short, it shows consistent differences from the
widespread A. borealis. The pronotum is 0.1 mm wider, the hind leg proportions of tibia to
tarsus are identical in both sexes instead of being sexually dimorphic, the male genitalic
capsule is 0.2 mm wider, the styles are more pointed (compare Fig. 52A to Fig. 51A) and the

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 19

aedeagus has a longer tip beyond the wide base (compare Fig. 41A to Fig. 40A).

Remarks. These two species appear to be glacial-age disjuncts of a formerly transcontinental
species.

Elachodelphax (Aschedelphax) unita sp.nov.
(Figs. 42A, 53A-B)
Etymology: Unita (noun in apposition), a mountain range in UT.

Adults. Colour and form as in A. borealis, but male pygofers narrower in lateral aspect, sternal
region oblique, producing apparent ventral extension in caudal aspect (Figs. 53A-B); styles
fingerlike, directed dorsad; aedeagus tapered, somewhat narrowed at midlength, sinuate to
pointed tip (Fig. 42A). Length: brachypterous male 2.4 mm, female 2.5-3.1 mm; macropterous
form unknown; width of crown between eyes 0.2 mm in male, 0.3 mm in female, of pronotum
0.8 in male, 0.9 mm in female, 0.1 mm wider than head; length of face above clypellus 0.6
mm, of antenna 0.3 mm, with 24 antennal segment much shorter than calcar, entire antenna
0.05 mm longer than calcar; apical 2 segments of rostrum 0.4-0.45 mm; medial length of fore
tibia 0.7-0.8 mm, of hind tibia 0.9 mm in male, 1.0 mm in female, of hind tarsus 0.8 mm in
both sexes.

Types. Holotype male, USA. UT- Cub Creek, Uinta [sic] Mts, 17 July 1952 (Bohart,
Knowlton). Paratypes: 10 females, same data as holotype. All types No. 22848 in CNCI.

Diagnosis. The male genitalia are distinctive.

Genus Eurybregma Scott, redefined

Eurybregma Scott, 1875. Type-species by monotypy: E. nigrolineata Scott, 1875: 92.
Diagnosis. The North American species formerly included in Eurysa Fieber and later,
tentatively placed in Chilodelphax Vilbaste (Wilson 1988), do not belong in either of these
genera. Chilodelphax is only superficially similar to one of the Nearctic species in male
genitalia, but the external characters (e.g., narrow crown) are quite different. Eurysa has a
spotted face found in the new world fauna only in Phyllodinus Van Duzee and Bakerella
Crawford. Nymphs of Eurysa have few abdominal pits (Ossiannilsson 1978), while nymphs
of Eurybregma (including the Nearctic species) have characteristic rows of numerous
abdominal pits (Anufriev 1987, fig. 3). This pit arrangement differs from those of Achorotile
and Laccocera in having the most medially located pit removed at a greater distance than the
others.

Included species. This data necessitates the removal of three North American species formerly
in Eurysa to new combinations: Eurybregma magnifrons (Crawford), Eurybregma montana
(Beamer) and Eurybregma obesa (Beamer).

Eurybregma eurytion sp.nov.

(Figs. 43A, 44A-B, 54B)

Etymology: eurys, widespread; -tion, result; in reference to the ubiquity of this species in
PNW grasslands wherever its sister species does not occur.

Adults. Colour and form as in E. magnifrons (Beamer 1952, fig. 2), but slightly narrower,
head slightly narrower than pronotum in male, as wide as pronotum to slightly wider in female;
anal process tiny; pygofer process inturned on upper margin (Fig. 54B); aedeagus straight

20 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

nearly to tip, upper margin at tip sinuate (Fig. 44A) or (in southern morph, Fig. 43A) “kinked,”
lower margin spiculate, convex on apical third, apex serrate around apical gonopore (Figs.
43A, 44A-B). Length: brachypterous male 2.5-3.2 mm, female 2.7-3.5 mm; macropterous male
4.1-4.4 mm, female 4.2-4.7 mm; width of crown between eyes 0.3-0.4 mm, of head and
pronotum 0.8-0.9 mm; length of face above clypellus 0.5-0.6 mm, of antenna 0.3 mm, with 2"
antennal segment subequal to calcar; apical 2 segments of rostrum 0.35-0.4 mm; medial length
of fore tibia 0.7 mm, of hind tibia 0.8-1.0 mm, of hind tarsus 0.8 mm.

Types. Holotype male, USA. /D- 23 km SW Darlington, 7 June 1992 (K.G.A. Hamilton).
Paratypes: 8 males, 10 females, same data as holotype; 1 male and 5 females, all macropterous,
from /D- Mud Lake, 19 June 1984 (K.G.A. Hamilton) on Agropyron; 4 males, 2 females, all
macropterous, WA- 9 km S of Methow, 10 June 1984 (K.G.A. Hamilton); 1 nymph, 38
brachypterous males and 41 brachypterous females, 1 macropterous male and 5 macropterous
females, 25 May-19 June and 22 July, from CANADA. BC- 7 km W Douglas Lake, 5 mi W
Hedley, Mission Flats [nr Kamloops], Princeton [airport]; USA. /D- 6 km S Baker, Galena
Summit 2800m [S of Obsidian], 10 km E Howe, 12 km N Leslie, Lowman, and 37 km NE;
MT- 5 km N Conner, Florence, Ketcham, 7 km W Lincoln, 8 km N Missoula, 18 km W
Ovando, 15 km W Philipsburg, 6 km SE St. Regis; OR- Bend (Pilot Butte), 13 km E Joseph;
WA-5 km SW Chesaw, 7 km N Havillah, Orient, 12 km E Teanaway. All types No. 22849 in
CNCI.

Additional material. Southern morph: 11 males and 8 females, all brachypterous, 8-13 June,
from USA- /D: 20 km N Malad City; UT: 4 km SW Lakeside Resort, 7 km E Laketown. No
associated males: 21 brachypterous females, 30 May-26 July, from CANADA- BC: Clinton,
Jesmond, Osoyoos, Kamloops, 2 mi S Louis Creek, Venables Valley Rd. at Hwy. 97; USA-
ID: Obsidian; MT: 8 km W Anaconda, Florence, 10 km N Three Forks, Cardwell; WA: 8 km
S Grand Coulee.

Diagnosis. This species differs from E. magnifrons (Figs. 45A-B, 55B) in its stouter aedeagal
shaft which is straight on the basal half, and in its inturned pygofer process. The ranges of
typical eurytion and magnifrons abut at Princeton, BC without any sign of introgression.

Genus Kosswigianella Wagner, redefined

Kosswigianella Wagner, 1963: 169. Type-species by original designation: Delphax exigua
Boheman, 1847.

Acanthodelphax LeQuesne, 1964: 57, syn.nov. Type-species by original designation: Del/phax
denticauda Boheman, 1847.

Diagnosis. Spatulate antennae with the apical segment not more than twice as long as wide,

and a contrastingly dark male abdomen, are synapomorphies linking the genera Kosswigianella

Wagner and Acanthodelphax LeQuesne. A new species, described below, shows the close

relationship between these taxa in having male genitalia combining the truncate pygofers and

drop-shaped basal aedeagal attachment of Kosswigianella and ventral pygofer tooth of

Acanthodelphax.

Included species. Synonymy of these genera creates new combinations for the species
formerly in Acanthodelphax: the North American K. analis (Crawford) and 3 Palaearctic
species, K. denticauda Boheman, K. spinosus (Fieber) and K. transuralica Anufriev.

Kosswigianella irrutilo sp.nov.
(Figs. 3, 46A, 56A-B)
Etymology: irrutilo, become ruddy.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 21

Adults. Redbrown, male (and sometimes female) abdomen deep chocolate brown. Head as
wide as pronotum in male, slightly narrower in female; face above clypellus broad, 25% longer
than wide, weakly inflated, smooth and shining with median carina scarcely evident; crown
slightly wider than long in male, as long as wide in female; antenna short, 2™ segment
spatulate as in Kosswigianella exigua (Boheman). Thorax with pronotal carinae short, curved
outwards, ending near midlength of segment; brachypterous tegmina truncate, 25% longer than
wide, covering only extreme base of abdomen, leaving 5 large abdominal segments exposed;
hind tibia bearing two minute spines on outer edge; calcar small and knife-shaped. Male anal
process strongly angled ventrad; genital ring with pygofers truncate, minute median process
on venter (Fig. 56B); diaphragm transverse, with inturned upper edge; styles abruptly
narrowed just beyond midlength, apices furcate (Fig. 56A); thecal ring drop-shaped, narrow
upper end articulated between bases of anal tube; aedeagus lamellate, in lateral aspect sinuate
with minute tooth near base and gonopore confined to narrowly rounded tip (Fig. 46A).
Length: brachypterous male 1.6 mm, female 1.7-2.1 mm (only specimen of macropterous form
with wing tips damaged); width of crown between eyes 0.3 mm, of head and pronotum 0.6 mm
in male, 0.7 mm in female; length of face above clypellus 0.4 mm, of antenna 0.25-0.3 mm,
with 2" antennal segment subequal to calcar; apical 2 segments of rostrum 0.25-0.3 mm;
medial length of fore tibia 0.3-0.35 mm, of hind tibia 0.5-0.55 mm, of hind tarsus 0.4-0.45
mm.

Types. Holotype brachypterous male, USA. CO- Tarryall, 17 June 1954 (H.H. Ross) GL 947.
Paratypes: 1 brachypterous male, 180 brachypterous and 1 macropterous females, same data
as holotype. All types No. 22850 in CNCI.

Diagnosis. The broad, shiny face, large anal processes, furcate styles and lamellate aedeagus
are distinctive.

Kosswigianella wasatchi sp.nov.
(Figs. 47A, 57A)
Etymology: Wasatch (patronym), a mountain range in UT.

Adults. Colour and form as in K. analis (Crawford) (Figs. 48A-B, 58A), but face above
clypellus broader, 45% longer than wide, crown less strongly produced; diaphragm narrowed
either side of broad, median process on dorsal edge, and bearing prominent “heel” between
bases of styles (Fig. 57A); aedeagus as in K. analis (Fig. 48A), but longer and more slender
(Fig. 47A). Length: brachypterous male 1.8-1.9 mm, female 2.3-2.4 mm; macropterous malé
unknown, female 3.2-3.3 mm; width of crown between eyes 0.2 mm in male, 0.25 mm female,
of male pronotum 0.7 mm, of female 0.8 mm, 0.1 mm wider than head; length of face above
clypellus 0.4 mm in male, 0.5 mm in female, of antenna 0.25 in male, 0.3 mm in female, with
2™ antennal segment subequal to calcar; apical 2 segments of rostrum 0.25-0.3 mm; medial
length of fore tibia 0.4 mm in male, 0.5 mm in female, of hind tibia 0.7 mm in male, 0.8 mm
in female, of hind tarsus 0.5 mm in male, 0.6 mm in female.

Types. Holotype male, USA. UT- 11 km SW of Scofield, 10 June 1992 (K.G.A. Hamilton).
Paratypes, 1 male, 5 brachypterous females, 2 macropterous females, same data as holotype.
All types No. 22851 in CNCI.

Diagnosis. This species is a sister-species of the transcontinental K. analis, from which it may
be distinguished by the slender aedeagal shaft and differently shaped diaphragm.

22 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Nothodelphax venusta (Beamer), comb.nov.
Delphacodes venusta Beamer, 1948: 115.
This is a sister-species to N. foveata, the type-species of the genus.

Genus Paraliburnia Jensen-Haarup, redefined
Paraliburnia Jensen-Haarup, 1917: 2. Type-species by original designation: P. jacobseni
Jensen-Haarup, 1917 [=Delphax concolor Fieber, 1866].

Diagnosis. This broad-headed genus has an unusually long rostrum (apical two joints together
0.55-0.7 mm, at rest extending at least to metatrochanters) and very large, foliaceous calcars
similar to those of the narrow-headed genus Megamelus Fieber. New World species also have
close-set arms of the forked median carina extending onto the face between the eyes, although
the smoothness of the apex of the head in some individuals makes these carinae hard to
distinguish.

Included species. There are two Palaearctic species (Nast 1972). One Nearctic species
Paraliburnia kilmani (Van Duzee: Liburnia) comb.nov. is transferred from Delphacodes.
Two new, related species are described below.

Paraliburnia furcata sp.nov.
(Figs. 49A-B, 59A-B)
Etymology: furcata, forked; in reference to the styles.

Adults. Colour and form as in P. kilmani (Van Duzee) (Wilson 1992, figs. 125-127), but much
larger; crown of head slightly wider than long in male; anal process more robust (Fig. 59B);
diaphragm narrowed mesally and produced caudad; styles small, abruptly narrowed on apical
third, appearing furcate (Fig. 59A); aedeagus unarmed, tip dorsoventrally flattened above
subterminal gonopore (Figs. 49A-B). Length to tip. of brachypterous tegmen 2.5 mm
[estimated overall length 3.0 mm]; width of crown between eyes 0.3 mm, of head 0.9 mm, of
pronotum 1.0 mm; length of face above clypellus 0.7 mm, of antenna 0.55 mm, with oe
antennal segment one-third length of calcar; apical 2 segments of rostrum 0.7 mm; medial
length of fore tibia 0.9 mm, of hind tibia 1.0 mm, of hind tarsus 0.5 mm; calcar 0.5 mm;
macropterous form and female unknown.

Type. Holotype male, CANADA. BC- Quesnel, 12 June 1949 (G.J. Spencer); No. 22852 in
CNCI.

Diagnosis. In size and genital characters this species is most similar to the European genotype,
but the styles are short and appear furcate, and the aedeagus is unarmed.

Paraliburnia lecartus sp.nov.

(Figs. 50A, 60A-B)

Etymology: /ecartus, arm (noun in apposition); in reference to the shape of the aedeagal
shaft.

Adults. Colour and form as in P. kilmani (Wilson 1992, figs. 125-127), but with head wider,
crown slightly wider than long in male (as in P. furcata) and as wide as long in female; anal
process smaller; pygofer strongly bowed outwards above lower margin (Fig. 60B); diaphragm
weakly sclerotized; styles short, strongly tapered, divergent (Fig. 60A); aedeagus armlike in
lateral aspect, tapered towards enlarged tip, with ventral margin straight, upper margin concave

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 23

and serrate, sides bearing unequally aligned serrate rows in a spiral pattern, that of the right
side being most dorsally situated near the tip, but more ventrally placed near the base (Fig.
50A); gonopore just below tip. Length: brachypterous male 2.3 mm, female 2.1mm, width of
crown between eyes 0.25 mm, of head 0.75 mm, of pronotum 0.8 mm; length of face above
clypellus 0.55 mm, of antenna 0.4 mm, with 2™ antennal segment slightly shorter than calcar;
apical 2 segments of rostrum 0.7 mm; medial length of fore tibia 0.7 mm, of hind tibia 0.9-0.95
mm, of hind tarsus 0.4 mm; calcar 0.35 mm.

Types. Holotype male, CANADA. BC- Fort St. John, Peace River, 10 June 1959 (G.G.E.
Scudder); paratype female, same data; both types No. 22853 in CNCI.

Diagnosis. No other species in this genus has as small a calcar. The spiral arrangement of
aedeagal serrations is also unusual.

ACKNOWLEDGEMENTS

This study is dedicated to the memory of H.H. Ross (1908-1978). It is the outcome of
many years of leafhopper surveys by “Herb” and his students to document the leafhopper
fauna of North American grasslands, with special focus on Rocky Mountain grasslands. The
manuscript was reviewed by D. Lafontaine of AAFC, Ottawa.

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1998b. New species of Hebecephalus DeLong from Bnitish Columbia, Idaho and adjacent states
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79-100.

26 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

y

Figures 1-4. Homoptera-Auchenorrhyncha of PNW grasslands. 1, Unoka dramatica, palest
form, Cicadellidae; 2, Unoka gillettei, darkest form; 3, Kosswigianella irrutilo, Delphacidae
(brachypterous form); 4, Bruchomorpha beameri, Caliscelidae (macropterous form).

Figures 5-9. Male genitalia of Cicadellidae. 5, Athysanella terebrans, variation in lateral
aspect of aedeagus (A-C) and distal part of style in widest aspect (D-E); 6, A. occidentalis
ladella, lateral aspect of aedeagus (A) and tip of aedeagus in caudal aspect (B); 7, same, of A.
occidentalis s.s.; 8, same, of A. occidentalis megacauda; 9, A. hyperoche, tip of pygofer (A)
and distal part of style in widest aspect (B). Scale line: 0.1 mm.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 27

Figures 10-14. Male genitalia of Cicadellidae: genitalic capsule in lateral aspect (A), same,
of aedeagus (B), same, parasitized individual (C), tip of aedeagus in dorsal aspect (D), same,
in caudal aspect (E), distal part of style in widest aspect (F) and in narrowest aspect (G). 10(A-
B, D-F), Athysanella lemhi; 11(B-D, F), A. castor ; 12(B, D-E), A. expulsa; 13(B, D), A.
attenuata; 14(B, D-G), A. repulsa. Scale line: 0.1 mm.

28 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Figures 15-25. Male genitalia of Cicadellidae. 15, Diplocolenus configuratus bicolor,
genitalic capsule in lateral aspect; 16, same, of D. configuratus nigrior; 17, Mocuellus
caprillus anfractus aedeagus in lateral aspect (A) and in caudal aspect (B); 18, same, of MW.
quinquespinus, with distal part of style in widest aspect (C); 19, M. caprillus strictus, distal
part of style in widest aspect ; 20, same, M. caprillus anfractus; 21, same, of M.
caprillus'\\anfractus from Idaho/Montana border; 22, same, of M. caprillus s.s. from Utah (A)
and from Alberta (B); 23, Sorhoanus involutus, genitalic capsule in lateral aspect (A), in
dorsal aspect (B), distal part of style in widest aspect (C), aedeagus in lateral aspect (D) and
in caudal aspect (E); 24, S. virilis, genitalic capsule in lateral aspect (A) omitting pygofer
setae, and distal part of style in widest aspect (B); 25, same, in S. debilis, with detail of pygofer
apex in ventrolateral aspect (C). Scale line: 0.1 mm..

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 29

ws r I
—--2 ee oe ee :

oe

Cs Sr ae

swabs eba eee
bs .

weet

Figures 26-37. Aedeagi of Cicadellidae, lateral aspect (A) and caudal aspect (B). 26,
Psammotettix diademata; 27, P. nesiotus; 28, P. lividellus; 29, Sorhoanus virilis; 30, S.
debilis; 31, S. orientalis; 32, S. xiphosura, holotype; 33. same, variety; 34-35, S. uhleri,
varieties; 36, Stenometopiellus vader; 37, Stenometopiellus cookei. Scale line: 0.1 mm.

30 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Figures 38-50. Male genitalia of Delphacidae: lateral aspect of aedeagal shaft (A), anal tube
(B), and gonopore aspect of aedeagal shaft (C). 38, Achorotile apicata; 39, A. acuta; 40,
Elachodelphax borealis; 41, El. mazama; 42, El. unita; 43, Eurybregma eurytion from Malad
City, ID; 44, Eu. eurytion, smallest specimen from same locality as holotype; 45, same, of Eu.
magnifrons; 46, Kosswigianella irrutilo; 47, K. wasatchi; 48, K. analis; 49, Paraliburnia
furcata; 50, P. lecartus. Scale line: 0.1 mm, same scale for each genus.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 31

Figures 51-60. Male genitalia of Delphacidae: caudal aspect of genitalia (A), omitting
aedeagal shaft and (in Figs. 57-58), omitting small styles; lateral aspect of genital capsule and
anal tube (B). 51, Elachodelphax borealis; 52, El. mazama; 53, El. unita; 54, Eurybregma
eurytion; 55, Eu. magnifrons; 56, Kosswigianella irrutilo; 57, K. wasatchi; 58, K. analis; 59,
Paraliburnia furcata; 60, P. lecartus. Scale line: 0.1 mm (Fig. 56A-B slightly larger).

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 33

Homoptera (Insecta) in Pacific Northwest grasslands.
Part 2 — Pleistocene refugia and postglacial dispersal
of Cicadellidae, Delphacidae and Caliscelidae

K.G. ANDREW HAMILTON

BIODIVERSITY PROJECT, RESEARCH BRANCH, A.A.F.C.
C.E.F. OTTAWA, ONTARIO, CANADA KIA 0C6

ABSTRACT

Biogeographic analysis suggests that 241 Cicadellidae, 33 Delphacidae and 1 Caliscelid
are restricted to grasslands in the Pacific Northwest. Nearly half of these (120 or 44%) are
endemics. This grassland endemic fauna is third only to those of the prairies and desert
grasslands. Of these, only six taxa probably postdate retreat of continental glaciers. The
present distribution of the older taxa indicates that this fauna are descended from nine
main glacial-era refugia (in descending order of importance): (1) east slopes of the
Cascade Range, from Washington state to southern Oregon; (2) Columbia basin including
Palouse hills of Washington and canyons of western Idaho; (3) south-facing slopes on the
Rocky Mountains of Montana and western Wyoming; (4) the headwaters of the Snake
River and south-facing slopes north of the Snake River in southern Idaho; (5) east of the
Coast Range of Oregon; (6) edges of glacial Lake Missoula in western Montana; (7) a
periglacial grassland near the ice front in Alberta; (8) the mountains of south-central
Oregon; and (9) the coast of the Queen Charlotte Islands. Additional refugia might have
been in the mountains of Colorado, Utah, eastern Arizona, and eastern Wyoming, where
there are an additional 22 endemics. Postglacial warming brought grasslands and their
endemic insects to British Columbia on the islands of the Strait of Georgia, to the East
Kootenay valley and to the upper Fraser River of BC. Faunal exchanges have occurred
across at least nine mountain passes on the continental divide. Three of these passes still
provide continuous grassland connections between the prairies and the intermontane
grasslands, yet not more than 14 slow-moving prairie species have surmounted any one
pass.

INTRODUCTION

Pacific Northwest grasslands, characterized in Part 1 of this study, are widely scattered
across a largely mountainous landscape extending across six states and two provinces (Fig. 1).
Specifically to the context of grasslands, Pacific Northwest (PNW) refers to the Great Basin
and adjacent deserts, grasslands and open forests west of the continental divide extending from
latitude 42° N both northwards and up slopes in mountainous areas to coniferous forests. This
area encompasses all but the mountains of the states of Idaho (ID), Oregon (OR) and
Washington (WA), plus the western parts of Montana (MT) and Wyoming (WY). Grassy
intermontane valleys of southern and central British Columbia (BC) and on its coastal islands,
plus foothills prairie of Alberta (AB) are also included in this faunal region. Similar grasslands
extend southeast (Fig. 2) into the mountains of eastern Arizona (AZ), Colorado (CO) and Utah
(UT). Low elevation grasslands of AZ, California (CA), western CO, Nevada (NV) New
Mexico (NM) and UT are considered to be desert grasslands of the Great Basin and Sonoran
subregions, or Mediterranean-zone shrublands.

PNW grasslands have undergone many vicissitudes during the last million years. Glaciers
carved out most of the valleys in BC down into northern WA, ID and MT; other valleys in
western MT have been inundated by glacial Lake Missoula as recently as 12,000 years ago;

34 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

and the Columbia Basin of WA has been repeatedly scoured by catastrophic floods emanating
from glacial Lake Missoula (Waitt and Swanson 1987). It has been thought that, during the
height of glaciation, tundra extended as far south on lowlands as WY and the Palouse hills of
eastern WA, and in the mountains, as far south as southern CO and UT (Brunnschweiler 1962).
Increased rainfall and colder temperatures also would have resulted in continuous spruce forest
throughout the rest of the PNW. Such enormous changes in the PNW should have resulted in
decimation of PNW grasslands by a combination of cool, wet weather and localized
catastrophic events. That any fauna could survive appears almost incredible. Wholesale
replacement of any grassland and its endemic insect fauna with widespread grassland species
from southern refugia is the expected outcome. During the warmest postglacial period (the
Hypsithermal) grasslands could have invaded the mountains from the prairies; but if so,
grasslands must have been more extensive and less discontinuous in the PNW than at present.

It would seem unlikely therefore that PNW grasslands could ever have constituted a
continuous, discrete ecozone. Yet many Homoptera are unique to these grasslands (for
examples of new taxa, see Part 1 of this study). Furthermore, some are distributed relatively
uniformly throughout the extent of the PNW. For example, Texananus extremus (Ball) not
only ranges throughout the Cordilleran region, but is also found on the foothills prairie of
southwestern AB. The purpose of this paper is to explore this paradox on the basis of evidence
of recent insect distributions, and to deduce the factors responsible for this distinctive faunal
assemblage. This study is the second detailed project emanating from a larger effort to study
the leafhoppers endemic to North American grasslands, initiated by H.H. Ross in 1952. An
overall summary was presented orally (Hamilton 1993) and a detailed study of the leafhopper
fauna of the Yukon has been completed (Hamilton 1997).

Leafhoppers (Cicadellidae) are particularly important in characterizing native grasslands.
They include the largest number of insect species endemic to the Great Plains (Ross 1970).
There are more than 200 species endemic to prairies, and another 136 known only from desert
grasslands, part of a total of some 800 grassland species in more than 80 genera (Hamilton and
Whitcomb 1993). These insects contrast with other grassland arthropods such as spiders and
ground beetles in being comparatively inefficient dispersers little influenced by microhabitat.
Their dispersal rates are usually slower than 1 km/yr (Hamilton 1999a). Smaller numbers of
Fulgoroidea (planthoppers) belonging to the families Delphacidae and Caliscelidae show
similar endemism although (on limited data available) their dispersal appears to be more rapid
than that of leafhoppers. Data are also available on grassland Delphacidae from the Yukon
(Wilson 1997).

METHODS

Materials. Ranges and host associations of Homoptera are based mainly upon surveys of
PNW grasslands carried out by the author in July-August 1976, August 1978, June 1984,
August 1985, May-June 1987, May-June 1992 and May 1995, which yielded samples of more
than 57,000 specimens. Particular attention was focussed on 15 of the lowest Rocky Mountain
passes between British Columbia and Wyoming (Table 1), plus adjacent grasslands at lower
elevations. These records were supplemented by collections made by R.F. Whitcomb of USDA
from 1977-1997 in most western states including MT, UT and WY, plus incidental records
from the H.H. Ross “GL” (grassland) survey and other material in the Canadian National
Collection of Insects (CNCI). Data from revisionary works (Lindsay 1940; Kramer 1971a;
Johnson and Blocker 1979; Blocker and Johnson 1988, 1990; Hamilton and Zack 1999;
Bartlett and Deitz 2000), and the only regional faunas, for WA (Wolfe 1955) and BC (Maw
et al., 2000) are also cited; these represent thousands of additional specimens. Host records
are those confirmed by my collecting, unless otherwise noted. For other recent revisions and

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 35
taxonomic methodology, see Part 1.

Table 1.
Rocky Mountain passes sampled for Homoptera, BC to WY, lettered as in Figure 3. Omitted
because not sampled: Deer Lodge Pass (1900 m), Elk Park, Flesher, Homesteak and Stemple
passes in MT (2000 m), and on the ID/MT border, Reynolds and Red Rock passes (2100 m)
and Bannack Pass (2300 m). Species numbered as in faunal list; hybrids (e.g., 14x87) are
recorded for both parental populations.

elevation direction | species in species through
pass near pass pass?
700 m 27, 30 1, 8, 10, 14
21500
Kicking 1700 m
Horse

“Crooked
River” (A)

Yellowhead

BC
BC
0).19526; 1, 2,7, 10, 24
28, 51, 68
M 1,11, 16 Wye 022;
75 02

Lolo

ie: ek eae a!
es ae
Rogers (C) Bu 1800 m
31
es i

MacDonald

(D)

MT/ID 1800 m 57, 67 eae

MT 2000 m 13, 30 Tele 12
57,68
“Grassy” (E) MT 2100 m N/S (open) 28, 43, 73 See) loa ie
22, 33, 38,
oy Paes

Lost Trail (F) ID/MT 2100 m 22, 43,68, 5, 32, 41, 57,
72 61, 69
Lemhi (G) ID/MT 2200 m E 5 ea ee? je 19,31, 51,
(open) 36, 43, 72, 8
73
8

3
7 4,5, 22

Crowsnest (B) | BC/AB 1400 m ae

N/S
E/W
E/W
E/W
N/S
N/S
E/W
N/S
N/S
E/W

/W

Bannock (H) | ID/MT | 2200m_ | N/S(open) | _ 3, 22, 29,

36, 41, 43,

51, 53, 72,

73
Monida (J) ID/M 2100m | N/S(open) | 33, 34, 43, 24, 51 0
57, 61
Great Divide WY 2000 m all (open) 3, 17, 20 + 9,15, 19, 21
Basin (K) 23, 25 29, 31, 50

Biogeography of Homoptera. The PNW grassland fauna includes three introduced
Eurasian species, the beet leafhopper Neoaliturus (Circulifer) tenellus (Zetterstedt), a sedge-
feeding leafhopper Euscelis obsoleta (Kirschbaum), and the planthopper Toya propinqua
(Fieber). They are excluded from this analysis because they were not part of the prehistorical
fauna of the region.

As a first step in this biogeographic analysis, prehistorical patterns of endemism must be
distinguished from stochastic patterns derived from rapid, opportunistic dispersal. For this
reason, wind-dispersed “‘microleafhoppers” (Cicadellidae of the subfamily Typhlocybinae) are
excluded from this biogeographic analysis. Other leafhoppers (“macroleafhoppers”’) include
a few migratory species, of which only Exitianus exitiosus (Uhler) is a grassland species.

Ril

28
5]
24
4,5
32
2

36 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Planthoppers of the families Delphacidae and Caliscelidae also are used to determine whether
the PNW fauna is essentially a prairie fauna, or an endemic intermontane fauna that survived
glaciation. Slowest of all are “flightless” species of the genus Errhomus (although males can
fly, their wingless females cannot have dispersed long distances: Hamilton and Zack 1999).
Such insects are important in elucidating where glacial refugia could have been located.

Second, native grassland species must be distinguished from those with wider ranges. The
former either feed exclusively on grasses, or are always associated with grassland sites
provided that they do not feed on trees. In cases where their hosts are unknown, they are
presumed to be similar to those of the most closely related species. This latter criterion is
usually sufficient to distinguish tree-feeding species which are almost always in separate
genera from those feeding on forbs and grasses. Many genera of leafhoppers and planthoppers
feed exclusively on grasses, but this is a less reliable criterion since such genera often include
sedge feeders of fens and bogs.

Almost all planthoppers of the families Delphacidae and Caliscelidae in the PNW are
restricted to grasslands, with the exception of the rush and sedge-feeding Delphacid genera
Achorotile Fieber, Kelisia Fieber, Megamelus Fieber, Pentagramma Van Duzee and
Stenocranus Fieber, plus eleven common and widely dispersing species: one sedge-feeding
Caliscelid, Peltonotellus histrionicus Stal; and ten Delphacids, Delphacodes campestris (Van
Duzee), D. consimilis (Van Duzee), D. lutulenta (Van Duzee), D. magna (Crawford), D.
puella (Van Duzee), Javesella pellucida (Fabricius), Liburniella ornata (Stal), Phyllodinus
nervatus Van Duzee, Pissonotus delicatus Van Duzee and Stobaera tricarinata (Say).

The 38 leafhopper target genera are discussed in the first part of this project. Three genera
are reported but not analysed here. Ballana (over 100 described species: DeLong 1964) needs
revision. Preliminary analysis of this genus suggests that most of the species are confined to
California and the Sonoran subregion, but that distribution patterns similar to those discussed
below will be found in the PNW. The leafhopper genera Deltocephalus Burmeister (13
described species: Kramer 19715) and Draeculacephala Ball (7 species complexes in northern
North America: Hamilton 1985) present unresolved taxonomic and biological problems, with
many (but not all) species restricted to sedge fens rather than to grasslands.

With these exceptions, the complete PNW fauna of “macro-leafhoppers” and planthoppers
was examined to determine quantitative degrees of endemism, and to deduce dispersal rates.
First, all native species of leafhoppers and planthoppers known from PNW grasslands were
sorted according to their overall distribution patterns and known biology. Species that range
into northern grasslands, prairies, or southern (“Great Basin” and “Sonoran subregion’)
grasslands are distinguished from those that appear to be truly endemic. Next, species that
occur along the continental divide are individually analysed to determine whether dispersal
patterns are discernible from present distributions. Finally, centres of endemism are compared
for evidence of glacial-age refugia.

Grassland characterization. PNW grasslands are diverse, both geographically and
botanically. The following is a synopsis drawn from many sources, including my own
observations of areas sampled from surveys extending over seven years. For example, good
collecting areas are usually found on south-facing slopes in Canada (Ross 1970) but farther
south these become too arid. West-facing slopes are preferred by leafhoppers in the PNW.

For this analysis, PNW grasslands are assumed to be biologically similar to small (possibly
relict) grasslands in areas outside the PNW, such as south-facing hillsides of the Yukon
(Hamilton 1997). This study identifies such grasslands in the drier valleys of western and
central British Columbia, and west-facing hillsides in the Rocky Mountains as far south as CO.
These grasslands are subdivided into nine disjunct areas (Fig. 2 A-J), from west to east:

(A) coastal areas of Pacific islands from the Queen Charlotte Islands (Fig. 2, star) of

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 37

northwestern BC, south to small islands in Puget Sound of northwestern WA;

(B) Pacific coastal grasslands, grassy “balds” on hills of the Coast Range, and valleys between
the Coast and Cascade ranges of OR;

(C) the eastern slopes of the Cascade Range, from WA to the California border;

(D) the mountains of south-central OR;

(E) the Columbia basin including Palouse hills of WA-OR, adjacent canyons of western ID,
and intermontane valley systems of central BC as far north as the Fraser River;

(F) the Snake River valley in southern ID and south-facing slopes north of the Snake River;

(G) the Upper Clark Fork valley system in western MT [for detailed analysis of dispersal
patterns, subdivided into the smaller Blackfoot (Fig. 3, GA), Bitterroot (GB), and Rock
Creek (GC) valleys];

(H) prairies east of the Continental Divide, from AB south to the Bighorn Mountains of
northern WY; and

(J) a southward chain of high altitude grasslands in the mountains of eastern ID, western and
southern WY, north-central CO, northern and central UT and eastern AZ.

Botanically, PNW grasslands form a complex of habitats (Munroe 1956; Kiichler 1985;
Ricketts et al. 1999) that are of four main types (Fig. 4):

(1) the largest area, extending from isolated valleys in BC south into WA and ID was
originally Palouse prairie, dominated by mixed grasses generically similar to those on the
prairies (Ecoregion 53 = Kiichler zone 43);

(2) extensive sagebrush steppe (Ecoregion 75 = Kiichler zone 49) has an ecotonal fringe of
Palouse grasslands dominated in less arid areas by drought-hardy wheat grass (Agropyron
spp.) and rice grass (Oryzopsis spp.);

(3) foothills prairie dominated with fescues (Festuca spp.) form a narrow altitudinal grassland
on the eastern slopes of the Rocky Mountains (Ecoregion 57 = Kiichler zone 56); and

(4) a limited area of oak savannah once occupied the Willamette Valley between the Coast
Range and the Cascades (Ecoregion 6 = Ktichler zone 24).

RESULTS

Using admittedly subjective criteria, the Homopterous fauna of PNW grasslands, including
adjacent montane grasslands, is calculated to consist of 215 species and 26 subspecies of
“macro-leafhoppers” plus 34 species of planthoppers. This totals 275 taxa with a wide variety
of dispersal characteristics (Faunal Synopsis and Index).

Species characteristic of boreal grasslands spread readily across mountain divides. There
are 36 such leafhoppers that range as far north as the Yukon (Hamilton 1997) but only 17
leafhoppers and 4 planthoppers range from boreal grasslands into the PNW (Faunal Synopsis
and Index). Their dispersal is stochastic; patterns are not discernible. They can be expected
to occur wherever cool-season grasses are found.

Eight Delphacids and 50 leafhoppers are found in PNW grasslands southwards into CA
and low elevations of AZ (Faunal Synopsis and Index). Some of these, notably Orocastus
tener (Beamer & Tuthill) and Psammotettix latipex (DeLong & Davidson), range far into
northern Canada (Hamilton 1997). These species most likely found a refugium in the
southwest because north-south geographic barriers are minimal and summer winds bring flying
insects such as migratory leafhoppers northwards from the Great Basin into western MT and
southern BC (Carter 1927). All such planthoppers and all but 16 (65%) such leafhoppers have
been found in Canada (Maw et al., 2000). By contrast, only one of four flightless southern
leafhoppers migrated north to Canada. One of these, Lystridea uhleri (Baker), was able to
cross the Columbia River only recently, almost certainly by human assistance (Hamilton and

38 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Zack 1999).

More than 50 prairie species are widespread across the continental divide. There is no clear
evidence for the direction of this spread, either onto the prairies from a montane refugium, or
into intermontane grasslands from a prairie refuge. This fauna includes the only grassland
Caliscelid in the PNW, Bruchomorpha beameri Doering, seven Delphacids, and 46
leafhoppers. Of these, 31 taxa range only a short distance into PNW grasslands. A comparison
of their distributions may serve to indicate where the continental divide has been crossed
recently and whether the territory available for colonization has been filled. Their dispersal
routes are inferred on the assumption that the most direct routes are the ones that were actually
taken.

This leaves 120 species and subspecies that are endemic to PNW grasslands plus 22
species which are endemic to montane grasslands adjacent to the PNW and are included for
comparison. These together constitute 142 taxa endemic to grasslands of the PNW and
adjacent mountains. Only six of these (Table 1, designated “?’’) are not known in sufficient
detail to permit analysis.

The 136 well studied endemic taxa (114 from the PNW plus 22 from adjacent mountains)
are compared to 25 prairie leafhoppers and six planthoppers that have limited distribution in
the PNW. Their probable refugia and dispersal routes are discussed in three main faunistic
sections.

First, the endemic taxa are divided into 130 ancient taxa and six subspecies which appear
to be of postglacial origin. All six, belonging to the genus Errhomus (Hamilton and Zack
1999), either form local “swarms” of intermediate forms between adjacent, closely related
species, or (in E. truncus Oman and E. similis medialis Oman) have unusual genitalic
characters that appear to serve as distinguishing specific markers enhancing species barriers
(“character displacement”) where their ranges came in contact.

Next, 20 of the 130 taxa are considered to be glacial in origin since they have sibling taxa
in adjacent areas, either separated by the continental divide, or occurring in isolated areas of
the PNW. Taxa without close relatives, or ones whose ranges overlap, are considered to be
preglacial in origin.

Finally, data from these 130 endemic taxa are synthesized for probable ecological
distributions during the glacial period. They may or may not cross the continental divide, but
have not invaded the prairies to any noticeable extent. Some are merely montane segregates
of species that once were transcontinental; they now appear as pairs of sister taxa across the
continental divide. Such species may have weathered Pleistocene glaciation in southerly
locations, in which case their present distribution may show how they arrived where they now
live. Other endemics represent pairs of sister taxa within the PNW, or as isolated species in
small areas of the PNW that have no close relatives elsewhere. It seems reasonable to assume
that these represent a fauna that survived the Pleistocene in intermontane valley refugia within,
or close to, PNW grasslands. The location of these refugia may be traced most easily where
modern distribution patterns are circular or elliptical, centred around a particular valley. Where
this is not the case, it is assumed that the species has migrated northwards and/or to higher
elevations in postglacial times, retreating from desertification to which it is not adapted.

DISCUSSION

Three species that are characteristic of true prairies are aggressive dispersers, occurring in
most suitable habitats: Auridius auratus (Gillette & Baker) (Hamilton 19995, map 1),
Paraphlepsius lascivius (Ball) (Hamilton 1975, map 9), and the planthopper Laccocera
lineata Scudder (from the Fraser River of central BC southeast in PNW grasslands to the
mountains of central UT, eastward to Saskatchewan, and found on the MT/ID border near

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 39

Rogers and “Grassy” passes, and in Bannock and Monida passes). Three species of
Hebecephalus are also widespread (Hamilton 1998a), but their distribution in PNW grasslands
is more erratic. All these species seem to be generalists on grasses. One other species, Hardya
dentata (Osborn & Ball) from the prairies west to UT (being found just north of “Grassy Pass”
and in Lemhi and Bannoch passes but not in the Snake River valley of ID), also has a “spotty”
distribution in south-central BC and adjacent WA, and in OR east of the Cascade mountains.
Still more disjunct are the known sites for two grass-feeding specialists of the leafhopper genus
Flexamia DeLong (Whitcomb and Hicks 1988, figs. 30, 41). Since two-thirds of these species
show disjunct distributions, the probability is great that some or all of these may represent the
descendants of PNW relict populations of pre-glacial widespread ranges which have spread
by opportunistic dispersal to other favourable sites.

Numerous species of the PNW appear to be endemic to the region. For example, of the 27
nearctic species of leafhoppers belonging to the genus Hebecephalus, 21 are endemic to the
PNW and adjacent areas (Hamilton 1998a); and of the 47 taxa of the flightless leafhopper
genus Errhomus, all but two are found exclusively in the PNW (Hamilton and Zack 1999).
Only a few leafhoppers are widely dispersed throughout the PNW (Fig. 5).

The 140 PNW-endemic taxa analysed below show two major influences: (1) mountain
chains, particularly the continental divide, which separate prairie-inhabiting species from those
in PNW grasslands, and (2) glacial-era grassland refugia, which predetermined the subsequent
dispersal and modern range of many PNW-endemic leafhoppers and planthoppers. The latter
can be subdivided into refugia within the PNW, and those to the south of the PNW which may
have contributed to the modern PNW fauna.

The most significant barrier between the eastern and western grasslands is the continental
divide, which permits only limited faunal exchanges (Fig. 6, stars). A second barrier is formed
by the Bitterroot Mountains along the northern half of the boundary between ID and MT.
Although the latter is pierced by the Clark Fork River, this valley lies deep in the coniferous
forest and is impassable to grassland flora and fauna. The only significant passes along this
ridge north of the continental divide are at Lookout Pass (1400 m elevation), also deep in
coniferous forest near Wallace, ID, and Lost Trail Pass (Fig. 6, F). An additional 21 main
passes penetrate the continental divide between northern BC and the mountains of CO (Fig.
6, circles). Two of these passes do not have any official name. The one near Summit Lake on
the Crooked River north of Prince George, BC is here referred to as “Crooked River Pass”
to distinguish it from the better-known Summit Lake Pass in northern BC. The second,
between Mt. Haggin and Grassy Mt. just south of Anaconda, MT is here called ‘Grassy Pass.”
The following faunal analysis details the evidence for 21 northern prairie leafhoppers and 3
planthoppers crossing the continental divide by way of these passes.

Species that penetrate the continental divide in CO or further southward, such as
Sorhoanus orientalis (DeLong & Davidson), do not occur in PNW grasslands and are not
included in this analysis.

Of the many species that are endemic to PNW grasslands, 14 pairs of sister taxa (16
leafhoppers, 10 planthoppers) occur on opposite sides of the continental divide. These sister-
taxa appear to be of most recent origin, and might possibly be kept geographically separate by
interspecific competition. Six taxa appear to be subspecies of postglacial origin. Another 28
species are widespread in the PNW but lack close relatives and are probably derived from
preglacial stock; six of these occur as three pairs of sister-taxa in disjunct areas of the PNW.
These PNW endemic faunas tend to have range extensions southeast into the mountains of CO,
UT and WY. In addition, there are 22 endemic species that only occur in these areas marginal
to the PNW: 12 in the mountains of CO and adjacent eastern WY (with two ranging north to
southern MT), eight in the mountains of UT and adjacent western WY, and one (Latalus
histrionicus Beirne) in the highest plateau of eastern AZ and also on Vancouver Island in BC.

40 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

The remaining 56 endemic species are limited to isolated parts of the PNW: 15 each in inland
OR and in inland WA, 10 along the ID/MT border, six along the west coast of OR and WA,
six in BC, and four in western MT, restricted to the upper reaches of the Clark Fork River and
its tributaries (the basin of glacial Lake Missoula).

Species crossing the continental divide.
(A) CALISCELIDAE

1. Bruchomorpha beameri Doering 1s a planthopper of the northern prairies (records from
southern Arizona and the Sacramento Valley of California [Doering 1940] must surely refer
to other species) that is also found in many of the intermontane valleys of BC (Fraser,
Okanagan and Kootenay rivers), northeastern WA (on the Pend Oreille River), and the Lake
Missoula basin of MT. It seems to have been able to surmount both “Crooked River” and
Crowsnest passes on the Alberta border, and has been taken in Rogers Pass, MT.

(B) CICADELLIDAE

2. Amblysellus wyomus Kramer is a leafhopper of the western prairies, a specialist on June
grass, Koeleria macrantha (Ledeb.) Schultes [=K. cristata (L.) Pers.], that has been taken in
the East Kootenay of southeastern BC and (Kramer 1971a) in the Bitterroot Valley of western
MT. It has probably reached these sites across the continental divide by way of Crowsnest and
Chief Joseph passes.

3. Athysanella attenuata Baker is a prairie leafhopper that feeds on wheatgrasses,
Agropyron spp. It has been taken in both Lemhi and Bannock passes as well just north of
“Grassy Pass” in MT (Fig. 7) and west of the Great Divide Basin in WY (Fig. 6 E, G, H, K).
From these locations it has spread further west along both sides of the Snake River valley of
ID.

4. Athysanella obesa Ball & Beamer is a grass-feeding prairie leafhopper that specializes
on June grass. It is found east of the continental divide at Badger Pass (2000m elevation) west
of Dillon, MT, and on the other side of the divide in the upper Snake River valley of eastern
ID, on the Lost River Range due south of Bannock Pass (Fig. 6, H), and also in MT due west
of “Grassy Pass” (Fig. 6, E) in Rock Creek valley.

5. Athysanella robusta Baker has the same range and grass host as A. obesa, but has not
been taken in the upper Snake River valley. Its migration route was probably the same as that
of A. obesa; neither is likely to have migrated from ID back into MT by way of Lost Trail Pass
as neither species has been found in the Bitterroot valley.

6. Athysanella terebrans (Gillette & Baker) is a leafhopper of the prairies, a specialist on
sand grass, Calamovilfa longifolia (Hook.) Schribn. which may have come to the upper Snake
River drainage basin in eastern ID by way of Monida Pass. Four females have been taken at
two sites.

7. Auridius helvus (DeLong), a June grass-feeding leafhopper of the western prairies and
eastern “prairie peninsula” from Minnesota to Michigan, has invaded southeastern BC
(probably through Crowsnest Pass) and the valleys of western MT (Hamilton 19995, map 2)
where it occurs in the headwaters of the Clark Fork River system. Populations have been found
on either side of “Grassy” and MacDonald passes, suggesting multiple invasions across the
continental divide. One population is found in Arizona near the Colorado River, and has
probably come there in the same way as Unoka gillettei Metcalf (#25).

8. Auridius ordinatus (Ball), a grass-feeding leafhopper of the northern prairies ranging
from the mountains of CO to isolated grasslands of Alaska (Hamilton 19995, map 4) has
invaded the upper Fraser valley of central BC by way of “Crooked River Pass.”

9. Ceratagallia arida (Oman) is a polyphagous prairie leafhopper that crosses the

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 41

continental divide in southeastern BC near Crowsnest Pass (Fig. 6, B) and in southern ID,
probably by way of the Great Divide Basin of WY (Hamilton 19985, map 3).

10. Ceratagallia viator Hamilton is a northern prairie species with remote populations in
the northwest and southwest (Hamilton 1998), map 2). It has entered PNW grasslands only
along the upper Fraser River and southeastern BC, probably by way of “Crooked River” and
Crowsnest passes.

11. Ceratagallia vulgaris (Oman) is a polyphagous leafhopper that is widespread in eastern
North America and also has populations on the far western plains including Rogers and
Pipestone passes on the continental divide in MT (Hamilton 1998), map 4).

12. Flexamia decora Beamer & Tuthill is a specialist on mat muhly, Muhlenbergia
richardsonis (Trin.) Rydb. It is widespread on the prairies and has penetrated the continental
divide at several places (Whitcomb and Hicks 1988, fig. 39). One population found in the
Clark Fork valley, MT, probably came by way of Chief Joseph Pass, since other MT
populations occur southeast of this pass. One population in Sublette Co., western WY,
suggests that other populations in UT came there by surmounting the Great Basin divide.

13. Diplocolenus c. configuratus (Uhler) is the typical subspecies of a grass-feeding
leafhopper of eastern North America that has crossed the continental divide on the ID/MT
border at MacDonald Pass, probably also at Rogers and “Grassy” passes, and also at Bannock
Pass (Fig. 8, open circles). Its tolerance of boreal conditions makes it the only prairie-
inhabiting leafhopper that is known to have crossed Kicking Horse Pass between Banff and
Yoho National Parks, having been found only 35 km west of the pass in BC. In more southerly
passes it hybridizes with other subspecies (see # 83, 139 and Fig. 8, bull’s-eyes).

14. Elymana circius Hamilton is a grass-feeding leafhopper of the Canadian prairies that
also occurs in the Peace River district of AB and the upper Fraser River of central BC
(Chiykowski and Hamilton 1985, fig. 8), probably by way of “Crooked River Pass,” BC (Fig.
6, A).

15. Idiodonus heidemanni (Ball) is a leafhopper of the western plains as far north as MT
that is also known from the upper Snake River in WY. It probably came there by way of the
Great Divide Basin of WY. It is not the species recorded as “/diodonus heidemanni” from CA
(DeLong and Severin 1948) which is actually Bonneyana schwarizi (Ball).

16. Mesamia ludovicia Ball is a specialist on prairie sage, Artemisia gnaphalodes Nutt.
(=A. ludoviciana Nutt.) that has been taken at Rogers Pass, MT.

17. Mocuellus caprillus Ross & Hamilton is a specialist on western wheatgrass, Agropyron
smithii Rydb., that occurs throughout the grasslands of MT and WY, and southeast of the
mountains of UT (Fig. 9). It has crossed the continental divide at least twice, probably at
MacDonald Pass and Great Divide Basin, judging by the proximity to known sites for this
species. It is also found at Lemhi Pass on the ID/MT border, where it hybridizes with
Mocuellus caprillus anfractus (#88).

18. Norvellina clarivida (Van Duzee) is a specialist of Atriplex spp. in southern AB and
SK, south in the mountains of CO and NM, and also occurs in UT (Lindsay 1940), having
crossed Great Divide Basin at Creston, WY.

19. Orocastus labeculus (DeLong) is a northern prairie leafhopper feeding on spear grasses
(Stipa spp.) It also has been found in southeastern BC (Cranbrook, Skookumchuck), eastern
ID (Carmen) and the Unitas of northern UT (Duchesne) and therefore must have penetrated
the continental divide in at least three places: Crowsnest, Lemhi and Great Divide Basin passes
seem indicated. A short series containing only females and nymphs from south-central BC
(Hedley) may prove to be an undescribed sister-species when males are found.

20. Orocastus perpusillus (Ball & DeLong) is an abundant northern prairie leafhopper
feeding on spear grasses. It has penetrated the continental divide at two locations (Fig. 10): in
the Lake Missoula valley and in the upper reaches of the Snake River. The most likely passes

42 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

it could have come through, considering its known range, are Rogers and Monida passes (Fig.
6, C, J). Its presence in western WY suggests that it has also crossed the Great Divide Basin.

21. Pinumius sexmaculatus (Gillette & Baker) is a grass-feeding leafhopper of the
northwestern plains that also occurs along the Fraser River of central BC, just north of “Grassy
Pass,” MT, and on the foothills of the Unita Mountains of UT. The northern and southernmost
populations west of the continental divide probably came there by way of “Crooked River
Pass,” BC and the Great Divide Basin of WY.

22. Prairiana kansana (Ball) is a prairie leafhopper that probably feeds on some woody
plant. It has been found just west of Rogers Pass and just north of “Grassy Pass” in MT, in
Bannock Pass and south of Lost Trail Pass in ID. As this species is not known from the
Bitterroot valley of MT, it probably has approached Lost Trail Pass from the south, via
Bannock Pass.

23. Psammotettix totalus (DeLong & Davidson) is a grass-feeding leafhopper of the
northwestern plains from northern MT to WY that also occurs in the Snake River valley of
south-central ID (DeLong and Davidson 1935), probably by way of the Great Divide Basin
of WY (DeLong and Davidson 1935: “Cane Tree” and “Cattail Spring,” WY are unknown
localities).

24. Rosenus cruciatus (Osborn & Ball) is a leafhopper specialist on June grass that is
widespread on the northern prairies including western MT. Populations throughout the Lake
Missoula Basin and south of Monida Pass in ID probably are derived from the only one found
to the east, on the upper reaches of the Missouri River. This would imply that the Lake
Missoula Basin populations came through one or more of the many passes near Butte. From
the dispersal patterns of other species, Pipestone Pass seems the most likely route. This
leafhopper also occurs on the foothills of the Unita Mountains of UT, and in southern BC.
Whether the southern populations arrived in UT from the Snake River plain of ID, or across
the Great Divide Basin of WY, cannot be ascertained at present. The population in the East
Kootenay of eastern BC must have come there through Crowsnest Pass. A female (incorrectly
recorded as Rosenus obliquus by Maw et al. [2000]) taken at Kelowna, BC and a male from
Myncaster (near the WA border) show that an isolated population also inhabits the Okanagan
Valley. The source for this population is unknown; possibly it is a Hypsithermal relict.

25. Unoka gillettei Metcalf is a leafhopper specializing on sand dropseed, Sporobolus
cryptandrus (Torr.) A. Gray, and ranging from southern Alberta and southwestern MT east to
Minnesota and south to Colorado. It is also found on the same host in southwestern WY, UT
and adjacent AZ and NV, and thus appears to have crossed the Great Divide Basin of WY
(Fig. 6, K) and travelled down the Green River to the Grand Canyon (Fig. 6, stars).
Populations in southern AZ are probably derived from a more southerly pass, as this species
specializes on gyp dropseed, Sporobolus nealeyii Vasey from TX to AZ.

(C) DELPHACIDAE

26. Elachodelphax pedaforma (Beamer) is a planthopper of the northern plains that also
occurs just beyond Crowsnest Pass at Fernie, BC.

27. Eurybregma magnifrons (Crawford) is a prairie planthopper that has invaded the upper
reaches of the Fraser River in central BC via “Crooked River Pass,” and migrated as far north
as Alaska, but has not crossed more southerly passes (Fig. 11).

28. Laccocera canadensis Beirne is a planthopper of the northern plains that also occurs
in the East Kootenay valley, BC, and in MT along Rock Creek and the Blackfoot River in
Lake Missoula Basin, and on “Grassy Pass.” The population in eastern BC must have come
there through Crowsnest Pass, while that on Rock Creek came by way of “Grassy Pass” and
the lower pass at Silver Lake just to the west. The population on the Blackfoot River probably
came there by way of Rogers Pass.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 a

29. Laccocera flava Crawford is a northern prairie planthopper that has entered the
headwaters of the Snake River of ID (Fig. 12, filled circles) by way of UT, probably traversing
the Great Divide Basin of WY, and is also in the Lake Missoula valley of MT. Whether it
came to the latter by way of MacDonald or by “Grassy Pass” cannot be determined. Its
presence in both Bitterroot and Rock Creek valleys suggests that it was once widespread in the
headwaters of the Clark Fork River. Its range may have become fragmented during the
formation of glacial Lake Missoula. It also reaches the continental divide at Bannock Pass.

30. Laccocera vittipennis Van Duzee is a common polyphagous planthopper of northern
prairies and Peace River grasslands on the western half of the continent, although not confined
to grasslands in eastern North America. It has been found in BC just beyond “Crooked River”
and Yellowhead passes, in MacDonald Pass and west of the pass in the upper reaches of the
Lake Missoula valley (Fig. 13).

31. Nothodelphax foveatus (Van Duzee) is a common planthopper of northern prairie
including the Peace River district. It also has been found in ID (the Lemhi Valley and in the
upper Snake River plain), MT (Blackfoot River Valley), OR (shore of the Columbia River)
and UT (foothills of Unita Mountains). Possibly it crossed the continental divide by way of
Rogers, Lemhi and Great Divide Basin passes; from the Snake River plain it has invaded the
Columbia River near its junction with the Snake.

Pairs of sister taxa separated by the continental divide.
(A) CICADELLIDAE

32. Athysanella occidentalis megacauda Hamilton (see Part 1) is a grass-feeding prairie
leafhopper known only from southern interior of BC and northern WA, south along the
Columbia as far as Vantage, WA. It was probably restricted to the Columbia canyon during
the Pleistocene. Typical occidentalis occurs only east of the continental divide. A short
distance to the west of the continental divide from the Bitterroot Valley of MT, south through
the headwaters of the Salmon River and the Camas valley in ID occur a hybrid swarm between
these two subspecies. To attain this distribution these hybrids must have crossed Lost Trail
Pass and the still higher Galena Summit (2700m elevation) between the headwaters of the
Salmon and Camas rivers. How typical occidentalis came across the continental divide is not
known, but the most likely route is through Lemhi Pass (Fig. 6, G) which lies halfway between
the extremes of its distribution to the west of the divide.

33. Attenuipyga (Dorycara) minor subspecies setosa Oman is a grass-feeding leafhopper
known from widely scattered PNW grasslands of central OR and adjacent WA, southern BC,
and the upper Snake River plain in ID. Its related subspecies, typical A. minor (Osborn), is
entirely east of the continental divide. Individuals that appear to be hybrids between these
subspecies have been taken just north of “Grassy Pass” in MT, and (Oman 1985) on the upper
Snake River plain not far from Monida Pass, in the mountains of CO and northern UT, and
central NV.

34. Auridius ordinatus subspecies crocatus Hamilton, a grass-feeding leafhopper, occurs
throughout PNW grasslands and as far south as northern NV (Hamilton 19994, map 4). It was
probably widespread in PNW grasslands during the Pleistocene. It has two related
geographical subspecies: typical A. ordinatus Ball (#8), a northern prairie subspecies, and
subspecies amarillo Hamilton in the mountains of NM. Hybrids between the typical subspecies
and crocatus are found at Monida Pass and adjacent areas of the upper Snake River valley.

35. Ceratagallia nanella subspecies zacki Hamilton, a polyphagous leafhopper in WA and
southernmost BC (Hamilton 19985, map 9), is related to two widely disjunct geographical
subspecies: typical C. nanella Oman of the prairies and southwestern grasslands from Canada
to AZ, and subspecies australis Hamilton in the mountains of Mexico. Subspecies zacki was
probably widespread on the Columbia basin during the Pleistocene.

at J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

36. Ceratagallia siccifolia subspecies compressa Hamilton, a polyphagous leafhopper
widespread in PNW grasslands, is related to two geographical subspecies: the widely disjunct
subspecies alaskana Hamilton in the far northwest, and typical C. siccifolia (Uhler) which is
mainly a prairie subspecies. Because the typical subspecies is widespread and aggressive,
having been taken at Bannock and Lemhi passes, and up to 3300 m elevation in CO, it is
assumed here to have been able to traverse all the mountain passes. It appears to have invaded
the PNW before postglacial times. Its populations have completely swamped the PNW
subspecies at higher elevations. Subspecies compressa is now confined mostly to valley
bottoms (Hamilton 19984, map 10).

37. Psammotettix beirnei Greene is a grass-feeding leafhopper known only from 7600'
[2500 m elevation] on Mt. Harry in the Selkirk Mountains near Revelstoke, BC, erroneously
recorded from Northwest Territories (Greene 1971). Its sister-species, P. alexanderi Greene
is known only from the White Mountains of New Hampshire (Greene 1971).

38. Stenometopiellus vader Hamilton (see Part 1) has a sister species S. cookei (Gillette)
on the northern prairies and mountains of CO. Both are sedge-feeding leafhoppers, the former
at present known only from the Lost River Range of eastern ID and just north of “Grassy Pass”
in MT. Its glacial-age refugium was probably the upper Snake River valley. Whether it spread
to southern MT by way of Lemhi or Bannock pass and later entered the Lake Missoula basin
by way of “Grassy Pass”, or whether it invaded the Lake Missoula basin directly by way of
Lost Trail Pass, cannot be determined at present.

39. Unoka dramatica Hamilton (see Part 1) is known only from southern BC (Fig. 6, filled
circle) widely disjunct from its sister species U. gillettei (#25). This PNW endemic of sandy
areas probably survived the Pleistocene in the upper reaches of the Columbia basin on sandy
outwashes from terminal moraines.

(B) DELPHACIDAE

40. Elachodelphax mazama Hamilton (see Part 1) is known only from the Methow Valley
in northwestern WA. Its presence near the arid inland valleys of southern BC suggest that its
parent species was once widespread throughout central BC. Its sister species is FE. borealis
Hamilton (see Part 1), a planthopper widespread from coastal Labrador and high mountains
of New Hampshire west to the Peace River district of Alberta that is not known to have
penetrated the Rocky Mountains.

41. Eurybregma eurytion Hamilton (see Part 1) includes in its extensive PNW range the
Lake Missoula valley in MT, indicating that it must have surmounted Lost Trail Pass, as it has
Bannock Pass (Fig. 11). It was probably widespread in PNW grasslands during the
Pleistocene. Its sister-species, the prairie-inhabiting E. magnifrons (#27) is found north of the
range of EF. eurytion, except where they co-occur at Princeton, BC.

42. Laccocera oregonensis Penner has a disjunct distribution, in eastern ID and around the
Columbia basin from southern BC to OR (Fig. 12, open circles). This suggests that it was
probably widespread in PNW grasslands during the Pleistocene. Its sister species, L. flava
(#29), may be inhibited in its westward spread by L. oregonensis just west of Lemhi Pass.

43. Laccocera vanduzeei Penner, sister species to Laccocera vittipennis (#30), 1s
widespread from central BC to ID, and has many populations in the mountains of AZ, CO and
UT (Fig. 13, open circles) where it probably found glacial refugia. It occupies the southern
part of the Lake Missoula valley as well as the headwaters of the Missouri River in MT. It has
been found in “Grassy”, Lemhi, Bannock and Monida passes and adjacent to Lost Trail Pass.
Multiple crossings of the continental divide seem indicated.

44. Nothodelphax venustus (Beamer), sister species to NV. foveatus (#31), is confined to the
mountains of southern BC and adjacent WA, southern WY and adjacent CO, and northern AZ.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 45

Endemic pairs of sister taxa in PNW (CICADELLIDAE only).

45. Athysanella expulsa Blocker is a grass-feeding leafhopper known only from central OR
east of the Cascade Mts. (Blocker and Johnson 1990).

46. Athysanella repulsa Hamilton (see Part 1), sister-species to 4. expulsa, is known only
from Flint Creek on the upper reaches of the Clark Fork valley of MT.

47. Auridius cosmeticus Hamilton (Hamilton 19996, map 3) is a sedge-feeding leafhopper
found only in the upper reaches of the Blackfoot and Bitterroot valleys of MT, where it seems
to have found a refugium from the waters of glacial Lake Missoula.

48. Auridius vitellinus Hamilton, sister species to A. cosmeticus (Hamilton 19995), is
known only from southernmost OR east of the Cascade Mts.

49. Latalus histrionicus Beirne is known only from Vancouver Island, BC, and the high
plateau of eastern AZ. Its range appears to reflect two isolated glacial-age refugia rather than
competition with its sister-species (see #50), because both species can be found near Alpine,
AZ.

50. Latalus intermedius Ross & Hamilton is a common grass-feeding leafhopper of
mountains in eastern AZ, CO, UT and western WY (which were probably its glacial-age
refugia). It has also established colonies in southeastern ID near the UT border, and at Rogers
Pass, MT, and three sites in the north: Peace River district of AB and BC, on the upper Fraser
River of central BC, and at Norman Wells on the Mackenzie River, Northwest Territories. It
must have spread from central BC through “Crooked River Pass” to reach the Peace and
Mackenzie rivers, and across the Great Divide Basin to reach UT and ID.

51. Sorhoanus debilis (Uhler) is a grass-feeding leafhopper from the eastern foothills of
the Cascade Mountains from northern OR to intermontane valleys of southern BC and ID, and
the mountains of central UT, eastern WY and CO (Fig. 14), all of which may have provided
glacial refugia. It has crossed the Crowsnest Pass to Watertown, AB, and has invaded the
headwaters of the Missouri River in MT by way of Bannock Pass and possibly also Lemhi and
Monida passes. From the vicinity of Helena it may have crossed MacDonald Pass to establish
a colony on the headwaters of the Clark Fork River; possibly it also crossed “Grassy Pass’’.

52. Sorhoanus virilis Hamilton (see Part 1), sister species to S. debilis, is known only from
the Cascade Mountains of southernmost OR.

Endemics without close relatives, widespread in PNW.
(A) CICADELLIDAE

53. Attenuipyga (Dorycara) omanae (Beamer) is a leafhopper specialist on bluebunch
fescue, Festuca idahoensis Elmer in widely scattered PNW grasslands from southern BC south
to the highlands of central OR, east to Bannock Pass (Fig. 6, H) and just north of “Grassy
Pass.” A nymph, possibly belonging to this species, has also been taken at Missoula, MT. It’
was probably widespread in WA-ID grasslands during the Pleistocene.

54. Auridius safra Hamilton is a grass-feeding leafhopper endemic to the hill country of
central and eastern OR and central ID. Its single site in the mountains of northeastern CA
(Hamilton 19995, map 1) probably represents the area of its glacial refugium.

55. Ceratagallia gallus Hamilton is a polyphagous leafhopper endemic to sagebrush-
grassland ecotone in southeastern ID and central UT. It probably survived the glacial period
and accompanying inundation of central UT on arid hillsides of the Wasatch Plateau, as has
the flightless, endemic sagebrush-feeding leafhopper Errhomus naomi (#155).

56. Chlorotettix similis DeLong is a grass-feeding leafhopper of the oak savannah of BC
and OR, and of Palouse grasslands from southern BC to western ID. It was probably
widespread in WA grasslands during the Pleistocene.

57. Endria montana (DeLong & Sleesman) is a leafhopper of the Columbia basin (Wolfe
1955) and high elevation grasslands and also occurs on oak savannah in Vancouver Island, yet

46 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

it has penetrated only a short distance into Canada (Fig. 15), probably by way of the Columbia-
Kootenay valley system. It apparently feeds on Letterman needlegrass, Stipa lettermanii
Vasey, and has been taken at Rogers, MacDonald and Monida passes, from whence it has
been able to invade the upper reaches of the Missouri and Yellowstone rivers. Presumably it
came to the Lake Missoula valley by way of Lost Trail Pass. It has also been taken at Lolo
Pass (1800 m elevation) in the Bitterroot Range and just beyond this pass on the upper reaches
of the Lochsa River, a tributary of the Clearwater. Oddly, it is not known from mountains south
of ID. Its glacial refugium was probably the Snake River valley of ID and MT, from whence
it retreated as postglacial temperatures rose.

58. Errhomus lineatus (Baker) is a flightless leafhopper specialist on balsamroot,
Balsamorhiza spp., endemic to Palouse grasslands of eastern WA, northeastern OR and
western ID (Hamilton and Zack 1999, map 6). Its typical subspecies is a hybrid of 3 peripheral
subspecies (see #100-102), suggesting that there were at least 3 glacial refugia. Since the
parental morphs are no longer adjacent, it is impossible to tell whether this hybrid swarm
originated before the latest glacial period and survived on the Palouse hills of WA, or whether
it represents a highly successful postglacial hybrid that has completely swamped the
intervening parental populations.

59. Errhomus similis kahlotus Oman is a subspecies of a widespread but flightless
balsamroot-feeding leafhopper species endemic to the Columbia basin in both WA and OR.
Populations of subspecies kahlotus Oman and its probable hybrid dubiosus Oman are
widespread but sparse on the Columbia basin of WA. They must have survived the glacial
period across most of the Columbia basin, except where intermittent Pleistocene inundations
swept away populations in low-lying areas. They represent 2 of 12 subspecies of E. similis (see
also # 106, 123-126) that reflect subdivision of the basin and eastern foothills of the Cascade
Mountains of WA and OR due to canyons formed by the glacial-age rerouted Columbia River
(Hamilton and Zack 1999, map 7).

60. Hebecephalus caecus Beamer is a grass-feeding leafhopper known only from the
eastern foothills of the Cascade Range in OR (Beamer 1936) and from the mountains of
southern ID just northeast of Boise. It was probably once widespread in OR-ID grasslands, but
during the Pleistocene found separate refugia on the eastern slopes of the Cascades in OR and
on the Snake River plains of ID.

61. Hebecephalus callidus (Ball) is a grass-feeding leafhopper known from the Palouse
hills of WA (Ball 1899a), the East Kootenay valley of BC, the mountains of southern ID north
of the Snake River plains, and the Bitterroot River valley of MT. It probably weathered the
Pleistocene in the mountains of northern WY, and must have come to the Bitterroot valley of
MT by way of Lost Trail Pass.

62. Hebecephalus crassus (DeLong) is a grass-feeding leafhopper known from
Yellowstone Park, WY (DeLong 1926), from the Lost River Range in southeastern ID, and
from Mt. Baldy (2000m elevation) in south-central BC. Its glacial-age refugium was probably
in the mountains of northern WY.

63. Hebecephalus firmus Beamer has been taken in WA (Beamer 1936) and I have taken
it near Yellowstone Park on both sides of the MT-WY border. Its glacial-age refugium was
probably in the mountains of WY.

64. Hebecephalus hilaris Beamer is a grass-feeding leafhopper known only from the
mountains of southeastern WY (Beamer 1936), and at Stevens Pass (1200m elevation) in the
Cascade Range of WA on redtop (Agrostis gigantea Roth.), an introduced grass.

65. Hebecephalus sagittatus Beamer & Tuthill has been found in small numbers in
scattered localities throughout the grasslands of the PNW, from the Blue Mountains of OR
(Beamer and Tuthill 1935) east to the Lost River Range of eastern ID, and from the foothills
of the Unita Mountains of UT north to the Okanagan Valley of BC. The Unita Mountains may

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 ull

have been its glacial-age refugium. Specimens have also been taken on south-facing slopes of
the Aishihik Canyon in southwestern Yukon, showing the close relationship between the
grasslands of these very widely separated areas. Its spread to the Yukon may well have been
accomplished before the present interglacial era (Hamilton 1997).

66. Laevicephalus salarius Knull is a specialist on alkali grass, Distichlis stricta (Torr.)
Rydb. It has been found most commonly in UT on the Wasatch Plateau and in the north, but
has also been taken at Kamloops in southern BC, and just north of Denver, CO. Its distribution
probably reflects lack of collecting on this grass at intervening localities.

67. Latalus curtus Beamer & Tuthill is a grass-feeding leafhopper widespread in PNW
grasslands that also has been taken at Pipestone Pass, at Lolo Pass on the MT/ID border west
of Missoula, and on the upper Fraser River in central BC.

68. Latalus mundus Beamer & Tuthill is a grass-feeding leafhopper of PNW grasslands
from the Okanagan Valley of southern BC to the Grand Teton Mountains of WY (which may
have been its glacial-age refugium). It is also found in southeastern BC not far from Crowsnest
Pass (probably by way of the Columbia-Kootenay valley system), at MacDonald Pass, MT,
and in the Bitterroot Valley, MT, below Lost Trail Pass.

69. Mocuellus larrimeri (DeLong) is a grass-feeding leafhopper of PNW grasslands from
scattered localities in south-central BC to western ID, and is also known from the East
Kootenay of BC (probably by way of the Columbia-Kootenay valley system), Missoula, MT
(DeLong 1926), and Rattlesnake Ridge in southern WA. It was probably widespread in WA-
ID grasslands during the Pleistocene but has moved northwards since then. Presumably it came
to MT by way of Lost Trail Pass.

70. Norvellina rubida (Ball) is a leafhopper that feeds on fleabane, Eriogonum spp.
(Asteraceae), which can grow at high elevation on south-facing mountain slopes. Nevertheless,
it is confined to the zone of PNW grasslands (Fig. 5, filled circles) and may have been
widespread in WA-ID grasslands during the Pleistocene.

71. Norvellina vermiculata Lindsay is found on the Snake River plain, ID and northern UT
to northwestern CO. It has also been taken just over the continental divide at White Sulphur
Springs in western MT.

72. Orocastus pinnipenis Ross & Hamilton is a leafhopper feeding on Sandberg’s blue
grass, Poa secunda Presl. that occurs on west-facing slopes at about the 2000m level
throughout most of ID and western MT (Fig. 16); elsewhere it is a valley or plateau species.
It was probably widespread in ID grasslands during the Pleistocene. It has been taken in
Rogers, Lemhi, Bannock and Monida passes and also close to Lost Trail and MacDonald
passes, but has invaded only the fringe of the prairies, preferring the higher elevation of the
Little Belt mountains of MT and the Bighorns of WY.

73. Psammotettix attenuens (DeLong & Davidson) 1s a grass-feeding leafhopper ranging
from the mountains of CO to coastal BC (Fig. 17). It was probably widespread in the CO
mountains and WA-ID grasslands during the Pleistocene. It has been taken in “Grassy”, Lemhi
and Bannock passes but has not penetrated the continental divide

74. Sorhoanus xiphosura Hamilton (see Part 1) is a grass-feeding leafhopper widespread
through the PNW. It is tolerant of high altitudes (e.g., it is found just beyond Lolo Pass on the
ID side of the border). Evidently it crosses passes readily.

(B) DELPHACIDAE

75. Caenodelphax atridorsum (Beamer) was described from the eastern slopes of the
Cascade Range in OR and has subsequently been discovered at two locations west of the
continental divide in MT (west side of Clark Fork Valley at Missoula, and in the Blackfoot
Valley).

76. Pissonotus rubrilatus Morgan & Beamer was described from Colorado and has

48 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

subsequently been found in widely scattered localities in WY, ID, and central BC (Bartlett and
Deitz 2000).

Taxa endemic to AB and inland BC.
(A) CICADELLIDAE

77. Ceratagallia okanagana Hamilton is known only from southernmost Okanagan Valley
in BC (Hamilton 1998)). It may have been limited to the northern part of the Columbia basin
during the Pleistocene.

78. Cuerna cuesta Hamilton. Although not strictly confined to grasslands (Fig. 18), this
forb-feeding leafhopper of BC is most often associated with grassy areas including lodgepole
pine stands. Its range extends into eastern WA and northern MT. It may have been limited to
the northern part of the Columbia basin during the Pleistocene. Its sister species C.
septentrionalis, which also ranges northwards into coniferous forest glades, is entirely on other
side of divide from BC to CO, crossing into the Pacific watershed only in the Yukon (Hamilton
1997).

79. Hebecephalus planaria Hamilton (1998a) is a grass-feeding leafhopper known only
from southern interior of BC. It may have been limited to the northern part of the Columbia
basin during the Pleistocene.

80. Rosenus decurvatus Hamilton & Ross is a grass-feeding leafhopper known only from
south-facing bluffs above the Peace River near the BC/AB border. It probably found a glacial-
age refugium on south-facing hillsides in MT and followed the retreating ice front northwards
during deglaciation.

(B) DELPHACIDAE

81. Paraliburnia furcata Hamilton (see Part 1) is a planthopper known only from the upper
Fraser River of central BC.

82. Paraliburnia lecartus Hamilton (see Part 1) is a planthopper known only from the
Peace River district of BC. Like the preceding species (#80, 81), it probably found a glacial-
age refugium on south-facing hillsides in MT.

Taxa endemic to ID/MT border (CICADELLIDAE only).

83. Athysanella castor Hamilton (see Part 1) is a leafhopper found on both sides of the
ID/MT border and also across the continental divide in the Rock Creek valley of MT. It has
been found just west of the divide at Lemhi Pass. It may have found a glacial-age refugium on
the MT foothills to the east of the pass.

84. Athysanella nielsoni Blocker is a leafhopper known only from the upper Snake River
valley of ID about 5 km north of Idaho Falls. A short series was taken on “sage” (Blocker and
Johnson 1990), but all other species in this genus are grass feeders, and this one probably is
also. It may never have moved from its glacial-age refugium, although a female that may
belong to this species has been found in western ID near the junction of the Salmon and Snake
rivers.

85. Diplocolenus configuratus bicolor Hamilton (see Part 1) has quite a restricted range
although technically this leafhopper occurs in three states (Fig. 8, stars). It is confined to the
continental divide and adjacent mountain ridges extending along only 400 km. During the
Pleistocene it may have been restricted to the upper end of the Snake River valley. Its range
now abuts that of the common eastern D. configuratus s.s. (#13) where they hybridize (Fig.
8, bull’s-eyes).

86. Hebecephalus crenulatus Hamilton (1998a) is a grass-feeding leafhopper that has been
taken from the upper Snake River plain. It may never have moved from its glacial-age
refugium. Unassociated females from near the Utah border may be conspecific with this

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 ae

species.

87. Hebecephalus ferrumequinum Hamilton (1998a) is a grass-feeding leafhopper that is
known only from just west of Bannock Pass. It may never have moved from its glacial-age
refugium.

88. Hebecephalus picea Hamilton (1998a) is a grass-feeding leafhopper that is known only
from a stony plain in the Lost River valley 30 km north of the Snake River plain. It may never
have moved from its glacial-age refugium.

89. Hebecephalus pugnus Hamilton (1998a) is a grass-feeding leafhopper that is known
from several sites in the upper Snake River valley of ID, and near the headwaters of Lost River
which flows into the Snake River plain. It probably found a glacial-age refugium on the upper
Snake River valley.

90. Hebecephalus veretillum Hamilton (1998a) is a grass-feeding leafhopper that is known
only from a west-facing hillside at Ketcham, ID on the north side of the Snake River plains.
It may never have moved from its glacial-age refugium.

91. Mocuellus caprillus anfractus Hamilton (see Part 1) is a grass-feeding leafhopper with
a nearly circular distribution centred around the upper Snake River valley of ID (Fig. 9, filled
circles) which is probably its glacial-age refugium. It has spread to hills within 200 km N or
S, and 150 km E or W. The typical subspecies (see #17) inhabits the prairies and hybridizes
with this subspecies at Lemhi Pass on the ID border.

92. Rosenus obliquus (DeLong & Davidson) 1s a grass-feeding leafhopper known only
from two valley sites in the mountains of south-central ID. It probably found a glacial-age
refugium on the upper Snake River valley.

Taxa endemic to Lake Missoula basin, MT (CICADELLIDAE only).

93. Errhomus bracatus Hamilton & Zack 1s a flightless balsamroot-feeding leafhopper
found only in the upper reaches of the Bitterroot valley of MT, where it seems to have found
a refugium from the waters of glacial Lake Missoula (Hamilton and Zack 1999, map 9).

94. Errhomus camensis Hamilton & Zack is a flightless balsamroot-feeding leafhopper
found only in the south of the Blackfoot valley of MT, where it seems to have found a
refugium from the waters of glacial Lake Missoula (Hamilton and Zack 1999, map 10).

95. Errhomus rivalis Hamilton & Zack is a flightless, polyphagous leafhopper found only
on the west bank of the Bitterroot River of MT, where it joins the Clark Fork River to the west
of Missoula, MT. (Hamilton and Zack 1999, map 9). Since this site was under 300m of water
only 12,000 years ago, it must have survived on nearby Black Mountain.

96. Errhomus solus Oman is a flightless, polyphagous leafhopper found on hillsides around
Missoula and in the Blackfoot valley of MT (Hamilton and Zack 1999, map 9). As these sites
are separated by the Clark Fork River, it must have found refugia from the waters of glacial
Lake Missoula in at least two localities.

Taxa endemic to inland OR and western ID (CICADELLIDAE only).

97. Ceratagallia acerata Hamilton is known only from southernmost OR east of the
Cascade Mts. (Hamilton 19985).

98. Ceratagallia clino Hamilton is known only from the eastern slopes of the Cascade
range in OR (Hamilton 19985).

99. Ceratagallia lophia Hamilton is known only from the 1500m high summit of the
Jackass Mts. in southeastern OR (Hamilton 1998)).

100-101. Errhomus affinis Oman 1s a flightless balsamroot-feeding leafhopper known only
from the high elevations on either side of Hells Canyon on the OR-ID border (Hamilton and
Zack 1999, map 9). The populations divided by this canyon on the ID side (typical subspecies,
#100) and on the OR side (subspecies attenuatus Hamilton & Zack, #101) must predate the

50 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

formation of the canyon at least 1.5 million years ago.

102. Errhomus josephi Oman is a flightless balsamroot-feeding leafhopper known only
from high elevations in the Grande Ronde canyon of northwestern OR and adjacent WA
(Hamilton and Zack 1999, map 8).

103. Errhomus lineatus cordatus Hamilton is now confined to the north shore of Coeur
d’Alene Lake in ID, and may have survived the glacial period along the adjacent Spokane
River of eastern WA where hybrids with the typical subspecies are now found.

104. Errhomus lineatus idahoensis Oman is now found east of the Snake River in ID and
probably was confined to canyons there during the glacial maximum.

105. Errhomus lineatus umatilla Oman occurs in scattered localities of northeastern OR
and may have been confined to the south banks of the lower Columbia canyon during the
Wisconsinan.

106. Errhomus ochoco Oman is a flightless balsamroot-feeding leafhopper known only
from the 1500m high summit of the Ochoco Mts. in central OR (Hamilton and Zack 1999, map
7).

107. Errhomus pallidus Oman is a flightless potentilla-feeding leafhopper known only
from the foothills of the Wallowa Mts. in northwestern OR (Hamilton and Zack 1999, map
10).

108. Errhomus serratus Oman is a flightless balsamroot-feeding leafhopper endemic to
widely separated PNW grasslands of northeastern OR and the eastern foothills of the Cascade
Mountains of WA (Hamilton and Zack 1999, map 9). It must have ranged around the entire
periphery of the Columbia basin before Pleistocene temperatures eliminated its northernmost
populations, leaving 3 subspecies in areas which might have provided glacial refugia (see also
#116, 117). The typical subspecies is found only on the southern flanks of the Blue Mts. of
OR.

109. Errhomus similis medialis Oman, together with possible hybrids (E. similis minutus
Oman, E. similis nanus Oman) and populations showing character displacement (E. similis
truncus Oman) occurs south of the lower Columbia canyon. All may be postglacial taxa
derived from a single glacial refugium.

110. Errhomus variabilis erratus Hamilton & Zack, is restricted to the Salmon River on
the west (OR) side of the canyon.

111. Errhomus variabilis gracilis Hamilton & Zack is restricted to the Salmon River on
the east (ID) side of the canyon.

112. Errhomus variabilis mimicus Hamilton & Zack is found on the southern flanks of the
Blue Mts. of OR.

113. Errhomus winquatt Oman is a flightless balsamroot-feeding leafhopper known only
from the south bank of the lower Columbia canyon where it penetrates the Cascade Mts. in
southern OR (Hamilton and Zack 1999, map 8).

114. Psammotettix greenei Hamilton (see Part 1) is a grass-feeding leafhopper known only
from southernmost OR east of the Cascade Mts. (Greene 1971).

Taxa endemic to inland WA (CICADELLIDAE only).

115. Ceratagallia vipera Hamilton is known only from the 1000m high summit of
Rattlesnake Ridge, an eastern outlier of the Cascade Mts. in southern WA (Hamilton 19985).

116-117. Errhomus brevis Oman is a flightless balsamroot-feeding leafhopper known only
from the north bank of the lower Columbia canyon and inland regions of southern WA
adjacent to the Cascade Mts. (Hamilton and Zack 1999, map 6). The canyon populations
(typical subspecies, #116) and inland ones (subspecies simcoe Oman, #117) form hybrid
swarms where they meet at the mouth of the Klickitat River (idem, map 11).

118. Errhomus calvus Oman is a flightless polyphagous leafhopper known only from the

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 51

Okanogan highlands of northern WA and adjacent BC (Hamilton and Zack 1999, map 2).

119. Errhomus paradoxus Oman is a flightless balsamroot-feeding leafhopper known only
from the northern bank of the lower Columbia canyon in southern WA just to the east of the
range of Errhomus brevis (Hamilton and Zack 1999, map 8).

120. Errhomus picturatus Hamilton & Zack is a flightless balsamroot-feeding leafhopper
known only from a single site near Wenatchee Lake in northwestern WA adjacent to the
Cascade Mts. just to the east of the range of Errhomus wolfei (Hamilton and Zack 1999, map
8).

121. Errhomus praedictus Hamilton & Zack is a flightless balsamroot-feeding leafhopper
known only from a single site on the south slopes of the Simcoe Mts. in southern WA just to
the east of the range of Errhomus brevis (Hamilton and Zack 1999, map 8, as inconspicuus
[sic]).

122. Errhomus reflexus Oman is a flightless balsamroot-feeding leafhopper known only
from two populations on either side of the Kittitas Valley in central WA east of the Cascade
Mts. (Hamilton and Zack 1999, map 8).

123. Errhomus satus Oman is a flightless balsamroot-feeding leafhopper known only from
just north of Satus Pass in the Simcoe Mts. in southern WA just to the east of the range of
Errhomus brevis (Hamilton and Zack 1999, map 8).

124. Errhomus serratus instabilis Oman is found only in the Naches valley of WA.

125. Errhomus serratus obliteratus Hamilton occurs on the arid eastern slopes of the
Wenatchee Mts. of WA.

126. Errhomus similis Oman, typical subspecies, and its probable hybrids E. similis
confinis Oman and Errhomus similis relativus Oman, occur on the eastern foothills of the
Cascade Mountains on WA.

127. Errhomus similis sobrinus Oman is confined to the angular bend made by the
Columbia in the Pascoe Basin, WA. This appears to be an area that was east of the Columbia
until the river shifted to its present location at least 5 million years ago.

128. E. similis socius Oman 1s confined to an area to the west of the Columbia canyon in
WA south of Lake Chelan and north of the Wenatchee River canyon.

129. Errhomus similis zonarius Oman is found north of the Columbia canyon and east of
the Okanagan valley in WA.

130. Errhomus variabilis Oman is a flightless balsamroot-feeding leafhopper endemic to
the northern edge of the Columbia basin of WA, and the Snake River valley on the ID-OR
boundary (Hamilton and Zack 1999, map 10). Its various subspecies reflect at least 4 glacial-
age refugia, the typical subspecies being found on the Columbia basin and the others (#107-
109) in peripheral areas.

131. Errhomus wolfei Oman is a flightless balsamroot-feeding leafhopper known only from
the Wenatchee River canyon of northwestern WA (Hamilton and Zack 1999, map 8).

132. Limotettix zacki Hamilton is known only from the 300m high summit of Badger Mt.
just west of the Columbia River in WA (Hamilton 1994a).

Taxa endemic to Pacific coast north of CA (CICADELLIDAE only).

133. Athysanella valla Blocker and Johnson is a grass-feeding leafhopper known only from
southernmost OR west of the Cascade Mts. (Blocker and Johnson 1990).

134. Ceratagallia omani Hamilton is known from Saturna I. off Vancouver Island, BC,
along the entire OR coast, and inland on the Coast Range as high as 1000m (Hamilton 19980).

135. Evacanthus lacunar Hamilton is a Scrophularia-feeding leafhopper known only from
Marys Peak (1100m) in the Coast Range of OR (Hamilton 1983).

136. Latalus occidentalis (DeLong) is known from Marys Peak and adjacent coast of OR,
and possibly also from Vancouver Island in BC (based on an unassociated female). This

52 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

species appears to be a closely related to an undescribed species from 1100m elevation in the
Cuyamaca Mts. of southernmost CA.

137. Limotettix beameri (Medler) is found along the coast of OR including the Coast
Range, and near Puget Sound in WA.

138. Psammotettix diademata Hamilton (see Part 1) is known only from the shore of the
Queen Charlotte Islands in BC (Fig. 2, star).

139. Psammotettix nesiotus Hamilton (see Part 1) is known only from both sides of the
Strait of Georgia that separates Vancouver Island from mainland BC.

Taxa endemic to mountains of CO and adjacent WY.
(A) CICADELLIDAE

140. Athysanella gardenia Osborn was described from the mountains of CO and has
subsequently been recorded from both CO and WY (Blocker and Johnson 1988) without any
more specific information. Its circumscribed distribution seems to indicate montane endemism.

141. Athysanella hyperoche Hamilton (see Part 1) is known only from a pass in the
Laramie Mountains of WY (see Part 1).

142. Diplocolenus configuratus nigrior Ross & Hamilton is a rather rare grass-feeding
leafhopper known only from mountains in AZ and CO (where it probably found glacial-age
refugia), and across about 200 km of the foothills of the Unita Mts. on the borders of ID, UT
and WY (Fig. 8, filled circles) to the west of D. configuratus. It appears to hybridize (Fig. 8,
bull’s-eyes) with both D. configuratus s.s. and D. configuratus bicolor.

143. Errhomus montanus (Baker) is a flightless potentilla-feeding leafhopper found on
high mountains from CO and western WY to southern MT (Hamilton and Zack 1999, map 1).
It was probably more widely distributed during the Pleistocene, and has since retreated into
its montane fastness as temperatures increased 10,000 years ago.

144. Hebecephalus chandleri Hamilton 1s a grass-feeding leafhopper known only from the
Bighorn Mountains of northern WY (Hamilton 1998a). It is a sister-species of H. atralbis
Emeljanov from Siberia.

145. Hebecephalus vinculatus Ball is a grass-feeding leafhopper found in the mountains
of CO (Ball 18995). It was incorrectly recorded from WY and Labrador (see Hamilton 1998a)

146. /diodonus josea (Ball) is a leafhopper of the mountains of CO and southern MT (near
Yellowstone N.Pk.) Nymphs that are also reddish, but evenly coloured instead of maculate,
perhaps may be conspecific; these are known from southern BC and adjacent ID.

147. Norvellina saucia (Ball) is known only from the mountains of CO (Lindsay 1940).

148. Orocastus hyalinus (Beamer) is a grass-feeding leafhopper known only from the
mountains of CO (Beamer 1938).

149. Psammotettix viridinervis Ross & Hamilton is a grass-feeding leafhopper known only
from the mountains of southeastern WY (Ross and Hamilton 1972).

150. Sorhoanus involutus Hamilton (see Part 1) is known only from the mountains of CO.

(B) DELPHACIDAE
151. Kosswigianella irrutilo Hamilton (see Part 1) is known only from the mountains of
CO. It has no known relatives.

Taxa endemic to northern UT and adjacent WY
(A) CICADELLIDAE

152. Athysanella obscura Johnson is known from “Antelope, UT” (Johnson and Blocker
1979) and an unspecified locality in WY (Blocker and Johnson 1990).

153. Hebecephalus abies is a grass-feeding leafhopper that is only known from the
foothills of the Unita Mountains and the West Tavaputs Plateau of UT (Hamilton 1998a).

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 53

154. Hebecephalus filamentus Hamilton & Ross is a grass-feeding leafhopper that occurs
in northeastern UT and also on the mountains of southeastern WY (Hamilton 1998a).

155. Mocuellus quinquespinus Hamilton (see Part 1) is a grass-feeding leafhopper that 1s
only known from a single locality in the foothills of the Unita Mountains of UT (Fig. 9, star),
an area particularly rich in PNW grassland leafhoppers, including Hebecephalus abies (# 149).

156. Norvellina curvata Lindsay is known only from the Grand Teton Mountains of WY
(Lindsay 1940).

(B) DELPHACIDAE

157. Achorotile apicata Hamilton (see Part 1) is a planthopper known only from the Unita
Mountains of northeastern UT. It is related to a transboreal species.

158. Elachodelphax unita Hamilton (see Part 1) is a planthopper known only from the
Unita Mountains of northeastern UT. It is related to E. mazama (#40) from the Cascade
Mountains of WA.

Taxa endemic to mountains of central UT
(A) CICADELLIDAE

159. Errhomus naomi Hamilton & Zack is known only from an isolated location in the Fish
Lake mountains of the Wasatch Plateau, UT. Its relatives inhabit western MT. It feeds on the
common species of the Great Plains, hoary sagebrush (Artemisia cana Pursh) rather than on
the common Rocky Mountain species (A. tridentata Nutt.)

(B) DELPHACIDAE
160. Kosswigianella wasatchi Hamilton (see Part 1) is a planthopper known only from the
Wasatch Plateau of central UT. It is related to a transcontinental species.

CONCLUSIONS

Endemism. Endemics account for approximately half (44%) of the entire PNW grassland
fauna of 275 taxa. If one adds the 22 taxa that are endemic to the mountains of CO, UT and
WY the total endemic fauna rises to 142 taxa (52% of the PNW fauna). This is one of the
highest percentages of regional endemism in North America. The total number of endemics
is third only to those of the prairies and desert grasslands. Clearly this represents an unique
fauna of great age that has managed to survive Pleistocene climate change in some refugium.
But where? This can be determined only after deducing how much the fauna moved in
response to postglacial (Hypsithermal) warming, and to what degree mountain passes proved
to be a barrier to dispersal.

Hypsithermal change. The only valleys in the PNW with few endemics and limited
southern faunal elements are the isolated glaciated valleys of central and eastern BC. These
valleys must have been repopulated largely by prairie insects coming over “Crooked River”
and Crowsnest passes, for they are essentially an Alberta fauna found mainly along the upper
Fraser River (#1, 8, 14, 21, 27, 31) and in the East Kootenay valley (#1, 2, 9, 19, 24, 26, 28).
This faunal invasion probably occurred during the Hypsithermal as these passes are now
completely forested for many kilometres. Hypsithermal grasslands probably extended as far
north as the Mackenzie River but not into the Yukon (Hamilton 1997; see also #50). Faunal
exchange this far north was probably at a still warmer time, possibly during the height of the
Sangamon interglacial 124,000 years ago (Matthews 1979).

It is unlikely that Hypsithermal warming ever brought together ranges of PNW grasslands
sufficiently to permit free faunal interchange. Widely dispersing leafhoppers seldom are found

54 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

throughout PNW grasslands. For example, the endemic leafhoppers Norvellina rubida (#70)
and Sorhoanus xiphosura (#74) range throughout much of the PNW grasslands from Wyoming
to southern British Columbia, but, even more than the grasslands themselves, they have
fragmented ranges (Fig. 5, filled circles and stars).

PNW-prairie passes. In the U.S.A. there is a sharp division between faunas of the PNW
grasslands and those of the prairies. As the two are connected across a number of grassy passes
(Fig. 6) and are botanically identical on both sides, this becomes hard to understand with our
present knowledge. Yet there are at least 59 prairie taxa that range up to the continental divide;
at least six do not cross it at all (the typical subspecies of #32-35, plus Stenometopiellus
cookei, sister-species of #38, and Elachodelphax borealis, sister-species of # 40), while 31
penetrate only a short way into the PNW (#1-31). Five taxa that inhabit the PNW do not cross
these passes although they range up to them (#53, 67, 68, 73, 91). Four PNW endemics have
crossed the divide to invade adjacent foothills prairie (#51, 62, 72, 83). Another 18 PNW taxa
which range up to the continental divide appear to be confined to mountains or high latitudes
(#62-64, 74, 76, 85, 87, 140-151) and are therefore unlikely to invade the prairies.

Yet despite the few cases of faunal exchange between these grasslands, there are many
passes in MT, ID and WY which show numerous cases of insect migrations (Table 1). Three
components seem to govern whether passes are suitable for leafhopper dispersal.

Predictably, “open” (grassland) passes have the largest number of grassland leafhopper
taxa. The wide pass at Great Divide Basin, WY was probably a conduit for 14 species, but
only a single grassland species has been taken actually in the pass which is now very arid. A
similar number probably crossed Bannock Pass, and 10 are still found there. Less certain
evidence has been found that “Grassy Pass” also has been the conduit for 14 species, of which
only three have been found in the pass. Ten to 12 taxa can be inferred to have crossed through
Lemhi, Monida and Lost Trail passes. Possibly “Grassy,” Lemi and Monida Trail passes have
been open grasslands only recently, having been forested during the Pleistocene, and then arid
during the Hypsithermal. Lost Trail Pass is currently forested on its southern flank, but might
have been open during the Hypsithermal.

Elevation is the second most important component governing whether leafhoppers can use
passes. Crowsnest Pass in southern BC (1400 m) and Rogers Pass, one of the lowest passes
in MT (1800 m), are both good leafhopper conduits (for 11 taxa each). MacDonald Pass (2000
m) has a substantially smaller number of taxa associated with it (eight taxa, four in the pass).
Increasing elevation of passes south of this is offset by the increasing elevation of the montane
grassland at this latitude, so that Lemhi and Bannock on the ID/MT border (both at 2200 m)
are “open” passes through which at least 12 taxa have passed.

Direction of the pass also seems to play an important role in dispersal of leafhoppers. The
passes most used by leafhoppers tend to have a north-south extent. East-west passes such as
MacDonald Pass (2000 m) and Chief Joseph, MT (2200 m) show much less influence in
leafhopper spread than one would expect. The direction of transfer has been mainly north to
south at “Crooked River,” Rogers and Monida passes, and south to north at “Grassy Pass”. in
each case the dominant direction is from the prairies into the intermontane grasslands, and not
vice versa. Probably the winds that aid most in leafhopper dispersal are the strong prairie
southerlies of summer. The prevailing northwesterlies of spring and fall appear to occur too
early and late to transfer gravid females. Such winds would carry Homoptera over local
barriers formed by north and east-facing slopes, which are cooler and not as suitable for
grassland insects.

Refugia. There are 32 widespread PNW grassland Homoptera (#34, 36, 41-43, 51, 52-76)
that may be either widely dispersing or merely disjunct relicts of a formerly widespread

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 eP)

distribution. These include 20 taxa (#34, 36, 41-43, 51, 53, 54, 63, 65-76) which fail to match
each other in distribution (e.g., Fig. 5) so no conclusions can be reached about their
Pleistocene distribution. The probability is greatest that these have each survived the
Pleistocene in more than one refugium, from whence they have spread to outlying areas in a
discordant manner. Three other such species are Endria montana (#57) whose range is divided
between grasslands on southern Vancouver Island, BC and those of the Rocky Mountains,
centred around the upper end of the Snake River Valley; Hebecephalus caecus (#60) whose
range is divided even today between the eastern slopes of the Cascades in OR and upper end
of the Snake River Valley; and Hebecephalus hilaris (#64) which is known only from the
Cascade Range of WA and the mountains of southeastern WY.

By contrast, there are 80 taxa endemic to just a part of the PNW (#45-52, 77-151). The
most strikingly different PNW grassland fauna is that of oak savannah and other Coast Range
grasslands such as the grassy “bald” above 1200 m elevation on Marys Peak, OR (Fig. 5, filled
triangle), the latter with two endemic species (#135, 136). This coastal grassland shares with
inland sites only three PNW leafhoppers (#56, 57, 73; see Figs. 15, 17). Seven grassland taxa
(#133-139) are restricted to the Pacific coast side of the Cascade range. One of these,
Psammotettix diademata (#138) is known only from the shores of the Queen Charlotte Islands
far to the north of other PNW grasslands. These islands also have one other endemic
leafhopper, Evacanthus grandipes Hamilton (1983) and an endemic spittlebug, Aphrophora
regina Hamilton (1982), both of which are forest insects, so the weather on the seaward side
of the islands was probably mild enough for all three of them to weather Pleistocene glaciation
there. Further evidence that the Queen Charlotte Islands were not glaciated during the last ice
advance is provided by Nearctic earthworms, that exist here and on Vancouver Island although
eradicated from the rest of Canada by glaciation (Hendrix and Bohlen 2001, fig. 1).

Other geographic areas with distinctive faunas, from north to south, are inland BC plus AB
with six taxa (#77-82), eastern ID plus adjacent MT with 10 taxa (#83-92), inland WA with
18 taxa (#115-132), and inland OR plus western ID with 18 taxa (#97-114). Do these four
geographic areas represent four additional Pleistocene refuges? Concurrence between
individual distribution patterns should resolve this question.

In addition to these 54 localized taxa endemic to inland PNW grasslands, there are six
leafhoppers that represent three pairs of sister taxa with widely disjunct ranges (#45-52). Three
of these, one from each pair, are associated either with the eastern slopes of the Cascade
Mountains in OR (#45, 48), or with the Cascades themselves in the south of OR (#52). Their
sister-taxa are found either on the upper reaches of the Clark Fork river in MT (#46, 47) or are
widely dispersed around the drainage basin of the Columbia River and the Clark Fork river
(#51). This suggests a PNW-wide dispersal pattern that became fragmented into an OR
component associated with the southern Cascades, a Columbia basin component, and a Clark
Fork component; subsequent catastrophic flooding of the Columbia basin in the late
Pleistocene may have wiped out intervening populations of all but the species most adapted
to high elevations, Sorhoanus debilis (#51). This disjunct distribution pattern is similar to that
of Hebecephalus caecus (#60), discussed above. Confirmation of a fragmented grassland
habitat is found in the several groups of local endemics that mirror this pattern, as well as some
widespread prairie species whose PNW range is separated by hundreds of kilometres, e.g.,
Flexamia inflata (Osborn & Ball) (Whitcomb and Hicks 1988, Fig. 41).

The easternmost of these disjunct grasslands occurs along the upper reaches of the Clark
Fork River from St. Regis, MT to the continental divide. This area has seven of the eight
endemic leafhoppers of MT (#46, 47, 93-96, plus Macrosteles skalkahiensis Beirne from 2200
m elevation in Skalkaho Pass, east of the Bitterroot Valley: Beirne 1952). Most of these
endemics are members of the flightless genus Errhomus. Three (#93, 94, 96) inhabit two of
the four main tributaries of the valley that was once Lake Missoula: the Bitterroot and

56 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Blackfoot rivers. A fourth (#95) is found on the west bank of the Clark Fork river where these
rivers converge at Missoula. This flightless species must have survived inundation of the valley
on south-facing mountain slopes at least 300 m higher that its present locality. Another genus
is represented along the third tributary, Rock Creek: Athysanella repulsa (#46).

A much larger number of local endemic taxa are associated with the land to the east of the
Cascade Range in WA and OR (#45, 48, 52, 60, 97, 98, 106, 109, 113, 114, 116-129, 131,
132), a total of 27 endemics in this area. Sixteen of these species are highly localized, flightless
leafhoppers of WA and adjacent OR (#107, 108, 116-129, 131), and two of these each have
two subspecies (#116, 117; 124, 125) in adjacent areas, suggesting occupancy of this area for
a considerable time. Three others appear to have had another refugium in southwestern OR
(#52, 97, 114). Two became stranded on isolated mountain crests east of the Cascades in WA
(#115, 132). Two species are known from single hills in south-central OR (#99, 106). It
appears clear that at least patches of grassland survived along with their leafhopper faunas in
numerous isolated situations, probably south-facing hillsides and canyon slopes. The only
species of Errhomus that inhabits the most arid parts of the Columbia plateau, FE. similis, now
favours north and east-facing slopes of coulees that are not so subject to drought as level
ground or south and west-facing slopes (Hamilton and Zack, 1999). This suggests that they
found glacial-age refugia on south-facing slopes. The grasslands of WA were probably more
severely modified than those of OR, resulting in a much more fragmented WA fauna.

The intervening area between these eastern and western grasslands is largely Columbia
basin. There, 13 endemic taxa are members of the flightless leafhopper genus Errhomus which
feed on arid-adapted perennial composites: four in WA and adjacent ID (#58, 59, 103, 130)
and nine along Hell’s Canyon on the ID/OR border, and up into the Blue Mountains of OR
(#100-102, 104, 105, 107, 108, 110-112). All other, more mobile endemic leafhoppers have
migrated northwards from the Columbia basin, usually into many adjacent areas (#32, 33, 35,
56, 78 and probably many of the widely dispersing species). Four (#39, 40, 77, 79) are known
only from glaciated valleys in southern BC and northern WA and must have come from refugia
further south. |

Nine species belonging to five genera (#38, 61, 84, 86, 88-92) are associated with the
headwaters of the Snake River and one (#60) has a disjunct population associated with a
tributary river northeast of Boise in the western part of the state. Curiously, none are members
of Errhomus although the genus is speciose both east and west of this valley. Pleistocene
volcanism that flooded much of the Snake River valley in ID may have decimated populations
of this arid-adapted genus. Other leafhoppers may have survived in this region through their
greater powers of dispersal. For example, Mocuellus caprillus anfractus (#91) has a nearly
circular range strongly suggestive of dispersal from a single valley bottom refuge. The other
species are scattered in various valleys along the south-facing slopes of the ranges crossing
central ID or the upper Snake canyon where it penetrates the WY border. This pattern is highly
suggestive of a Pleistocene rain-shadow grassland in the upper Snake River plains, together
with isolated grassland patches on south-facing slopes from the Lost River Range westward
to the present site of Boise.

Other leafhoppers may have adapted to cooler conditions during the Pleistocene, and
subsequently sought refuge after glaciers melted by migrating to higher elevation grasslands.
Some PNW endemics (e.g., #55) are now limited to a few valley bottom sites in a linear
pattern suggesting migration north from a glacial refugium. At least four widespread species
(#20, 27, 43, 73) have migrated far northwards into glaciated areas. Four others (#29, 51, 62,
72) are now found in mountains or grasslands at lower elevation less far north. One (#37) is
known only from a glaciated mountain. Fifteen other endemic taxa are characteristic of limited
areas of the Rockies of southern MT/ID (Bitterroot Range) south to CO (#83, 85, 87, 140-
151). These may have shifted their ranges vertically rather than longitudinally, in the same

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 57

manner that nine endemics in UT and adjacent WY (#152-160) almost certainly have done.
One flightless species (#143) and two others (#142, 146) are presently found in isolated
mountains in three states, so the refugium may have extended a considerable distance along
the mountain range. One species (#144) is presently confined to the Bighorn Mountains of
north-central WY, which may represent an isolated grassland refugium.

Three species are presently known only from the “Crooked River” Pass vicinity, on the
upper Fraser River of BC (#81) and south-facing bluffs along the Peace River in Alberta (#80,
82). These may once have been inhabitants of the eastern slopes of the Rockies; but this would
require a very extensive displacement northwards during the Hypsithermal not evident among
other Homoptera.

Pleistocene climate. There is ample evidence that PNW grasslands and their fauna are
endemic phenomena. Indeed, there is only evidence for a Hypsithermal invasion of prairie
grasslands into two northern, deglaciated mountain valleys. How, then, can one reconcile the
biological evidence for extensive endemism with the geological evidence for extensive
permafrost patterns across the breadth of the PNW?

First, glacial-age circular features known as “stone rings” may not be indicative of
permafrost at all. That they have been cited as evidence of permafrost (Brunnschweiler 1962)
may be due to superficial similarities to ice mounds in modern tundra. This view has not found
favour among modern geologists (Matthews 1979). Stone rings are found only on tablelands
in the PNW (Fig. 19). These structures may point towards yearly alternations of extremes of
heat and cold. They were probably formed of rock fragments forced outwards around the edges
of ponds that froze solid each winter, then completely melted again each summer, leaving a
depression to collect yet more water and thus grow to a larger size each successive year.

Second, land slope and direction are obvious components of microclimate. South-facing
slopes are particularly subject to aridity and high summer temperatures; west-facing slopes
show similar but less severe conditions and are the modern preferred habitat of PNW grassland
leafhoppers. A number of endemic grassland leafhoppers in the Yukon are associated with
south-facing slopes (Hamilton 1997). That such a phenomenon was very much a part of
glacial-age ecology is shown by pollen and subfossil assemblies in the Saint Lawrence valley
of Québec (Mott et al. 1981). At a time when the whole valley was extensive “spruce-aspen
forest” following the retreat of “tundra” conditions 10,000 years ago, Mont St. Hilaire had high
levels of oak pollen and even an oak-feeding treehopper (Membracidae), presumably from the
sun-warmed slopes.

Thirdly, timing is probably a factor. Permafrost soil structures indicate the overall coldness
of a site, but not the duration of that cold. Thus, a long and severe winter could well be
succeeded by a mild (perhaps even hot) but short summer. This is probably the explanation of
why the majority of endemic flightless leafhoppers in Errhomus have survived the Pleistocene
on the Columbia Basin by feeding on a spring-flowering host, balsamroot.

In total, 38 PNW-endemic taxa belong to Errhomus (#58, 59, 93-96, 98-105, 109-114,
116-131, plus six subspecies of postglacial origin). These often occupy adjacent territories that
follow geological features erased long before the Pleistocene (Hamilton and Zack 1999). The
evidence is thus strong that these regions suffered little climate change during the Pleistocene,
although the extent of the arid areas of the Columbia basin were probably more limited than
at present. Errhomus are mostly insects that emerge early in spring to take advantage of
perennials that dry up in summer. During the Pleistocene their hosts may have grown in mid-
summer instead of springtime. If these leafhoppers are active during cool weather, they would
have been able to adjust their seasonality so that their ranges were little affected by Pleistocene
conditions.

Nine of the 12 balsamroot-feeding species of Errhomus in WA and OR are found very

58 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

close to (but not on) ring-patterned ground (Fig. 19, # 1-9). Six exceptions to this lifestyle are
only found in the most northerly or montane species: E. calvus (Fig. 19, # 10) north of the
Columbia, FE. pallidus in the Blue Mountains of OR (Fig. 19, # 12), E. solus and E. rivalis
(Fig. 19, # 14-15) near the deepest part of the Lake Missoula valley, FE. montanus in the
highest mountains of CO and WY, and E£. naomi above 3000 m in the Wasatch Plateau of UT.
All have shifted to other hosts, probably when balsamroot became scarce under Pleistocene
conditions. In all of the glaciated area east of OR, only the highly localized E. camensis (#94)
still feeds on balsamroot.

Additional evidence for short but warm Pleistocene summers close to ice margins comes
from eastern Canada. There, the endemic Gulf of St. Lawrence aster, Aster lautentianus
Fernald is presumed to have weathered the Pleistocene on the ice-free Magdalen Is and
subsequently spread to adjacent deglaciated coasts of New Brunswick and Prince Edward I
(Hamilton 2002, fig. 2). This aster flowers after two to three months, as contrasted to its
continental relatives A. brachyactis Blake and A. frondosus (Nutt.) T. & G. at six to eight
months each (Houle and Haber 1990).

How hot was the PNW glacial age summer? East of the Rockies and south of the ice sheet,
summer was much cooler than at present except in the far south (Brunnschweiler 1962). This
was probably due to extensive cloud cover associated with increased precipitation. Prairie-like
conditions (“periglacial grassland’) existed only close to the ice front (Hamilton 19945) where
a permanent high pressure system over the glacier field suppressed summer rainfall. South of
the PNW there is also evidence of reduced evaporation and increased precipitation: glacial-age
Lake Bonneville completely filled the northwestern valley of UT to overflowing (it is now
shrunk to Salt Lake). But, as in eastern North America, summer temperatures near the ice front
may well have been close to spring temperatures today. Evidence for this in western North
America comes from four endemic species (#80-82, 138) associated with the northern
grasslands of the Peace River in AB, the upper Fraser River and on the Queen Charlotte
Islands of BC. Such a suppression of rainfall is indicated by the distribution of loess in the
PNW, which would have been driven from glacial moraines southward with strong winds off
the ice sheet across treeless areas, and deposited where open vegetation (grasslands or open
woodlands) broke the force of the wind. Major loess deposits on the Columbia basin as far
south as the Blue Mountains of OR, and on the Snake River plains of ID, suggest the presence
of open forests or grasslands there (Fig. 20).

Conversely, the numerous glacial-age lakes across UT, NV and southern OR show that
these areas were cooler and wetter than at present. This beit must have been a zone of constant
frontal activity between a stationary high-pressure system over the ice sheet, and a stationary
low-pressure system over the semideserts of Arizona and New Mexico (Fig. 20). Between
these two pressure systems the prevailing winds would have been driven from east to west,
drawing moisture from the Gulf of Mexico northwestward towards UT. This would have
constituted a real monsoon system, reversing the usual air flow patterns of the winter season.

Summary. At least nine PNW Pleistocene grasslands can be deduced. These are listed in
descending importance as refuges for leafhoppers and planthoppers:

(1) east slopes of the Cascade Range (Fig. 2C), from Washington state to northern California,
with 27 endemics, two of which are sister species of Montana endemics;

(2) Columbia basin (Fig. 2E) including Palouse hills of Washington and canyons of western
Idaho, with 22 endemics, mostly with flightless females incapable of traversing deep
canyons;

(3) Rocky Mountains of Montana south to Colorado (Fig. 2J) where 19 endemics must have
persisted on grassy south-facing slopes close to their present highly restricted ranges;

(4) the Snake River and south-facing slopes north of the Snake River in southern Idaho (Fig.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 59

2F) must have been grassland near the headwaters, as all 12 endemics are found in that end
of the valley;

(5) the Pacific coast, Mary’s Peak (Fig. 5, triangle) and the Willamette Valley (Fig. 2B) west
of the Cascade Range of Oregon have nine endemics;

(6) the Upper Clark Fork valley in western Montana (Fig. 2G) has six grassland endemics
(including ones with flightless females) which must have survived the Pleistocene on the
edges of glacial Lake Missoula;

(7) grassland patches along the upper Fraser and Peace rivers of BC have three endemics
which are deep in glaciated areas and must have come from grasslands east of the
Continental Divide (Fig. 2H, northern end); these and possibly a single widespread PNW
species (#50) with a similar distribution along the Fraser and Peace rivers to the Mackenzie
River are probably remnants of a periglacial grassland near the ice front in Alberta;

(8) the mountains of south-central OR (Fig. 2D) have two endemics; and

(9) the coast of the Queen Charlotte Islands (Fig. 2, star) has a single endemic grassland
leafhopper. Other grasslands on islands further south (Fig. 2A) were repopulated by
hypsithermal invasion.

Additional refugia might have been in the mountains of Utah and adjacent Wyoming, but these

may have been further southwards.

ACKNOWLEDGEMENTS

This study is the outcome of 23 years of leafhopper surveys (1948-1971) by H.H. Ross to
document the leafhopper fauna of North American grasslands, and is dedicated to the memory
of “Herb.” He and his students took more than 1,400 samples, beginning on July 16, 1948 at
Ozark, Illinois (note code GL 109). This work was continued by R.F. Whitcomb and myself.
The Pacific Northwest maps were based on an original prepared by Bonnie Hall for Oregon
State University, used by permission. The manuscript was reviewed by D. Lafontaine of
AAFC, Ottawa.

REFERENCES

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18995. Some new species of Deltocephalus. The Canadian Entomologist 31: 188-192.

Bartlett, C.R. and L.L. Deitz. 2000. Revision of the New World Delphacid planthopper genus Pissonotus
(Hemiptera: Fulgoroidea). Thomas Say Publications in Entomology: Monographs. 234 pp.

Beamer, R.H. 1936. New leafhoppers from the western United States (Homoptera-Cicadellidae). The Canadian
Entomologist 68(11): 252-257

1938. Some new species of leafhoppers (Homoptera-Cicadellidae). Journal of the Kansas
Entomological Society 11(3): 77-84.

Beamer, R.H. and L. Tuthill. 1935. A monograph of the genera A/apus and Hebecephalus (Homoptera,
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Circular 277.

FAUNAL SYNOPSIS AND INDEX

to species numbered in text
(*) indicates taxa of probable post-glacial origin

CALISCELIDAE A. nielsoni Blocker #84 (ID)
Bruchomorpha beameri Doer. #1 (prairie) A. obesa Ball & Beamer #4 (prairie)
A. obscura Johnson #152 (UT-
CICADELLIDAE WY)
Acinopterus viridis Van Duz. prairie A. occidentalis megacauda H. #32 (PNW)
Amblysellus grex (Oman) southern A. repulsa Hamilton #46 (MT)
A. wyomus Kramer #2 (prairie) A. robusta Baker #5 (prairie)
Athysanella acuticauda (Bak.) northern A. terebrans (Gillette & Bak.) #6 (prairie)
A. attenuata Baker #3 (prairie) A. strobila Blocker southern
A. castor Hamilton #83 (MT) A. utahna Osborn prairie
A. expulsa Blocker #45 (OR) A. valla Blocker and Johnson #133 (OR)
A. gardenia Osborn #140 (CO- Attenuipyga omanae (Beamer) #53 (PNW)
WY) A. platyrhynchus setosa Om. #33 (PNW)

A. hyperoche Hamilton #141 (WY) Auridius auratus (Gill. & Bak.) prairie

A. cosmeticus Hamilton #47 (MT)
A. helvus (DeLong) #7 (prairie)
A. ordinatus (Ball) #8 (prairie)
A. o. crocatus Hamilton #34 (PNW)
A. safra Hamilton #54 (PNW)
A. vitellinus Hamilton #48 (OR)
Balclutha confluens (Rey) northern

B. manitou (Gillette & Baker) northern

B. neglecta DeLong & Dav.) prairie

B. punctata (Fabricius) northern
Ballana spp. (11) southern

B. atridorsum (Van Duzee) prairie

B. chrysothamna DeLong ? ID)

B. curvidens DeLong ? (ID)

B. ortha DeLong prairie

B. spinosa DeLong ? (ID-OR)
Carsonus aridus (Ball) southern

C. furcatus (Oman) southern

C. irroratus (Ball) southern
Ceratagallia acerata Ham. #97 (OR)

C. arida (Oman) #9 (prairie)
C. artemisia Oman southern

C. californica (Baker) southern

C. clino Hamilton #98 (OR)

C. gallus Hamilton #55 (ID-UT)
C. lophia Hamilton #99 (OR)

C. nanella zacki Hamilton #35 (BC-WA)
C. okanagana Hamilton #77 (BC)

C. robusta poudris (Oman) — southern

C. omani Hamilton #134 BC/OR)
C. siccifolia (Uhler) northern

C. siccifolia compressa H. #36 (PNW)
C. vipera Hamilton #115 (WA)

C. viator Hamilton
C. vulgaris (Oman)
Chlorotettix similis DeLong

#10 (prairie)
#11 (prairie)
#56 (BC/ID/
OR)
Colladonus aureolus (Van D.) southern
Commellus sexvittatus (V. D.) prairie

Cuerna cuesta Hamilton #78 (BC-MT-
WA)
Deltocephalus artemisiae
Gillette & Baker prairie
D. fuscinervosus Van Duzee — southern
D. minutus Van Duzee southern

D. vanduzeei Gillette & Baker ? (PNW)

D. vanfus Kramer southern
Diplocolenus brevior R. & H. northern

D. configuratus (Uhler) #13 (prairie)
D. c. bicolor Hamilton #85 (ID-MT-

WY)

D. c. nigrior Ross & Hamilton #142 (AZ-ID)
D. evansi (Ashmead) northern
Draeculacephala bivoltina H. southern

D. borealis orea Hamilton ? (PNW)

D. crassicornis Van Duzee —_? (PNW)

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

D. robinsoni Hamilton
Elymana circius Hamilton
Endria inimica (Say)

E.. montana (DeLong & SI.)
Errhomus affinis Oman

E. affinis attenuatus Hamilton
E. bracatus Hamilton

E. brevis Oman

E. brevis simcoe Oman

E. calvus Oman

E. camensis Hamilton
E. josephi Oman

E. lineatus (Baker)

E. lineatus cordatus Hamilton
E. lineatus idahoensis Oman
E. lineatus umatilla Oman

E. montanus (Baker)

E. naomi Hamilton

E. ochoco Oman

E. pallidus Oman

E. paradoxus Oman

E. picturatus Hamilton

E. praedictus Hamilton

E. reflexus Oman

E. rivalis Hamilton

E. satus Oman

E. serratus Oman

E. s. instabilis Oman

._ S. obliteratus Hamilton

. similis Oman

. Ss. confinis Oman*

_ Ss. dubiosus Oman*

. S. kahlotus Oman

. medialis Oman

. minutus Oman*

. nanus Oman*

_ relativus Oman*

. sobrinus Oman

. socius Oman

. truncus Oman*

. zonarius Oman

. Solus Oman

E. variabilis Oman

E. v. erratus Hamilton

E. v. gracilis Hamilton

E. v. mimicus Hamilton

E. wolfei Oman

E. winquatt Oman

Euscelis relativa (Gill. & B.)
Evacanthus lacunar Hamilton
Flexamia decora Beam. & T.
F. inflata (Osborn & Ball)
F. serrata Beamer & Tuthill

E
ie
18)
ie

iB
E.
Ee
Hs)

HRA AAA AHA UNA YA

Ve
EE
Ee
E
E
E

northern
#14 (prairie)
prairie
#57 (PNW)
#100 (ID)
#101 (OR)
#93 (MT)
#116 (WA)
#117 (WA)
#118 (BC-
WA)
#94 (MT)
#102 (OR-
WA)
#58 (WA)
#103 UID)
#104 (ID)
#105 (OR)
#143 (CO/
MT-WY)
#159 (UT)
#106 (OR)
#107 (OR)
#119 (WA)
#120 (WA)
#121 (WA)
#122 (WA)
#95 (MT)
#123 (WA)
#108 (OR)
#124 (WA)
#125 (WA)
#126 (WA)
#126 (WA)
#59 (WA)
#59 (WA)
#109 (OR)
#109 (OR)
#109 (OR)
#126 (WA)
#127 (WA)
#128 (WA)
#109 (OR)
#129 (WA)
#96 (MT)
#130 (WA)
#110 (OR)
#111 (ID)
#112 (OR)
#131 (WA)
#113 (OR)
northern
#135 (OR)
#12 (prairie)
prairie
prairie

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Frigartus frigidus (Ball) prairie
Gyponana hasta DeLong southern
Hardya dentata (Osborn & B.) prairie
Hebecephalus abies Hamilton #153 (UT)

63
N. curvata Lindsay #156 (WY)
N. perelegantis (Ball) southern
N. rubida (Ball) #70 (PNW)
N. saucia (Ball) #147 (CO)

H. caecus Beamer #60 (ID/OR)

H. callidus (Ball) #61 (PNW)

H. chandleri Hamilton #144 (WY)

H. crassus (DeLong) #62 (BC/
ID-WY)

H. crenulatus Hamilton #86 (ID)

H. ferrumequinum Hamilton #87 (ID)

H. filamentus Hamilton & R. #154 (ID)

H. firmus Beamer #63 (WA/
MT-WY)

H. hilaris Beamer #64
(WA/WY)

H. occidentalis Beamer & T. prairie

H. planaria Hamilton #79 (BC)

H. picea Hamilton #88 (ID)

H. pugnus Hamilton #89 (ID)

H. rostratus Beamer & Tut. _ prairie

H. sagittatus Beamer & Tut. #65 (PNW)

H. truncatus Beamer & Tut. _ prairie

H. veretillum Hamilton #90 (ID)

H. vinculatus Ball #145 (CO)

Hecalus montanus (Ball) northern

H. viridis (Uhler) prairie

Idiodonus heidemanni (Ball) #15

(southern)
I. josea (Ball) #146
(CO/MT)

Laevicephalus salarius Knull ? (BC/UT)

Latalus curtus Beamer & Tut. #67 (PNW)

L. histrionicus Beirne #49 (BC/AZ)

L. intermedius Ross & H. #50 (PNW)

L. missellus (Ball) northern

L. mundus Beamer & Tuthill #68 (PNW)

L. occidentalis (DeLong) #136 (OR)

N. vermiculata Lindsay #71 (ID/CO)
Orocastus hyalinus (Beamer) #148 (CO)
O. labeculus (DeLong) #19 (prairie)
O. pinnipenis Ross & H. #72 (ID-MT)
O. perpusillus (Ball & DeL.) #20 (prairie)
O. tener (Beamer & Tuthill) southern
Paluda gladiola (Ball) northern
Paraphlepsius lascivius (Ball) prairie
P. occidentalis (Baker) southern
Pinumius sexmaculatus

(Gillette & Baker) #21 (prairie)
Prairiana kansana (Ball) #22 (prairie)
Psammotettix amplus (D. & D.southern

P. attenuens (DeLong & D.) #73 (PNW)
P. beirnei Greene #37 (BC)
P. dentatus Knull southern

P. diademata Hamilton #138 (BC)
P. greenei Hamilton #114 (OR)
P. lividellus (Zetterstedt) northern
P. nesiotus Hamilton #139 (BC)
P. obesus Knull southern

P. shoshone (DeLong & Dav.) southern

P. totalus (DeLong & Dav.) #23 (ID-
MT-WY)
P. viridinervis Ross & H. #149 (WY)
Rosenus cruciatus
(Osborn & Ball) #24 (prairie)

R. decurvatus Hamilton & R. #80 (BC)
R. obliquus (DeLong & Dav.) #92 (ID)

Lebradea flavovirens (G. & B.)northern
Limotettix beameri (Medler) #137

(OR/WA)
Limotettix bullatus (Ball) southern
L. finitimus (Van Duzee) southern
L. glomerosus (Ball) southern
L. symphoricarpae (Ball) prairie
L. zacki Hamilton #132 (WA)
Lonatura salsura (Ball) prairie
Lystridea uhleri Baker southern
Mesamia frigida (DeL. & H.) southern
M. ludovicia Ball #16 (prairie)

Mocuellus caprillus Ross & H.#17 (prairie)

M. c. anfractus Hamilton #91 (ID)
M. c. strictus Ross & H. northern
M. larrimeri (DeLong) #69 (PNW)
M. quinquespinus Hamilton #155 (UT)

Norvellina clarivida (Van D.)
N. columbiana (Ball)

#18 (prairie)
prairie

Scaphytopius diabolus

(Van Duzee) southern
S. oregonensis (Baker) southern
Sorhoanus bimaculatus

(Gillette & Baker) northern
S. debilis (Uhler) #51 (PNW)
S. involutus Hamilton #150 (CO)
S. uhleri (Oman) northern
S. virilis Hamilton #52 (OR)
S. xiphosura Hamilton #74 (PNW)

Stenometopiellus vader Ham. #37 (ID-MT)

Stragania atra (Baker) prairie
S. rufoscutellata (Baker) prairie
Texananus cumulatus (Ball) southern
T. dolus DeLong prairie
T. extremus (Ball) southern
T. lathropi (Baker) southern
T. latipex (DeLong) southern
T. oregonus (Ball) southern
T. proximus Crowder southern

Twiningia fasciatus (Beamer) southern

T. pellucidus (Ball) southern
T. scrupulosus (Ball) southern
Unoka dramatica Hamilton #39 (BC)

64

U. gillettei Metcalf
Xerophloea zionis Lawson

DELPHACIDAE

Achorotile apicata Hamilton

Caenodelphax atridorsum
(Beamer)

Delphacodes anufrievi Wils.

D. gillettei (Van Duzee)

D. lineatipes (Van Duzee)

D. neocclusa Muir & Gifford

D. occlusa (Van Duzee)

Elachodelphax mazama Ham.

E. pedaforma (Beamer)
E. unita Hamilton
Eurybregma eurytion Ham.
E. magnifrons (Crawford)
Javesella lutulentella

(Muir & Gifford)
Kosswigianella irrutilo Ham.
K. wasatchi Hamilton

#25 (prairie)
southern

#157 (UT)
#75
(MT/OR)
northern
southern
northern
southern
southern
#40 (WA)
#27 (prairie)
#158 (UT)
#41 (PNW)
#27 (PNW)

southern
#151 (CO)
#160 (UT)

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Laccocera canadensis Beirne #26 (prairie)
L. flava Crawford #29 (prairie)
Laccocera lineata Scudder _ prairie

L. oregonensis Penner #42 (PNW)
L. vanduzeei Penner #43 (PNW)
L. vittipennis Van Duzee #30 (prairie)
Megamelanus bicolor Ball southern
Nothodelphax foveatus

(Van Duzee) #3 1 (prairie)
N. glacia Wilson northern
N. venustus (Beamer) #44 (PNW)
Parkana alata Beamer southern

Paraliburnia furcata Hamilton#81 (BC)

P. kilmani (Van Duzee) northern

P. lecartus Hamilton #82 (BC)
Pissonotus frontalis (Crawford) southern
P. rubrilatus Morgan & B. #76 (PNW)
Prokelisia carolae Wilson southern

P. salina (Ball) prairie

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 65

SCALE IN MILES
50

100

50 100 150
SCALE IN KILOMETERS

Figures 1-3. Distribution of Pacific Northwest grasslands and widespread endemic
leafhoppers. 1, provinces and states of the Pacific Northwest (boxed area); 2-3, geographical
areas of PNW grasslands, with ID/MT border (box) enlarged to show Clark Fork valley system
(shaded in Fig. 2, GA-GC in Fig. 3). Bold line: continental divide indicated by dividing area
H (prairies) from areas F (Snake River plains) and G (Clark Fork valley); grey lines: county
boundaries. For abbreviations, see text.

66 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Figures 4-5. Distribution of types of PNW grasslands (4) and leafhoppers (5). Black, Palouse
grasslands; stippled, grassland-sagebrush ecotone; outline, fescue grasslands; dashed outline,
oak savannah; filled triangle, Evacanthus lacunar on Mary’s Peak; filled circle, Norvellina
rubida; star, Sorhoanus xiphosura.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 67

SCALE IN MILES
50 100

0 50 100 150
SCALE IN KILOMETERS

Figure 6. Distribution of Unoka gillettei (stars) and U. dramatica (filled circles) west of
101°W, showing invasion of montane area across Pacific continental divide (dashed line).
Major passes from British Columbia to Wyoming (circles), from N to S: A, “Crooked River
Pass,’ BC; B, Crowsnest Pass, BC/AB; (continued in detail) C, Rogers, MT (1800 m); D,
MacDonald Pass; E, “Grassy Pass”, MT; F, Lost Trail Pass, ID/MT; G-H, Lemhi and Bannock
passes, ID/MT; J, Monida Pass, ID/MT; K, Great Basin Divide, WY. Open circles represent
most important passes for leafhopper dispersal; split circles, other passes.

68 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

SCALE IN MILES
100

50 100 150
SCALE IN KILOMETERS

Figure 7. Distribution of the leafhopper Athysanella attenuata west of 101°W. Box, area of
detail; solid line, continental divide; open circles, dispersal routes across Lemhi and Bannoch
passes.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 69

SCALE IN MILES
ie) 50 100

ie) 50 100 150
SCALE IN KILOMETERS

24 oe ite

a

Figure 8. Distribution of the leafhoppers Diplocolenus configuratus (dashed line and open
circle), D. configuratus bicolor (star) and D. configuratus nigrior (filled circle) west of
101°W. Box, area of detail; oval, populations confined to mountains; bull's eye, hybrid

populations.

70 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

SCALE IN MILES
fe) 50 100

ie) 50 100 150
SCALE IN KILOMETERS

Figure 9. Distribution of the leafhoppers Mocuellus quinquespinus (star), M. caprillus (dashed
line), M. caprillus anfractus (filled circles) and M. caprillus strictus (open circles). Box, area

of detail; ovals, populations confined to mountains; arrows, inferred dispersal route across
Lemhi and “Grassy Pass.”

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 Al

SCALE IN MILES
100

50 100 150
SCALE IN KILOMETERS

Figure 10. Distribution of the leafhopper Orocastus perpusillus west of 101°W. Box, area of
detail; ovals: southern populations confined to mountains; arrows, inferred dispersal route
across Rogers and Monida passes.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

i2

o
= on
—«
wW
a =
oi Ww
= 382
eal
Oo a
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Distribution of the planthoppers Eurybregma eurytion (open circles) and

Eurybregma magnifrons (filled circles). Ovals

inferred dispersal routes.

11

igure

F

arrows,

2)

fined to mountains

10ns con

populat

>

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 i

: el :

SCALE IN MILES

0 50 100
ie) 50 100 150

SCALE IN KILOMETERS

Figure 12. Distribution of the planthoppers Laccocera oregonensis (open circles) and
Laccocera flava (filled circles). Box, area of detail; ovals, populations confined to mountains;
arrows, inferred dispersal route across MacDonald or “Grassy Pass.”

ies J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

’
.
‘
i

SCALE IN MILES
100

50 100 150
SCALE IN KILOMETERS

Figure 13. Distribution of the planthoppers Laccocera spp. (vittipennis group) west of 101°W.
Open circles, L. vanduzeei; filled circles, Laccocera vittipennis; outlines, populations confined
to mountains; arrows, inferred dispersal route across MacDonald, “Grassy Pass’, Lost Trail,
Lemhi, Bannock, Monida and Raynolds passes.

dS

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

5

—x
o ||
ud Ww
| Os
= 96
J
= =

wW
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q wow
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populations confined to mounta

ovals

: area of detail;

Figure 14. Distribution of the leafhoppers Sorhoanus debilis (c

(star). Box

pass.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

76

Wise:
2 pd ere

Oe ee
pr ales s
Oras &

5)
tJ
Ww
Wy ff
= 82
a
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Ww
ad Oo
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,

arrows,

Rogers and MacDonald passes (open circles).

b)

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p)

Distribution of the leafhopper Endria montana. Box, area of detail

15

igure

nferred dispersal route across Lost Trail

F
1

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 77

SCALE IN MILES
50 100

50 100 150
SCALE IN KILOMETERS

Figure 16. Distribution of the leafhopper Orocastus pinnipenis. Outline, eastern populations
confined to mountains; open circles, taken in passes; arrows, inferred dispersal routes through
Lost Trail and MacDonald passes.

78 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

v4

[ ee
| se

Ny
oe

SCALE IN MILES
50 100

50 100 150
SCALE IN KILOMETERS

Figure 17. Distribution of the leafhopper Psammotettix attenuens. Outline, populations
confined to mountains; open circles, samples taken in passes; arrow, inferred dispersal route
through Lost Trail Pass.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 79

IN MILES
SCALE oH 100

50 100 150
SCALE IN KILOMETERS

Figure 18. Distribution of the leafhopper Cuerna spp. west of 101°W. Box, area of detail;
dashed line, western extent of C. septentrionalis; filled circle, C. cuesta; arrows, inferred

dispersal route.

Figure 19 (following page). Distribution of glacial-age patterned ground (stippled) in the
PNW and probable glacial refugia of relict populations of the flightless genus Errhomus
(stars). 1, E. picturatus; 2, E. serratus obliteratus; 3, E. reflexus; 4, E. serratus instabilis; 5,
E. satus; 6, E. praedictus; 7, E. paradoxus; 8, E. winquatt; 9, E. ochoco; 10, E. calvus; 11, E.
Josephi; 12, E. pallidus; 13, E. bracatus; 14, E. rivalis; 15, E. solus; 16, E. camensis. Circled:
species that have shifted to non-balsamroot hosts. All species shown in modern sites, except
for 3, 10, 11 and 15 which are shown with contracted ranges.

80 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

SCALE IN MILES
50 100

° so ite, 0} 150
SCALE IN KILOMETERS

E25 ee AY E> SS BE Ps Oi a a

i

ag HULU

es o|o:
un
I

Th
ant ;
Wi
7

go
MI
vt

al!

Figure 20. Hypothetical summer monsoon over PNW and areas to the south during the height
of Pleistocene glaciation, showing permanent high pressure system (cold front) over WA, ID
and MT and permanent low pressure system (warm front) over AZ; prevailing winds between
these would be east to west (reverse of winter direction), bringing wetter than normal
conditions to UT, NV and southern OR. Outline, ice cover; stippled, loess deposits and

canyons; striped, glacial-age lakes.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 81

Occurrence of the spider mite predator Stethorus punctillum
(Coleoptera: Coccinellidae) in the Pacific Northwest

D.A. RAWORTH AND M.C. ROBERTSON

AGRICULTURE AND AGRI-FOOD CANADA,
P.O. BOX 1000, AGASSIZ, BRITISH COLUMBIA, CANADA V0M 1A0

Stethorus punctillum Weise is a Palearctic species first reported in North America by
Brown (1950), but found in the 1940's in collections from Ontario (Putman 1955). The beetle
was collected on Lulu Island near Vancouver BC in 1950 (Tonks 1953). Since 1997 it has
been mass reared and sold (Applied Bio-nomics Ltd., Saanich, BC) as a biological control
agent for two-spotted spider mites, Tetranychus urticae Koch (Acari: Tetranychidae) (Raworth
2001).

According to Gordon (1985) S. punctum picipes Casey, a species native to North America,
and the introduced species S. punctillum have overlapping distributions on the west coast of
North America. Concern about the potential impact of commercial releases of S. punctillum
on the species complex in the Lower Fraser Valley prompted a review of our recent collections
of Stethorus, and collections of two other researchers, from the Pacific Northwest.

Stethorus spp. were collected during various studies from: raspberry, Rubus idaeus L.
(Rosaceae) at Abbotsford BC (1986, 1996), Summerland BC (1997), and Snohomish and
Skagit Counties WA (1991); corn, Zea Mays L. (Gramineae) at Vancouver BC (1992); and
greenhouse cucumber Cucumis sativus L. (Cucurbitaceae) at Abbotsford (1995) and Cobble
Hill BC (1996). The beetles were maintained in 70% ethanol and the genitalia of the males
were dissected and mounted on slides. Identifications were based on Gordon (1985) using a
phase contrast microscope, and confirmed by Y. Bousquet at the Eastern Cereal and Oilseed
Research Centre, Ottawa. Forty-three male beetles collected from the Pacific Northwest were
all S. punctillum. Five male beetles collected from Summerland, BC were all S. punctum
picipes.

Stethorus punctillum was the only species found in collections from agricultural habitats
in the Pacific Northwest during a 12-year period. Although the sample size was not large (43
males), the samples were consistent among years and locations, suggesting that S. punctillum
is the dominant species. Continued inundative and augmentative releases of S. punctillum will
probably not alter the species complex in the Lower Fraser Valley. Stethorus punctillum may
have displaced S. punctum picipes in agricultural areas of Vancouver Island and the Lower
Fraser Valley prior to commercial releases. Putman (1955) described a rapid displacement in
southern Ontario between 1930 and 1940. No mass reared beetles have as yet been distributed
to growers in the Okanagan Valley, although 500 have been sold to a grower further east, at
Winlaw (Bob Macadam, Westgro Sales Inc., personal communication). Further work is
needed to determine the nature of the complex in the Okanagan.

ACKNOWLEDGEMENTS

The authors thank B. Congdon and D. Gillespie for additional samples of Stethorus, and Y.
Bousquet for confirming our identifications. The work was funded in part by the BC
Raspberry Growers' Association, the BC Greenhouse Vegetable Research Council and the
Federal MII and PERD programs. Pacific Agri-Food Research Centre, Agassiz contribution
no. 677.

82 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

REFERENCES

Brown, W.J. 1950. The extralimital distribution of some species of Coleoptera. The Canadian Entomologist 82:
197-205.

Gordon, R.D. 1985. The Coccinellidae (Coleoptera) of America North of Mexico. Journal of the New York
Entomological Society. 912 pp.

Putman, W.L. 1955. Bionomics of Stethorus punctillum Weise (Coleoptera: Coccinellidae) in Ontario. The
Canadian Entomologist 87: 9-33.

Raworth, D.A. 2001. Development, larval voracity, and greenhouse releases of Stethorus punctillum
(Coleoptera: Coccinellidae). The Canadian Entomologist 133: 721-724. [Note Erratum, The Canadian
Entomologist 133: 895.]

Tonks, N.V. 1953. Annotated list of insects and mites collected on brambles in the lower Fraser Valley, British
Columbia. Proceedings of the Entomological Society of British Columbia 49: 27-29.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 83

Species of Alaska Scolytidae:
Distribution, hosts, and historical review

MALCOLM M. FURNISS
UNIVERSITY OF IDAHO, MOSCOW, ID 83843.

EDWARD H. HOLSTEN

U.S. DEPARTMENT OF AGRICULTURE, FOREST SERVICE, FOREST HEALTH
PROTECTION, ANCHORAGE, AK 99503

MARK E. SCHULTZ

U.S. DEPARTMENT OF AGRICULTURE, FOREST SERVICE, FOREST HEALTH
PROTECTION, JUNEAU, AK 99508

ABSTRACT

The species of Scolytidae in Alaska have not been compiled in recent times although
many of them are included in earlier broader works. The authors of these works are
summarized in a brief history of forest entomologists in Alaska. Fifty-four species of
Alaskan Scolytidae are listed (23 species in the subfamily Hylesininae and 31 in the
subfamily Scolytinae) belonging to 24 genera (11 in Hylesininae and 13 in Scolytinae).
They infest 15 species of trees and shrubs of which 10 are conifers (host to 48 species of
Scolytidae) and 5 are angiosperms (host to 6 species of Scolytidae). Fifty species are
bark beetles that inhabit phloem and four species are ambrosia beetles that live in
sapwood. All are species native to Alaska.

Keywords: Bark beetles, Scolytidae, conifers, angiosperms, Alaska

INTRODUCTION

The biographic regions of Alaska reflect the great expanse of that state, extending over
a wide range of latitude, longitude, and elevation. In broad terms, these regions are
characterized by a relatively mild and moist coastal climate, particularly in the southeast
and coastal south-central areas; an extensive drier, colder climate in interior and northern
Alaska; and numerous mountain ranges that attain the highest elevation on the continent.

Except for willows (Salix spp.), the species of Alaskan trees are somewhat more
diverse in the coastal environment. Because species of Scolytidae are host-specific to a
marked degree, the extent of their diversity generally reflects that of woody plant species
that are needed and available for their existence. Warmer climate also enhances scolytid
diversity by exerting less selective pressure as is evident in tropical regions. Thus, more
than half of the known Alaskan scolytid fauna occur in rather close proximity to the coast.
Nonetheless, white spruce, Picea glauca (Moench) Voss, a tree that occurs widely in the
interior, is host of more species of scolytids than any other tree species in Alaska (Table
1).

The species of Scolytidae in Alaska have not been compiled in recent times although
many of them are included in the broader works of Bright (1976, 1981), and Wood (1982).
The first record of Alaskan Scolytidae (12 species) was reported by C. G. Mannerheim in
the period 1843-1853 (Hamilton 1894). Beckwith (1972) listed 17 species infesting spruce
(Picea spp.), two of which have been placed in synonymy by Wood (1982). Werner and

84 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Holsten (1984) listed 29 species attracted to pheromone baited traps and spruce trap trees.
Werner (2002) lists 23 species in relation to various environmental disturbances in interior
Alaska white spruce forests. Gara et al. (1995) list 12 scolytid species associated with the
spruce beetle, Dendroctonus rufipennis (Kirby), in white and Lutz spruce (Picea x lutzii
Little). The biology of the more common scolytids of Alaska is described by Holsten et al.
(2001). Records of some secondary scolytids are obscure, being mentioned rather
incidentally in studies of more prominent primary species (e.g., Baker et al. 1977; Furniss
et al. 1976, 1979; Werner 1986; Werner et al. 1977, 1981). This work draws together
published and unpublished records since 1946. For perspective, a brief history is presented
of the location and staffing of forest entomologists in Alaska.

Table 1

Hosts of Alaskan Scolytidae.

Species of
Scolytidae

Chamaecyparis nootkatensis
Tsuga heterophylla
Populus tremuloides

Tsuga mertensiana
Betula papyrifera

Picea mariana
Picea sitchensis
Pinus contorta
Thuja plicata
Salix alaxensis
Salix scouleriana

Larix laricina

Picea glauca

Picea x lutzii

H

Alniphagus aspericollis heal de aah |
| Carphoborus andersoni_ | ae ee ee ee ee ee
| Carphoboruscarri_ | [as Ge sa eH fe a
| Carphoborus intermedius || x i ee ee ee
| Dendroctonus punctatus | ae a Ga [es [ES]
| Dendroctonus rufipennis | CX | i ec a ee A P
| Dendroctonus simplex || FG (ii Fae a De Ja P|
| Hylastes nigrinus | Pe a Fe ee ee ev

Hylurgops rugipennis rugipennis Laie xX | Xx ie ae ae | |
| Phioeosinus cupressi dT x] | Txt | | | | | | | | TT
aehloeosiniths pint S25 PE Le
| Phloeosinus punctatus | | Ct CT UT | TT x} | fT | TT
| Phloeotribustecontei S| SL x] | | | | TE TUT | | TT |
Pi ee asa esas
| Polygraphus rufipennis | ST xt xt [xix] | | | | | | ft ft
| Polygraphus convexifrons | =| xt x] | | | | | ft | | fT | ft
| Pseudohylesinus granulaus | | | | | | TT TT TT TT
| Pseudohylesinus sericeus | =| OT | ct CT TT TT TT | TT
| Pseudohylesinus sitchensis | | | Txt xt [ | | ft | | | | tt
| Pseudohylesinusisugae | S| OT | oT TT | xt TT TT
| Scierusannectens | txt | TT TT TE TT TP
| Scierus pubescens | CT xt TT TT TT

Xylechinus montanus aes eae eee

Scolytinae

| Cryphatus pubescens | ot xt | | | | | | | tT | | |
| Cryphalus ruficollis ruficollis | [x] | | | [ [| | | | fT | fT 7 Tt |
| Crypturgus borealis | CT xt TT xt | fT fT tT ft tt tT Tt
sir pam
| Dryocoetes affaber ST CT xt x] [xt x] | [ [| [ | | | Tt ft
Sr

Dryocoetes caryi eas ee Fa De el

Ips borealis borealis anna ae eee eee eee

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 85

Species of
Scolytidae

| Ipsconcinnus
| Ipsperturbatus |
[ipspint
Pai a a ars
| Lymantor alaskanus |
| Orthotomicus caelatus |
Pityophthorus bassetti |

| Pityophthorus carinulatus Swe |
| Pityophthorus murrayanae |__|
| Pityophthorus nitidulus |
| Pityophthorus nitidus |
| Pityophthorus opaculus |
| Pityophthorus recens |
| Pityophthorus tuberculatus |
| Procryphalus mucronatus |
irocrphales wean, J
a

a

ea

a

[4

a

ee Picea mariana
| | |x| Picea sitchensis

| | Picea glauca
PLT ET LETT eT LT de] Pinus contort

Cl aie

HISTORY OF BARK BEETLE SURVEYS IN ALASKA

After Mannerheim’s early work (Hamilton 1894), infestations of Alaskan bark beetles
were reported only sporadically beginning in 1922 (Zogas and Holsten 2002). Forest
entomologists were not directly involved until 1946 when Robert L. Furniss of the
Portland, Oregon, Forest Insect Laboratory was requested by regional forester, B. Frank
Heintzleman, to investigate dying Sitka spruce, Picea sitchensis (Bong.) Carr., on the
Tongass and Chugach National Forests (Furniss 1946, 1948, 1950; Furniss and Jones
1946). William F. McCambridge was the first forest entomologist to be stationed in Alaska
(Juneau, 1952-1956). He was replaced by George L. Downing; others have followed there
to this time. Since 1976, Forest Service Alaska Region entomologists have also been
stationed at Anchorage but mainly concerned with spruce beetle infestations on the Kenai
Peninsula. Richard A. (Skeeter) Werner was stationed at the Institute of Northern Forestry
at Fairbanks from 1974 until that facility closed in 1996. During that time, the Alaska State
Division of Forestry hired entomologist Roger E. Burnside who is stationed at Anchorage.

Pseudips (=Ips) mexicanus

PEt TET TT ETT EEE TTT I cers caring

MATERIALS AND METHODS

Records of Alaskan species of Scolytidae were obtained from specimens available to
us, from reviewers credited in Acknowledgments, and from the literature, including reports
listed by Zogas and Holsten (2002). Such reports are on file at USDA, Forest Service,
Alaska Region, Anchorage. Sources of specimens, locality information, and host data were

86 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

primarily the Forest Service Alaska Region collections located at Anchorage and Juneau,
and the senior author’s collections. His specimens were collected during various studies
and recreational visits since 1967 throughout all regions of the state where scolytids occur.
Specimens are deposited at the USDA Forest Service, Alaska Region, in Anchorage and
Juneau; in the W.F. Barr Entomological Museum, University of Idaho, Moscow, ID;
Canadian National Collection of Insects, Arachnids and Nematodes (CNC), Ottawa;
Brigham Young University, Provo, UT; Oregon State University, Corvallis, OR; and
USDA, Agriculture Research Service, Beltsville, MD.

RESULTS AND DISCUSSION

Fifty-four species of Alaskan Scolytidae are listed here (23 species in the subfamily
Hylesininae and 31 in the subfamily Scolytinae). They belong to 24 genera (11 in
Hylesininae and 13 in Scolytinae). They infest 15 species of Alaskan trees and shrubs of
which 10 are conifers (host to 48 species of Scolytidae) and five are angiosperms (host to
six species of Scolytidae). Fifty species are bark beetles that inhabit phloem and four
species are ambrosia beetles that live in sapwood. All are native to Alaska. Four species
new to Alaska are: Phloeotribus lecontei Schedl, Cryphalus pubescens Hopk.,
Pityophthorus recens Bright, and Trypodendron betulae Sw.

The following list is ordered alphabetically within each subfamily. Distributions
include place names that may be found on road maps and in Orth (1967). For some species
such as the spruce beetle we have found it more expedient to describe the extent of their
occurrence in general terms. Where relevant, a remark is inserted to call attention to some
aspect of a record. Such is the case with Lymantor alaskanus Wood for which the host is
unknown. In this way we hope to stimulate additional collecting and study. Also, selected
references are provided where available.

Hylesininae

Alniphagus aspericollis (LeC.)
Distribution: Hollis, Prince of Wales Is.
Host: Alnus sp.
Carphoborus andersoni (Sw.)
Distribution: Bonanza Cr. (nr. Fairbanks), Copper Center, Fort Yukon, John R..,
Salmon R., Sheenjek R.
Host: Picea glauca.
Carphoborus carri Sw.
Distribution: Bonanza Cr. (nr. Fairbanks), John R.
Host: Picea glauca.
Carphoborus intermedius Wood
Distribution: Bonanza Cr. (nr. Fairbanks).
Host: Picea glauca.
Dendroctonus punctatus LeC.
Distribution: Interior drainages. (Furniss 1995)
Host: In the base of living Picea glauca.
Dendroctonus rufipennis (Kirby)
Distribution: Throughout the distribution of its hosts. This is by far the most
economically important scolytid in Alaska.
Hosts: Picea sitchensis, P. glauca, P. x lutzii and P. mariana.
Dendroctonus simplex LeC.
Distribution: Interior, throughout distribution of its host in the middle Yukon R.
and in the Tanana R. drainage.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 87

Host: Larix laricina.
Hylastes nigrinus (Mannerheim)
Distribution: Sitka Is.
Remarks: Holotype female in Helsinki Museum (Wood 1982), host not specified.
Hylurgops rugipennis rugipennis (Mannerheim)
Distribution: Farragut Bay, Ft. Yukon, Juneau, Kethikan, Kodiak Is., Loring,
Sitka, Skagway.
Hosts: Picea sitchensis, Pinus contorta.
Hylurgops subcostulatus subcostulatus (Mannerheim)
Distribution: Kenai Penin.
Remarks: Holotype presumably lost (Wood 1982).
Phloeosinus cupressi (Hopk.)
Distribution: Chichagof Is., Juneau, Petersburg, Wrangell.
Host: Chamaecyparis nootkatensis.
Phloeosinus pini Sw.
Distribution: Bonanza Cr. (nr. Fairbanks), Chichagof Is., Eagle, Fairbanks, John
R.
Host: Picea glauca, Pinus contorta.
Phloeosinus punctatus LeConte
Distribution: Edna Bay, Hollis, Kosciusko Is., Petersburg.
Host: Thuja plicata.
Phloeotribus lecontei Schedl
Distribution: Sheenjek R., Bonanza Cr. (nr. Fairbanks).
Host: Picea glauca.
Remarks: New state record.
Phloeotribus piceae Sw.
Distribution: Bonanza Cr. (nr. Fairbanks), Eagle R., Hope, John R., Summit L.
(Kenai Penin.).
Host: Picea glauca.
Polygraphus convexifrons Wood
Distribution: Arviriag, Bonanza Cr., John R., Matanuska, McKinley N.P., Noatak
R., Quartz Cr. and Summit L. (Kenai Penin.), Walker L.
Hosts: Picea glauca, P. x lutzii.
Polygraphus rufipennis (Kby.)
Distribution: Anchorage, Bonanza Cr. (nr. Fairbanks), Bullfrog Is. (Yukon R.),
Chandalar, Douglas, Dry Gulch and Cooper Landing (Kenai Penin.),
Fairbanks, Ft. Yukon, Juneau, Klutina L., Homer, Hughes, John R.,
Kasilof, Kenai National Moose Range, Lawing (Kenai Lk.), McKinley,
Mile 34 Seward Hwy., Montana Cr. (Mat-Su Valley), Moose Pass,
Northway Jct., Russian R. (Kenai Penin.), Sheenjek R., Skilak L.,
Summit L.(Kenai Penin.), Talkeetna.
Hosts: Picea glauca, P. x lutzii, P. mariana, P. sitchensis, Pinus contorta.
Pseudohylesinus granulatus (LeConte)
Distribution: “Nauacin” (Wood 1982).
Host: Not specified, presumably Tsuga heterophylla.
Pseudohylesinus sericeus (Mannerheim)
Distribution: “Sitka” (Wood 1982).
Host: Not specified, presumably Tsuga heterophylla.
Pseudohylesinus sitchensis Sw.
Distribution: Kodiak Is., Juneau, Prince of Wales Is.
Host: Picea sitchensis.

88 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Pseudohylesinus tsugae (Sw.)
Distribution: Chichagof Is., Dry Gulch (Kenai Penin.), Echo Cove (nr. Juneau),
Glacier Highway, Hollis, Juneau, Ketchikan.
Host: Tsuga heterophylla
Scierus annectens LeC.
Distribution: Bonanza Cr. (nr. Fairbanks), Bullfrog Is., Fairbanks, John R.,
Kasilof, Sumit L. (Kenai Penin.), Klehini R., (N of Haines), Willow.
Host: Picea glauca
Scierus pubescens Sw.
Distribution: Bonanza Cr. (nr. Fairbanks), Summit L. and Swanson R. road
(Kenai Penin.).
Host: Picea glauca
Xylechinus montanus Blackm.
Distribution: Bonanza Cr. (nr. Fairbanks), Swanson River road, Summit L. (Kenai
Penin.).
Host: Picea glauca.

Scolytinae

Cryphalus pubescens Hopk.
Distribution: Iliamna L.
Host: Picea glauca.
Remarks: New state record.
Cryphalus ruficollis Hopk.
Distribution: Bonanza Cr. (nr. Fairbanks), Charley R., John R., Summit L. (Kenai
Penin.).Host: Picea glauca.
Crypturgus borealis Sw.
Distribution: Bonanza Cr. (nr. Fairbanks), Charley R., Coleen R., Echo Cove (nr.
Juneau), Fairbanks, John R.
Hosts: Picea glauca, P. sitchensis.
Dolurgus pumilus (Mann.)
Distribution: Chichagof Is., Echo Cove and Gold Cr. (nr. Juneau), Hollis, Juneau,
Montana Cr. (SE AK), Naukati Bay, Sitka Island, Yakutat.
Hosts: Picea sitchensis, Tsuga heterophylla.
Dryocoetes affaber (Mann.)
Distribution: Arviriag (nr. Noatak R.), Bonanza Cr., Coleen R., Cooper Landing,
Douglas, Dry Gulch (Kenai Penin.), Echo Cove (nr. Juneau), Fairbanks,
Ft. Yukon, Hollis, Homer, John R, Juneau, Kandik R., Kasilof, Kenai
National Moose Range, McKinley N.P., Mentasta, Mile 24, Mile 50,
Montana Cr. (Mat-Su Valley), Moose Pass, Naukati Bay, Noatak R.,
Northway Junction, Patterson, Russian R., Sheenjek R., Summit L.
(Kenai Penin.), Talkeetna, Venetie, Walker L., Yakutat.
Hosts: Picea glauca, P. x lutzii, P. sitchensis, Pinus contorta.
Dryocoetes autographus (Ratz.)
Distribution: Anchorage, Bonanza Ck. (nr. Fairbanks), Chandalar, Cooper
Landing and Dry Gulch (Kenai Penin.), Edna Bay, Glacier Bay, Hollis,
John R., Kandik R., Kenai National Moose Range, Ketchikan, M.34
Seward Hwy, Naukati Bay, Northway Junction, Patterson, Russian R.,
Summit L. and Sunrise (Kenai Penin.), Yakutat.
Hosts: Picea glauca, P. sitchensis, Tsuga heterophylla.
Dryocoetes caryi Hopk.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 89

Distribution: Homer, Mile 34 Seward Hwy, Moose Pass and Russian R (Kenai
Penin.).
Host: Picea x lutzit.
Ips borealis borealis Sw.
Distribution: Bonanza Cr. (nr. Fairbanks), Cooper Landing and Dry Gulch (Kenai
Penin.), John R., Kenai National Moose Range, Nanana Hwy., Summit
L.(Kenai Penin.), Wasky (nr. Togiak L.).
Hosts: Picea glauca, P. x lutzit.
Ips concinnus (Mann.)
Distribution: Echo Cove (nr. Juneau), Edna Bay, Hollis, Homer, Hoonah, Juneau,
Kodiak Is., Mile 20, Montana Cr. (nr. Juneau), Rogers Point, Seagull
Creek, Granite Cr. (Kenai Penin.), Seward.
Host: Picea sitchensis, P. x lutzii.
Ips perturbatus (Eichh.)
Distribution: Arctic Village, Arviriaq (nr. Noatak R.), Bonanza Cr. (nr.
Fairbanks), Chandalar, Circle City, Coleen R., Cooper Landing and Dry
Gulch (Kenai Penin.), Fairbanks, Ft.Yukon, Hope, Hughes, John R..,
Kandik R., Kasilof, Kenai National Moose Range, Klutina L., Lawing
(Kenai Penin.), Mentasta, McKinley River drainage (24 mi E of Lake
Minchumina), Mile 50, Northway Jct., Noatak R., Pah R., Porcupine R.,
Richardson Highway, Russian R. (Kenai Penin.), Sheenjek R. Summit L.
(Kenai Penin.), Venetie, Walker L.
Host: Picea glauca, P. x lutzii.
Ips pini (Say)
Distribution: Douglas Is., Ketchikan, Petersberg.
Host: Pinus contorta.
Ips tridens (Mann.)
Distribution: Bonanza Cr. (nr. Fairbanks), Cooper Landing and Dry Gulch (Kenai
Penin.), Echo Cr. (nr. Juneau), Richardson Highway, Rodgers Point
(Chichagof Is.), Seward, Cove (nr. Juneau), Fick Cove (Chichagof Is.),
Hollis, Homer, Hood Bay, John R., Juneau, McKinley N.P., Mile 34
Seward Hwy., Montana Cr. (Mat-Su Valley), Moose Pass and Quartz Cr.
(Kenai Penin.), Sheenjek R., Sitka Is., Summit L. and Sunrise (Kenai
Penin.), Willow.
Hosts: Picea glauca, P. x lutzii, P. sitchensis.
Remark: Two subspecies, /. tridens tridens (Mann.) and /. tridens engelmanni
Sw., occur in Alaska (Wood 1982); however, the distributions given
therein appear to overlap and further collecting is needed to clarify this
point. According to Wood (1982), females of the latter subspecies tend
to have a more protuberant and generally more pubescent frons.
Lymantor alaskanus Wood
Distribution: Bonanza Creek, 42 km W of Fairbanks.
Hosts: Unknown
Remark: This species was described (Wood 1982) from 19 specimens collected
18-VII-1978 on a sticky trap baited with Ipsenol (pheromone of /ps spp.)
and alpha-pinene (resin constituent). Its host is unknown. The other
North American species, L. decipiens LeC., infests dead, dry, branches
of living maple trees. Its gallery penetrates the sapwood and is associated
with black stain (Wood 1982).
Orthotomicus caelatus (Eichh.)

90 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Distribution: Bonanza Creek (nr. Fairbanks), Circle City, Douglas, John R.,
Juneau, Kandik R., McKinley River drainage (24 mi E of Lake
Minchumina), Mile 7, Petersburg, Porcupine R.
Hosts: Picea glauca, P. sitchensis, Pinus contorta.
Pityophthorus bassetti Blkm.
Distribution: Bonanza Cr. (nr. Fairbanks), Fairbanks.
Host: Picea glauca.
Pityophthorus murrayanae murrayanae Blackm.
Distribution: Bonanza Cr. (nr. Fairbanks), Hope, Mile 295 Richardson Hwy.,
Sheenjek R., Summit L. (Kenai Penin.).
Hosts: Picea glauca, P. x lutzii.
Pityophthorus nitidulus (Mann.)
Distribution: Anchorage, Bonanza Cr. (nr. Fairbanks), Cooper Landing and Dry
Gulch (Kenai Penin.), Eagle R., Hollis, Homer, Iliamna L., Juneau,
Kenai National Moose Range, Kodiak Is., Lawing, Moose Pass, Quartz
Cr. (Kenai Penin.), Seward, Skilak L., Summit L. (Kenai Penin.).
Hosts: Picea glauca, P. x lutzii, P. mariana, P. sitchensis.
Pityophthorus nitidus Sw.
Distribution: Bonanza Cr. (nr. Fairbanks), Fort Yukon, Hope, John R., Lawing
(Kenai Penin.), Summit L. (Kenai Penin.).
Hosts: Picea glauca, P. x lutzii.
Pityophthorus opaculus LeC.
Distribution: Bonanza Creek (nr. Fairbanks), Fairbanks, Hope, John R., Summit
L. (Kenai Penin.).
Hosts: Picea glauca, P. x lutzii.
Pityophthorus recens Bright
Distribution: Granite Cr. (Kenai Penin.).
Host: Picea x lutzii.
Remark: New state record. Hundreds of specimens were captured in funnel traps
during field tests of Jps perturbatus pheromones.
Pityophthorus tuberculatus Eichh.
Distribution: Haines, Juneau, Skagway (Bright 1981).
Host: Pinus contorta.
Procryphalus mucronatus (LeConte)
Distribution: Hope.
Host: Unknown.
Procryphalus utahensis (Hopk.)
Distribution: Bonanza Creek (nr. Fairbanks), Galena.
Hosts: Salix scouleriana, Salix sp.
Pseudips (=Ips) mexicanus (Hopk.)
Distribution: Douglas Is., Juneau.
Host: Pinus contorta.
Remark: Cognato (2000) transferred /ps mexicanus to the new genus Pseudips.
Scolytus piceae (Sw.)
Distribution: Bonanza Cr. (nr. Fairbanks), Fairbanks, Ft.Yukon, John R.,
Northway Junction, Walker L.
Hosts: Picea glauca, Larix laricina.
Trypodendron betulae Sw.
Distribution: Anchorage, Chichagof Is., Fairbanks.
Host: Alnus sinuata, Betula papyrifera.
Remarks: New state record.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 91

Trypodendron lineatum (Oliv.)

Distribution: Admiralty Is., Auke Bay, Bonanza Cr. (nr. Fairbanks), Chichagof
Is., Cooper Landing, Cordova, Dry Gulch (Kenai Penin.), Edna Bay,
Fairbanks, Hollis, Homer, John R., Kenai National Moose Range,
Petersburg, Russian R. and Summit L. (Kenai Penin.), White Bay,
Yakutat.

Hosts: Picea glauca, P. sitchensis, Pinus contorta, Tsuga heterophylla.

Trypodendron retusum (LeC.)

Distribution: Anchorage, Bonanza Cr. (nr. Fairbanks), Russian R., Summit L. and
Sunrise (Kenai Penin.), Willow.

Hosts: Betula papyrifera, Populus tremuloides.

Trypodendron rufitarsus (Kby.)
Distribution: Russian R. (Kenai Penin.), Talkeetna, Sunrise (Kenai Penin.).
Hosts: Picea glauca, Tsuga mertensiana.

Trypophloeus striatulus (Mann.)

Distribution: Interior, coastal NW, N Slope; wherever its host occurs.

Host: Salix alaxensis.

Remarks: Collected at Shublik Spring, Canning River, Lat. N 69° 30', possibly the
farthest north of any scolytid. A fungus, Cytospora sp. is universally
present in infested stems indicating a possible symbiotic relationship.
The same stem is infested by several generations of the beetle before
killing it (Furniss 1997).

ACKNOWLEDGEMENTS

We thank Kenneth P. Zogas, USDA Forest Service, Forest Health Protection,
Anchorage, Alaska for curating the Alaska insect collection for the last twenty years and
for compiling the list of accessions used herein. The manuscript was reviewed by Donald
E Bright, Jr. and Stephen L. Wood who also identified most of the bark beetles recorded
here. Other reviewers who contributed to improvement of the paper were Roger E.
Burnside and Richard A.Werner.

REFERENCES

Baker, B.H., B.B. Hostetler, and M.M. Furniss. 1977. Response of eastern larch beetle (Coleoptera:
Scolytidae) in Alaska to its natural attractant and to Douglas-fir beetle pheromones. The Canadian
Entomologist 109: 289-294.

Beckwith, R.C. 1972. Key to adult bark beetles commonly associated with white spruce stands in interior
Alaska. U.S. Department of Agriculture, Forest Service Research Note PNW-189. Portland, OR. 6 p.

Bright, D.E., Jr. 1976. The bark beetles of Canada and Alaska. The insects and arachnids of Canada, Part
2. Biosystematics Research Institute, Research Branch, Canada Department of Agriculture Publication
1576. Ottawa.

Bright, D.E., Jr. 1981. Taxonomic monograph of the genus Pityophthorus Eichhoff in North and Central
America (Coleoptera: Scolytidae). The Canadian Entomologist Memoir No. 118.

Cognato, A.J. 2000. Phylogenetic analysis reveals new genus of Ipimi bark beetle (Scolytidae). Annals of
the Entomology Society of America 93: 362-366.

Furniss, M.M. 1995. Biology of Dendroctonus punctatus (Coleoptera: Scolytidae). Annals of the
Entomology Society of America 88:173-182.

Furniss, M.M. 1997. Biology of the willow bark beetle, Trypophioeus striatulus (Mann.), in feltleaf willow
in interior Alaska. U.S. Department of Agriculture Forest Service, Alaska Region, Progress report (in
cooperation with E.H. Holsten). (On file, Anchorage).

Furniss, M.M., B.H. Baker, and B.B. Hostetler. 1976. Aggregation of spruce beetles (Coleoptera) to
seudenol and repression of attraction by methylcyclohexenone in Alaska. The Canadian Entomologist
108:1297-1302.

O72 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Furniss, M.M., B.H. Baker, R. A. Werner, and L.C. Yarger. 1979. Characteristics of spruce beetle
(Coleoptera) infestation in felled white spruce in Alaska. The Canadian Entomologist 111:1355-1360.

Furniss, R.L. 1946. Memorandum regarding an outbreak of Dendroctonus on Kosciusko Island., U.S.
Department of Agriculture, Bureau of Entomology, Plant Quarantine, Forest Insect Laboratory,
Portland, Oregon. (Cited in Zogas and Holsten 2002).

Furniss, R.L. 1948. [Afognak Island outbreak, summary]. Letter to Regional Forester, Alaska Region. U.S.
Department of Agriculture, Bureau of Entomology, Plant Quarantine, Forest Insect Laboratory,
Portland, Oregon. (Cited in Zogas and Holsten 2002).

Furniss, R.L. 1950. Forest insect situation in Alaska. U.S. Department of Agriculture, Bureau of
Entomology, Plant Quarantine, Forest Insect Laboratory, Portland, Oregon. (Cited in Zogas and
Holsten 2002).

Furniss, R.L. and I.H. Jones. 1946. A second report concerning the bark beetle outbreak on Kosciusko
Island. U.S. Department of Agriculture, Bureau of Entomology, Plant Quarantine, Forest Insect
Laboratory, Portland, Oregon. (Cited in Zogas and Holsten 2002).

Gara, R.I., R.A. Werner, M.C. Whitmore, and E.H. Holsten. 1995. Arthropod associates of the spruce
beetle Dendroctonus rufipennis (Kirby) (Coleoptera, Scolytidae) in spruce stands of south-central and
interior Alaska. Journal of Applied Entomology 119: 585-590.

Hamilton, M.D. 1894. Catalogue of the Coleoptera of Alaska, with the synonymy and distribution. Trans
American Entomology Society Vol. XXI.

Holsten, E.H., P.E. Hennon, L. Trummer, and M. Schultz. 2001. Insects and diseases of Alaskan forests,
U.S. Department of Agriculture, Forest Service Alaska Region R10-TP-87.

Orth, D.J. 1967. Dictionary of Alaska place names. Geology Survey Professional Paper 567. U.S.
Government Printing Office, Washington, D.C.

Werner, R.A. 1986. The eastern larch beetle in Alaska. U.S. Department of Agriculture, Forest Service
Research Paper PNW-357.

Werner, R A. 2002. Effect of ecosystem disturbance on diversity of bark and wood-boring beetles
(Coleoptera: Scolytidae, Buprestidae, Cerambycidae) in white spruce ecosystems in Alaska. U.S.
Department of Agriculture, Forest Service General Technical Report. PNW-GTR-546.

Werner, R.A., H.H. Baker, and P.A. Rush. 1977. The spruce beetle in white spruce forests of Alaska. U.S.
Department of Agriculture, Forest Service General Technical Report PNW-GTR-61.

Werner, R A., M M. Furniss, L C. Yarger, and T. Ward. 1981. The effect of methylcyclohexenone and
trans-verbenol on response of eastern larch beetle, Dendroctonus simplex Lec. (Coleoptera:
Scolytidae) to its natural attractant. U.S. Department of Agriculture, Forest Service Research Note
PNW-371.

Werner, R.A. and E.H. Holsten. 1984. Scolytidae associated with felled white spruce in Alaska. The
Canadian Entomologist 116: 465-471.

Wood, S.L. 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a
taxonomic monograph. Great Basin Naturalist Memoir No. 6.

Zogas, K. and E H. Holsten. 2002. Bibliography. Alaska Region Forest Health Protection, 1919 — 2001.
Region 10, Anchorage, Aik. U.S. Department of Agriculture, Forest Service Tech. Rep. R10 —TP-107.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 93

Searching behaviour of 7richogramma wasps (Hymenoptera:
Trichogrammatidae) on tomato and pepper leaves

ROBERT R. MCGREGOR

DEPARTMENT OF BIOLOGY, DOUGLAS COLLEGE,
P.O. BOX 2503, NEW WESTMINSTER, B.C., V3L 5B2

RENEE P. PRASAD and DEBORAH E. HENDERSON

E.S. CROPCONSULT LTD., 3041 33RD AVE WEST, VANCOUVER, B.C., V6N 2G6

ABSTRACT

Walking speeds of both Trichogramma brassicae and T. sibericum were substantially
lower on tomato than on pepper leaf disks. The difference may be due to the presence of
glandular trichomes on tomato foliage. Total time spent on leaf disks during behavioural
trials was lower on tomato than on pepper leaf disks for both species of wasps. This may
indicate a higher propensity to disperse from tomato foliage than from pepper foliage.
Lower walking speeds and shorter residence times on tomato leaves could result in a
lower searching efficiency of wasps on tomato than on pepper. The subsequent efficacy
of Trichogramma for biological control of cabbage loopers in greenhouses may be lower
on tomato crops than on pepper crops.

Key words: Trichogramma brassicae, Trichogramma sibericum, Trichoplusia_ ni,
searching behaviour, glandular trichomes, egg parasitoids, biological control, greenhouse
vegetables

INTRODUCTION

Foliar pubescence on agricultural crops may interfere with the host location activities
of small entomophagous insects and mites and reduce their subsequent effectiveness for
biological control of arthropod pests (Obrycki 1986). Such effects have been documented
for a variety of taxa. The predatory mite, Phytoseiulus persimilis Athias-Henriot (Acarina:
Phytoseidae), was trapped by glandular trichomes more frequently on a tomato variety
with a high trichome density than on a relatively hairless tomato variety (van Haren et al.
1987). Walking speed and parasitism of greenhouse whitefly (7rialeurodes vaporariorum
(Westwood) (Homoptera: Aleyrodidae)) by Encarsia formosa Gahan (Hymenoptera:
Aphelinidae) was lower on cucumber varieties with higher levels of foliar pubescence (van
Lenteren et a/. 1995). Walking speeds of E. formosa recorded on pubescent tomato and
gerbera foliage were slower than those recorded on hairless sweet pepper (Sutterlin & van
Lenteren 1997). The estimated searching efficiency of Orius insidiosus (Say)
(Heteroptera: Anthocoridae) was lowest on pubescent tomato foliage compared to corn and
bean foliage (Coll et al. 1997). Higher prey handling times were recorded for Podisus
nigrispinus (Dallas) (Heteroptera: Pentatomidae) on tomato plants compared to those
recorded on eggplant or pepper plants (De Clerq et a/. 2000). In this study, we compare the
searching behaviour of 7richogramma brassicae Bezdenko and T. sibericum Sorokina
(Hymenoptera: Trichogrammatidae) on tomato and pepper leaf disks in the laboratory.

Wasps of the genus Trichogramma are egg parasitoids that have been used widely for
biological control of Lepidopteran pests (Li 1994). Searching behaviour and parasitism of
hosts by a number of 7richogramma species are impeded by high levels of foliar
pubescence. The time required to walk across leaf disks, the percentage of wasps initiating

94 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

flight before crossing leaf disks, and the number trapped by trichomes were all higher for
T. pretiosum Riley on tomato varieties with higher densities of glandular trichomes and
containing either 2-tridecanone or 2-undecanone (Kashyap ef al. 1991). In a field study,
the highest levels of parasitism of Heliothis spp. eggs (Lepidoptera: Noctuidae) by T.
pretiosum and T. exiguum Pinto & Platner were observed on tomato varieties with the
lowest density of glandular trichomes (Kauffman & Kennedy 1989). Walking speeds of T.
exiguum were lowest on highly-pubescent mullein, intermediate on intermediately-
pubescent tomato, and highest on relatively hairless maize and soybean (Keller 1987).
Parasitism of Heliocoverpa armigera (Hiibner) (Lepidoptera: Noctuidae) eggs by T.
chilonis Ishii on pigeonpea varies widely across plant parts depending on the presence of
trichomes. Parasitism is much lower on pods and calyxes that have a higher trichome
density than on leaves (Romeis et a/. 1998; 1999).

T. brassicae are released in British Columbia vegetable greenhouses for biological
control of cabbage loopers (7richoplusia ni (Huebner) (Lepidoptera: Noctuidae)) (Portree
1993). T. sibericum has been evaluated for management of cabbage loopers in vegetable
greenhouses (E.S. Cropconsult Ltd., unpublished report). Although Trichogramma wasps
are released in both tomato and pepper crops, no previous comparisons of searching
behaviour on tomato and pepper foliage have been done. Previous work has shown that
yellow sticky traps (that are used to monitor dispersal by 7. brassicae) capture fewer
wasps in tomato greenhouses than in pepper greenhouses (E.S. Cropconsult Ltd.,
unpublished data). In addition, numerous dead wasps have been found near release sites
on tomato stems, presumably trapped by glandular trichomes (Prasad, personal
observation). Both of these observations indicate that Trichogramma wasps may be less
efficient for biological control of cabbage loopers on tomato than on pepper crops.

For the purposes of this paper, searching behaviour in Trichogramma is considered to
be all activities that facilitate the location of suitable host eggs including walking on and
examining host-free leaf surfaces. Here, we report measurements of walking speed on,
time elapsed before exiting from, and proportion of time spent searching on excised
tomato and pepper leaf disks in the laboratory by T. brassicae and T. sibericum females.
Our objective was to identify differences in the behaviour of wasps between these two
plant species caused by the presence of glandular trichomes on tomato leaves.

MATERIALS AND METHODS

Insects. Trichogramma sibericum were reared on sterilized Ephestia kuehniella eggs.
T. sibericum females used in bioassays were between 1 and 3 d old. T. brassicae were
obtained from Beneficial Insectary (Oak Run, California) and were 3 to 5 d old when used
in trials. All wasps were provided continuously with dilute honey (50% by volume)
applied as droplets to the inner surface of holding containers.

Bioassay arenas.Tomato and pepper leaves of unknown variety were collected from
commercial greenhouses, wrapped in damp paper towelling, stored in Zip-lock® bags at 5°
C, and used in trials within 5 d of collection. Arenas for measurement of searching
behaviour consisted of leaf disks (2 cm in diameter) cut with a cork borer from tomato or
pepper leaves and attached (underside up) to the bottom half of 9 cm plastic petri-dishes
using double-sided tape. Behavioural observation and video recording were facilitated by
confining wasps to this simple two-dimensional arena in the absence of host eggs.
Trichogramma females search for hosts on both upper and lower leaf surfaces on entire
plants (McGregor, personal observation). Here, we assume that the behaviour wasps
display in this simple bioassay is analogous to behaviour displayed on whole plants in the
field. The underside of leaves was used for trials because it has a higher density of
trichomes on tomato leaves than the upper surface. Ten arenas (five tomato and five
pepper) were prepared at a time for use in bioassays. After ten trials were completed, ten

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 5)

more arenas were prepared. This prevented leaf disks from deteriorating before use in
bioassays.

General bioassay procedure. For each trial, a female wasp was released in the center of
a tomato or pepper leaf disk from the tip of a fine paintbrush. The wasp was videotaped
using a Panasonic WV-CD110 camera fitted with a Tamron 72mm camera lens and run by
a Panasonic WV-PS10 drive unit., The position of the camera and zoom lens were adjusted
until the two-cm leaf disk filled most of the screen on a video monitor. Wasps were
videotaped for 5 min or until they flew or walked off the leaf disk. All trials were run
under ambient conditions in the laboratory (temperature: 21-25° C; relative humidity: 37-
65%). The trial area was illuminated from above with a single incandescent bulb (60
watt). Trials were alternated between plant species. Twenty trials were completed for
each wasp species on each plant species (ie. total of 80 trials).

T. sibericum bioassays. All T. sibericum bioassays were run as above. However, trials
that lasted less than 60 s were discarded. This was done to maximize the number of
walking tracks that could be analysed for each wasp in order to increase the precision of
estimates of walking speed. The first 20 trials for each plant species that lasted more than
60 seconds were retained for analysis.

T. brassicae bioassays. The majority of trials for this species lasted less than 60
seconds so none was discarded. The wasps walked or flew off the leaf shortly after release
in most cases. The first 20 trials for each plant species were retained for further analysis
no matter how short in duration.

Estimation of walking speed. An acetate sheet was taped to the screen of the video
monitor. Walking tracks of Trichogramma females were traced with a felt pen on the
acetate sheet. Tracks were only traced and analysed for walking on the surface of the leaf
disk. Walking along the edge of the leaf disk was not recorded. One to five tracks of at
least 1 cm were traced for each wasp. Tracing of a particular track was stopped when the
wasp reached the edge of the leaf disk. The time a wasp spent walking along a traced path
was recorded with a stopwatch. The length of the path was measured on the acetate sheet
using a cartographer’s odometer. The odometer consists of a rotating wheel (and
associated accumulating scale) that is rolled along a curved line to measure its length in
odometer units. Odometer units were calibrated to the image from the video display by
measuring a tracing of a video clip of a centimeter ruler filmed at the same position that
bioassays were conducted. Walking speed for each wasp was calculated by dividing the
sum of the distance travelled in all measured tracks by the sum of the times recorded for
walking those tracks.

Measurement of searching time and residence time. The videotape of each trial was
viewed from the time of release of the wasp until the end of the trial. The total time each
wasp spent on the leaf disk during the trial ("residence time") was recorded using a
stopwatch (ie. the time from release until the time the wasp walked or flew off the leaf or
until the trial was stopped at 5 m). As such, residence time ranged from zero to 300 s.
Searching time was recorded on a second stopwatch as the time the wasp spent walking on
the leaf disk during the trial. Time spent resting and cleaning wings or antennae was not
included. The proportion of time spent searching during a trial was calculated by dividing
searching time by residence time.

Data analysis. Data for all trials for both Trichogramma species and both leaf species
were combined for analysis. The variables walking speed, total time spent on the leaf disk
and proportion of time spent searching during the trial were analysed by two-factor
analysis of variance (ANOVA) where the factors were Trichogramma species (WASP)
and leaf species (LEAF). Data for proportion of time spent searching were arcsin-square
root-transformed before analysis to normalize data. Untransformed means are reported for

96 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

proportion of time spent searching. All analyses were conducted using Sigmastat version
2.0 (Fox et al., 1995).

RESULTS

Walking speed. Walking speed was significantly higher (approximately twice as fast)
on pepper leaves than on tomato leaves for both Trichogramma species (Table 1, Table 2).
There was no significant difference in walking speed between the two Trichogramma
species. There was also no difference in how variation in walking speed in the two wasp
species was affected by the two plant species as indicated by the non-significant
interaction term in the ANOVA.

Table 1
Mean walking speed, residence time and proportion of time spent searching (Mean + SE)
by female 7richogramma wasps on tomato and pepper leaf disks.

Variable T. sibericum T. brassicae

Tomato Pepper Tomato Pepper
Walking speed 0.9+0.1 2.002 je) ts (I 2302
(mm/sec)
Residence time 219 ED 248 + 19 48+8 892 20
(seconds)
Proportion of time 0.68 + 0.05 0.63 + 0.03 0.82 + 0.06 0.73 + 0.05

spent searchin

Table 2
Two-factor analysis of variance of walking speed, residence time and proportion of time
spent searching by female 7richogramma wasps. Factors for analysis are wasp species
(WASP; either 7. sibericum or T. brassicae) and plarit species (PLANT; either tomato or
pepper leaf disks).

Variable Factor F df p
Walking speed WASP 2.1 1 Oils
LEAF 64.5 l <0.001
WASP*LEAF 0.3 l 0.62
Residence time WASP 89.0 1 <0.001
LEAF 4.0 1 0.048
WASP*LEAF 0.1 1 ORs:
Proportion of time WASP 10.1 1 0.002
spent searching LEAF 3.3 1 0.07
WASP*LEAF 0.5 l 0.49

Residence time on leaf disk. Both Trichogramma species and leaf species affected the
residence time of wasps on leaves (Table 1, Table 2). 7. brassicae females spent
significantly less time than 7. sibericum on both tomato and pepper leaf disks. However,
this reflects the manner in which trials were conducted for the two Trichogramma species.
T. sibericum trials of less than 60 s were rejected and this artificially inflated the mean
residence times for this species. Because T. brassicae females dispersed from leaf disks
more readily than 7. sibericum females, shorter trials were retained for analysis. For both
wasp species, LEAF was a significant effect (ie. more time was spent on pepper than on

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 DF

tomato leaf disks). Wasps exited earlier from tomato foliage than from pepper foliage in
trials. Variation in residence time in both wasp species was affected similarly on the two
plant species as indicated by the non-significant interaction term in the ANOVA.

Proportion of time spent searching. Leaf species did not significantly affect the
proportion of time that females spent searching while on leaf disks during trials (Table 1,
Table 2). However, the proportion of time that females spent searching on leaf disks was
significantly higher for 7. brassicae than for T. sibericum. Again, this reflects the
difference in the manner in which trials were conducted for the two species. 7. brassicae
trials often consisted of the wasp walking to the leaf edge and directly off the edge after
release. This sort of trial has a proportion of time spent searching of 1 (or 100% of time
searching), and was much more common for 7. brassicae than for T. sibericum. Again,
the interaction term was non-significant indicating that both wasp species react in a similar
fashion to the two leaf species.

DISCUSSION

There are clear effects of leaf species on the behaviour of both 7. brassicae and T.
sibericum. First, wasps walk at approximately half the speed on tomato leaf disks as on
pepper leaf disks. This difference was consistent in two Trichogramma species of
different ages and physiological conditions. Such a reduction of walking speed on tomato
is similar to what has been reported for other Trichogramma species and 1s probably
caused by the presence of glandular trichomes which impair the walking behaviour of
wasps (Keller 1987; Kauffman & Kennedy 1989; Kashyap et a/. 1991). Second, wasps
leave tomato foliage (with trichomes) sooner than they leave pepper foliage (without
trichomes). This result may indicate a lower preference for tomato foliage than pepper
foliage as a habitat for host search. A higher propensity to disperse from foliage with
higher trichome densities has previously been reported for 7. pretiosum (Kashyap et al.
1991). Because the wasps walked faster and stayed longer on pepper leaf disks, they likely
examined more of the leaf surface available than wasps on tomato leaf disks.

The T. brassicae females used in this study were more behaviourally active and
dispersed from leaf disks much more readily than T. sibericum females. This behavioural
difference could be genetically based, caused by the difference in age between the groups
of wasps (T. brassicae were 3-5 d old; T. sibericum were 1-3 d old), or caused by some
other difference in physiological condition that influences dispersal. Whatever the
explanation, the consistency of walking speed measurements for these two behaviourally-
distinct groups of insects is remarkable. Also, despite differences in residence time
between 7. brassicae and T. sibericum caused by different bioassay methods, we still
detected lower residence times on tomato leaf disks than on pepper for both species.

We have assumed that our measurements of behaviour on a two-dimensional excised
leaf disk are analagous to behaviour on whole plants under field conditions. In addition,
we have assumed that wasps are searching for hosts or evaluating leaf surfaces as host
habitat when walking on leaf surfaces in the laboratory. Our method allowed the detection
of behavioural differences between two plant species that are likely caused by the presence
of trichomes on tomato. Measurement of walking speed in the laboratory has been
recommended to evaluate performance quality for mass-reared Trichogramma wasps
(Cerutti & Bigler 1994), although this measure of quality does not always correlate with
parasitism success (van Hezewik et al. 2000). In light of the results presented here, it
seems critical to measure walking speed on the appropriate plant substrate. Prediction of
performance on tomato crops using walking speed data from pepper leaves would
overestimate searching efficiency of both species of wasps.

Our results show that both 7. brassicae and T. sibericum behave differently on tomato
leaf disks than on pepper leaf disks. If these behavioural differences also occur during

98 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

searching behaviour in commercial greenhouses, the resulting level of biological control
could be lower in tomato greenhouses vs. pepper greenhouses. Wasps might discover
fewer hosts on tomato because lower walking speed and a higher tendency to exit tomato
foliage would reduce the number of hosts encountered per unit time. This hypothesis has
important implications for release rates of Trichogramma in the two crops (ie. higher
release rates may be required in tomato greenhouses). However, our laboratory-derived
results should be validated by conducting releases of Trichogramma into tomato and
pepper crops under operational conditions.

ACKNOWLEDGEMENTS

We thank the British Columbia Greenhouse Research Council for financial assistance.
Many thanks to Bernie Roitberg for the use of his laboratory and video equipment for the
purposes of the study.

REFERENCES

Cerutti, F. & F. Bigler. 1994. Quality assessment of Trichogramma brassicae in the laboratory.
Entomologia Expermentalis et Applicata 75: 19-26.

Coll, M., L.A. Smith & R.L. Ridgway. 1997. Effects of plants on the searching efficiency of a generalist
predator: the importance of the predator-prey spatial association. Entomologia Experimentalis et
Applicata 83: 1-10.

De Clerq, P., J. Mohaghegh & L. Tirry. 2000. Effect of host plant on the functional response of the
predator Podisus nigrispinus (Heteroptera: Pentatomidae). Biological Control 18: 65-70.

Fox, E., Shotton, K. & Ulrich, C. 1995. SigmaStat Statistical Software, Version 2.0. Jandel Corporation,
Chicago, Illinois.

Haren, R.J.F. van, M.M. Steenhaus, M.W. Sabelis & O.M.B. De Ponti. 1987. Tomato stem trichomes and
dispersal success of Phytoseiulus persimilis relative to its prey Tetranychus urticae. Experimental &
Applied Acarology 3: 115-121.

Hezewik, B.H. van, R.S. Bourchier & S.M. Smith. 2000. Searching speed of Trichogramma minutum and
its potential as a measure of parasitoid quality. Biological Control 17: 139-146.

Kashyap, R.K., G.G. Kennedy & R.R. Farrar. 1991. Behavioural response of Trichogramma pretiosum
Riley and Telonomus sphingis (Ashmead) to trichome/methyl ketone mediated resistance to tomato.
Journal of Chemical Ecology 17: 543-556.

Kauffman, W.C. & G.G. Kennedy. 1989. Relationship between trichome density in tomato and parasitism
of Heliothis spp. (Lepidoptera: Noctuidae) eggs by Trichogramma spp. (Hymenoptera:
Trichogrammatidae). Environmental Entomology 18: 698-704.

Keller, M.A. 1987. Influence of leaf surfaces on movements by the hymenopterous parasitoid
Trichogramma exiguum. Entomologia Experimentalis et Applicata 43: 55-59.

Lenteren, J.C. van, L.Z. Hua, J.W. Kamerman & X. Rumei. 1995. The parasite-host relationship between
Encarsia formosa (Hym., Aphelinidae) and Trialeurodes vaporariorum (Hom., Aleyrodidae) XXVI.
Leaf hairs reduce the capacity of Encarsia to control greenhouse whitefly on cucumber. Journal of
Applied Entomology 119: 553-559.

Li, L.Y. 1994. Worldwide use of Trichogramma for biological control on different crops: a survey, pp.
37-54. In E. Wajnberg & S.A. Hassan (eds.), Biological control with egg parasitoids. C.A.B.
International, Oxon, UK.

Obrycki, J.J. 1986. The influence of foliar pubescence on entomophagous insects, pp. 61-83. In: D.
Boethel & R.D. Eikenbary (eds.), Interactions of plant resistance and parasitoids and predators of
insects. Wiley, New York.

Portree, J. 1993. Greenhouse vegetable production guide for commercial producers. Province of British
Columbia, Ministry of Agriculture, Fisheries and Food. Victoria, B.C.

Romeis, J., T.G. Shanower & C.P.W. Zebitz. 1998. Physical and chemical plant characters inhibiting the
searching behaviour of Trichogramma chilonis. Entomologia Experimentalis et Applicata 87: 275-284.

Romeis, J., T.G. Shanower & C.P.W. Zebitz. 1999. Trichogramma egg parasitism of Helicoverpa
armigera on pigeonpea and sorghum in southern India. Entomologia Experimentalis et Applicata 90:
69-81.

Sutterlin, S. & J.C. van Lenteren. 1997. Influence of hairiness of Gerbera jamesonii leaves on the
searching efficiency of the parasitoid Encarsia formosa. Biological Control 9: 157-165.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 9

Stylops shannoni (Stylopidae, Strepsiptera): A New species
for Canada, with comments on Xenos peckii

REX D. KENNER

SPENCER ENTOMOLOGICAL MUSEUM,
UNIVERSITY OF BRITISH COLUMBIA, VANCOUVER, BC

ABSTRACT

The collection of a male Stylops shannoni Pierce and a number of stylopized bees,
Andrena hippotes Robertson, containing both male and female strepsipterans 1s reported.
This appears to be the first host record and the first association of males and females for
this strepsipteran species. This also appears to be the first record for S. shannoni in
Canada. In addition, a specimen of Polistes fuscatus stylopized by Xenos peckii Kirby
was found in the collection of the Spencer Entomological Museum at the University of
British Columbia.

INTRODUCTION

Strepsiptera is an order of peculiar parasitic insects. Taxonomically, strepsipterans have
had an unsettled history, having been included in at least five different orders with a rank
ranging from subfamily to full ordinal status (Bohart 1941). Strepsipterans are now usually
placed in their own order, closely allied with Coleoptera (Kukalova-Peck & Lawrence
1993).

Strepsipterans are not often observed. The females of most species are neotenic and
remain permanently in the host; the adult males, though free-flying, are small and short-
lived. I report the collection of an adult male strepsipteran and a number of stylopized bees.
In addition, I report on a stylopized specimen of paper wasp found in the Spencer
Entomological Museum collection (SEMC) at the University of British Columbia.

MATERIALS AND METHODS

In early April, 2000, during an informal biodiversity survey of my yard in Richmond,
British Columbia, I collected a free-flying male strepsipteran. In the same area was a
flowering bush which was being visited by a number of species of Hymenoptera and
Diptera. Among the Hymenoptera were bees belonging to the families Apidae, Halictidae
and Andrenidae, the last being Andrena sp., some of which were stylopized (Fig. 1). Over
the next three weeks, I monitored the bees visiting the bush. Because the neighborhood is
residential, I was unable to follow the bees leaving the bush to determine the location of
their nests. One bee was found under a clod of earth near the base of the bush but I could
find no evidence of a nest in that area. Three of the female strepsipterans were dissected
from their hosts and fixed in gluteraldehyde for electron microscopic examination. The
puparium of one of the males was opened and the male removed. Several stylopized bees
were coated with gold-palladium and photographed using a scanning electron microscope
at 20 kV.

RESULTS

Between 9-18 April 2000, I collected 88 Andrena sp. in my front yard, of which 24
were parasitized. During the warmest part of the day, 1200—1400 hours, the stylopization
rate of these Andrena sp. was about one in three. This rate dropped to about one in ten for
collections before 10:00. No more free-flying males were observed after the initial

100 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

collection but several bees contained open puparia, indicating males had emerged, and
three males still in their puparia were collected. The latter three specimens were in multiply
stylopized bees, one with a female and the other two each with a second male which had
emerged prior to collection of the bee (Fig. Ic).

Figure 1. Scanning electron micrographs of Andrena (Trachandrena) hippotes Robertson
stylopized by Stylops shannoni Pierce. a and b: female S. shannoni; ¢: male S. shannoni in
its puparium (left) with an open puparium (right) from which a second male had emerged
prior to collection of the bee; d: close up of male puparium.

Using standard keys (Borrer et al. 1989), I was able to confirm that the male
strepsipterans were Stylops sp. as expected from the fact that they were parasitizing
Andrena sp. Two mature males and the fixed female specimens were submitted to J.
Kathirithamby who determined the males to be Stylops shannoni Pierce. W. E. LaBerge
and R. Brooks determined the host bees to be Andrena (Trachandrena) hippotes
Robertson. Voucher specimens of both the strepsipterans and the host bees have been
deposited in the SEMC.

While collating data with respect to paper wasps in the SEMC, a stylopized specimen
with a locality label of Vancouver was found. It carried a female strepsipteran with its head
extruded between tergites three and four.

DISCUSSION

Stylops shannoni was known previously only from a free-flying male collected on
Plummer’s Island, Maryland (Pierce 1918, Bohart 1941). As far as I have been able to

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 101

determine, this is the first time males and females of S. shannoni have been collected
together and this is the first host record for this enigmatic species. Although the host
species for S. shannoni in Maryland was not determined, A. hippotes is found there
(LaBerge 1973). If S. shannoni is generally associated with A. hippotes, it could be found
in much of North America since A. hippotes has a transcontinental range extending from
southern British Columbia to central California on the west coast and Nova Scotia to
Georgia on the east coast (LaBerge 1973).

Stylops erigeniae Pierce is known only from a female also collected on Plummer’s
Island (Pierce 1918). Bohart (1941) suggested that “it is probable that shannoni represents
the male of erigeniae”’. Pierce, who was under the mistaken impression that Sty/ops spp.
are strictly monospecific with respect to host species, erected a new species, S. hippotes
Pierce, for a female stylopizing A. hippotes which was collected in Ohio (Pierce 1909).
Bohart (1941) in his revision of North American Strepsiptera lists S. hippotes as a species
of “uncertain position”. Perhaps S. shannoni is the male of S. hippotes.

Due to the lack of distinctive morphological characters, female Stylops spp. are difficult
to identify (Kathirithamby 1989). Added to that, the poor condition of many of Pierce’s
type specimens (Bohart 1941) will make resolution of these possible synonymies very
difficult. Kathirithamby (personal communication) suggests that DNA analysis will be
required to match males and females.

The stylopized paper wasp found in the SEMC 1s Polistes fuscatus (Fabricius). It is not
P. f. aurifer Saussure, the subspecies which commonly occurs in BC as it has no yellow
spots on tergite two. It is darker overall than typical P. f aurifer even taking into account
the variant formerly called P. f montanus Bequaert (Bequaert 1940). The wasp keys out to
P. f. fuscatus (Bequaert 1940, 1942). Although some records of P. f fuscatus in BC have
been attributed to “assisted transport” (Buckell & Spencer 1950), Leech (1966) reports the
collection near Vernon in 1947 of a stylopized paper wasp which was determined as P. f.
variatus Cresson. Polistes f. variatus has subsequently been synonymized with P. f
fuscatus (Snelling 1974). It seems unlikely that Leech’s record is the result of “assisted
transport”.

The wasp Leech collected had two puparia projecting between tergites five and six, one
laterally and one ventrally. Leech extracted the two male strepsiperans and they were
determined as Xenos peckii Kirby (Leech 1966). One of these specimens was submitted to
the SEMC in 1958 although its current location is not known. The female strepsipteran in
the SEMC P. fuscatus specimen also appears to be X. peckii. The positions of the
strepsipterans in both cases appear to be unusual. According to Salt & Bequaert (1929), in
Polistes sp., male strepsiptera are usually located under tergites 3 or 4 while females are
usually under tergite 5. Further, all examples cited by those authors of extrusion on the
ventral surface of the wasp involve wasps with three or more parasites.

Based on the records presented here, both Stylops shannoni and Xenos peckii should be
added to the Canadian and BC species lists given in Peck (1991).

ACKNOWLEDGEMENTS

I thank J. Kathirithamby for helpful comments and for determining the strepsipterans,
W. E. LaBerge and R. Brooks for determining the host bees, A. Behennah for finding the
stylopized Polistes sp., E. Humphrey for use of supplies and facilities of the BioSciences
Imaging Centre, G. S. Kenner for taking the micrographs in Figure 1.

REFERENCES

Bequaert, J. 1940. An introductory study of Polistes in the United States and Canada with descriptions of
some new North and South American forms (Hymenoptera; Vespidae). Journal of the New York
Entomological Society 48: 1-31.

Bequaert, J. 1942. A new color form of Polistes fuscatus from Canada. The Canadian Entomologist 74:

102 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

159-161.

Bohart, R. M. 1941. Revision of the Strepsiptera with special reference to the species of North America.
University of California Publications in Entomology, 7: 91-159.

Borror, D. J., C. A. Tnplehorn and N. F. Johnson. 1989. An Introduction to the Study of Insects, 6th Ed.
Harcourt Brace College Publishers, New York.

Buckell, E. R. and G. J. Spencer. 1950. The social wasps (Vespidae) of British Columbia. Proceedings of
the Entomological Society of British Columbia 46: 33-40.

Kathirithamby, J., 1989. Review of the order Strepsiptera. Systematic Entomology 14: 41-92.

Kukalova-Peck, , J. and J. F. Lawrence. 1993. Evolution of the hind wing in Coleoptera. The Canadian
Entomologist 125: 181-258.

LaBerge, W. E. 1973. A revision of the bees of the genus Andrena of the western hemisphere. Part VI.
Subgenus Jrachandrena. Transactions of the American Entomological Society 99: 235-371.

Leech, H.B. 1966. A British Columbia record for Xenos peckii Kirby. Journal of the Entomological
Society of British Columbia 63: 40.

Peck, S. B. 1991. Order Strepsiptera, twisted-winged parasites. Jn Y. Bousquet (Ed.), Checklist of Beetles
of Canada and Alaska. Research Branch, Agriculture Canada, Publication 1861/E. pp. 366-367.

Pierce, W. D. 1909. A monographic revision of the twisted winged insects comprising the order
Strepsiptera Kirby. Bulletin of the U. S. National Museum 66.

Pierce, W. D. 1918. The comparative morphology of the Strepsiptera together with records and
descriptions of insects. Proceedings of the U. S. National Museum 54: 391-501.

Salt, G. and J. Bequaert. 1929. Stylopized Vespidae. Psyche 36: 249-282.

Snelling, R. R. 1974. Changes in the status of some North American Polistes (Hymenoptera: Vespidae).
Proceedings of the Entomological Society of Washington 76: 476-479.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 103

Short-range horizontal disruption by verbenone in
attraction of mountain pine beetle (Coleoptera: Scolytidae)
to pheromone-baited funnel traps in stands of lodgepole pine

DANIEL R. MILLER

USDA FOREST SERVICE, SOUTHERN RESEARCH STATION,
320 GREEN STREET, ATHENS GA USA 30602-2044

ABSTRACT

Verbenone interrupted the attraction of mountain pine beetle, Dendroctonus ponderosae
Hopkins, to baited multiple-funnel traps at a distance of <4 m. Catches of beetles in traps
placed >4 m from traps with verbenone were not significantly lower than catches in
control traps. These results are consistent with the short-range phenomenon of
“switching” exhibited by mountain pine beetle in the formation of a spot infestation in
stands of lodgepole pine.

Key words: Dendroctonus ponderosae, Coleoptera, Scolytidae, verbenone,
antiaggregation pheromone, multiple funnel trap

DISCUSSION

Non-destructive semiochemical-based tactics to control populations of the mountain
pine beetle, Dendroctonus ponderosae Hopkins, are appealing where resource management
objectives cannot tolerate removal of infested trees (Borden and Lindgren 1988).
Verbenone is an antiaggregation pheromone used by D. ponderosae, to interrupt attraction
of beetles to trees already colonized by beetles (Lindgren and Borden 1989). However,
operational trials of verbenone to disrupt populations of mountain pine beetles have had
mixed success in stands of lodgepole pine, Pinus contorta var. latifolia Engelmann,
(Amman and Lindgren 1995). Verbenone has been ineffectual in stands of ponderosa pine,
P. ponderosae Laws. (Bentz et al. 1989). Two methods of application have been used in
these verbenone trials: (1) aerially-applied verbenone-impregnated polyethylene beads; or,
the most common method, (2) bubblecaps containing verbenone and attached to trees in a
grid system. The concentration of bubblecap devices deployed in most trials in British
Columbia has been about 100/ha (10 by 10 m spacing regime) (Amman and Lindgren
1995). My objective was to assess the effective range of verbenone, released from
bubblecaps, in interrupting the attraction of D. ponderosae to its aggregation
semiochemicals over a range of 10 m.

On 17 August 1991, seventy 12-unit Lindgren multiple-funnel traps (Phero Tech Inc.,
Delta BC) were deployed in stands of mature lodgepole pine near Penticton, British
Columbia, in ten sets of seven traps per set. The infestation levels of pines by D.
ponderosae in these stands varied from 10-20% of live trees. One trap in each set was
randomly positioned, serving as the first trap for the subsequent array of traps along a
randomly selected compass direction. Five additional traps within each set were positioned
in a linear array, starting 2 m from the first trap, with a space of about 2 m between traps.
The seventh trap in each set was set 10 m from the last trap in the array (20 m from the first
trap) and in the same line, to test if the effect of verbenone was similar across a distance of
10 m. Each trap was hung between trees with twine such that the bottom of each trap was
about 0.5 m above ground. No trap was within 2 m of any tree. Sets of traps were spaced 1

104 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

— 5 km apart. All traps were baited with the primary aggregation semiochemicals for D.
ponderosae (Skillen et al. 1997): (+)-exo-brevicomin flex lure (chemical purity >98%);
polyethylene bubblecap lure containing a 13:87 mixture of cis-and trans-verbenol
[chemical purities 98%, enantiomeric composition 83:17 (—):(+)]; and polyethylene bottle
containing myrcene (chemical purity >95%). In five randomly selected sets of traps
(designated as treated), the first trap in the array was baited additionally with a verbenone
bubblecap (black polyethylene bubblecap containing verbenone [chemical purity >98%,
enantiomeric composition 83:17 (—):(+)]). All lures were supplied by Phero Tech Inc.
(Delta, British Columbia). The remaining five sets of traps were designated as controls.
The verbenols were released at a combined rate of about 1.74 mg/d at 24 °C whereas
verbenone and myrcene were released at about 14 and 281 mg/d at 24-28 °C, respectively
(determined by weight loss) (PheroTech Inc. unpublished data). exo-Brevicomin was
released at about 0.01 mg/d at 24 °C (determined by collection of volatiles) (PheroTech
Inc. unpublished). Catches of insects were collected on 7 September 1991, terminating the
experiment. Voucher specimens were deposited at the Entomology Museum, Pacific
Forestry Centre (Victoria, British Columbia). The data were analyzed with the SYSTAT
statistical package version 9.0 (SPSS 1998). Trap catch data [transformed by In(y + 1)] for
control and treatment were subjected separately to one-way analysis of variance (ANOVA)
using position of trap as the model factor. Fisher’s least significant difference (LSD)
multiple range test was performed when P < 0.05. Paired t-tests were used to compare
mean catches in control and treatment traps for traps >4 m from the first trap.

Table 1.
Total catches of Dendroctonus ponderosae in baited Lindgren multiple-funnel traps from

20 August to 7 September 1991 near Princeton, British Columbia.
Mean (+SE) number of beetles “

Distance of trap Control Verbenone treatment
from 1" trap (m) (n =5) (n = 5)

0 22 oa Dea oa

2 86+ 13.3a 46+15.6 b

4 784193 a 70 + 20.4 be

6 86+17.6a 56 + 10.7 be

8 86+22.4a O72 20S 8c

10 59 16:3)a 81+ 19.0 bc

20 98 2124 70 + 19.7 be

Means within the same column followed by the same letter are not significantly
different at P < 0.05 (LSD test).

The effective range of verbenone in this study seemed to be less than 4 m from the
release point (Table 1). There was a significant difference among mean trap catches of D.
ponderosae in the treated set of funnel traps (F623 = 6.733, P < 0.001), with catches of
beetles significantly lower in traps baited with verbenone than in any of the other traps in
the treated set. There was no significant difference among mean trap catches of D.
ponderosae in the control set of funnel traps (F623 = 0.634, P = 0.702) with a mean (+SE)
catch of 84 + 7 beetles per trap. The mean (+SE) total catch of D. ponderosae in traps 24
m in the control set (403 + 83 beetles/set) was not significantly different from the mean
total catch of beetles in traps >4 m from the verbenone-baited trap in the treated set (374 +
69 beetles per set) ( ¢ test, df = 8, P = 0.795). Results from this study suggest that using the
existing type of bubblecap device to disrupt aggregations of D. ponderosae in stands of
lodgepole pine would require deployment of devices at a much higher density, and

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 105

subsequently would require an excessively high number of bubblecaps (>600-700 devices
per ha at a spacing of 2-4 m). Further tests of verbenone to disrupt attacks by D.
ponderosae should consider devices releasing verbenone at rates much higher than that of
the black verbenone bubblecap used in past trials (about 14 mg/d at 24-28 °C).

ACKNOWLEDGEMENTS

I thank B. Bentz, B.S. Lindgren and K.O. Britton for reviews of the manuscript. J.E.
Brooks and D. Bartens provided field assistance. This study was supported by Forest
Renewal British Columbia (FRBC).

REFERENCES

Amman and Lindgren. 1995. Semiochemicals for management of mountain pine beetle: Status of research
and application. pp. 14-22. In S.M. Salom and K.R. Hobson (eds.). Application of semiochemicals for
management of bark beetle infestations — Proceedings of an informal conference. U. S. Department
of Agriculture Forest Service General Technical Report INT-GTR-318.

Bentz, B., C. K. Lister, J. M. Schmid, S. A. Mata, L. A. Rasmussen and D. Haneman. Does verbenone
reduce mountain pine beetle attacks in susceptible stands of ponderosa pine? U. S. Department of
Agriculture Forest Service Research Note RM-495.

Borden J.H. and B.S. Lindgren. 1988. The role of semiochemicals in IPM of the mountain pine beetle. pp.
247-255. In T.L. Payne and H. Saarenmaa (eds.). Integrated control of scolytid bark beetles. Virginia
Polytechnic Institute and State University, Blacksburg VA.

Lindgren, B. S. and J. H. Borden. 1989. Semiochemicals of the mountain pine beetle (Dendroctonus
ponderosae Hopkins). pp. 83-88. In G.D. Amman (compiler). Proceedings — Symposium on the
management of lodgepole pine to minimize losses to the mountain pine beetle. U.S. Department of
Agriculture Forest Service General Technical Report INT-262.

Skillen, E. L., C. W. Berisford, M. A. Camann and R. C. Reardon. 1997. Semiochemicals of forest and
shade tree insects in North America and management applications. U.S. Department of Agriculture
Forest Service Publication FHTET-96-15.

SPSS Inc. 1998. SYSTAT 9.01 Statistics. Chicago, IL.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 107

Flight Tunnel and Field Evaluations of Sticky Traps for
Monitoring Codling Moth (Lepidoptera: Tortricidae) in Sex
Pheromone-treated Orchards

A. L. KNIGHT, D. LARSON, AND B. CHRISTIANSON

YAKIMA AGRICULTURAL RESEARCH LABORATORY, AGRICULTURAL RESEARCH
SERVICE, USDA 5230 KONNOWAC PASS RD. WAPATO, WA 98951

ABSTRACT

Delta, diamond, and wing style sticky traps baited with codlemone were evaluated in both
flight tunnel and in field trials to determine their performance in capturing male codling
moth Cydia pomonella (L.). Flight tunnel studies found no differences among trap types in
terms of moth orientation behaviors. However, the proportion of moths contacting each trap
type that were caught varied significantly. The 1CP wing trap caught a lower proportion of
moths than the IIB diamond trap due to a significantly lower efficiency in retaining moths
that landed on the trap. The position of a moth’s first contact varied among traps with a
significantly higher proportion landing on the outside of the wing style versus the delta and
diamond traps. A significantly lower proportion of moths first landing on the outside of the
delta trap were caught than for moths landing on the outside of the 1C wing trap. A
significantly lower proportion of moths landing on the front opening of the 1CP wing trap
were captured than for the other traps. No differences were found among trap types for
either the proportion of moths flying into traps or the proportion of these moths captured.
A majority of moths orienting to the diamond and delta traps first landed on the front flap
and walked into the trap. The removal of the front flap from these traps did not affect their
efficiency. However, a significantly greater proportion of moths flew directly into the delta
trap when the flap was removed. Lure position within a delta trap did not affect moth catch,
but it did affect the position of a moth’s first contact with the trap. Lures placed high in the
trap elicited moth landing on the inside surface of the trap’s side or on the outside of the
trap. Moths tended to land on the front flap when lures were placed in the adhesive. The
relative field performance of traps in a sex pheromone-treated apple orchard was consistent
with the flight tunnel studies, however, it was also influenced by moth population density.
The 1CP trap caught significantly fewer moths than the other traps in an orchard with low
codling moth density. The mean cumulative moth catch of each trap type was proportional
to its adhesive-treated surface area within orchards receiving releases of sterile moths.

Key words: codling moth, traps, monitoring, mating disruption, sex pheromones

INTRODUCTION

Traps baited with codlemone, the major sex pheromone component of codling moth, Cydia
pomonella L. (Roelofs et al. 1971), have been used for > 25 yr to monitor populations in tree
fruit orchards (Butt et al. 1974, Maitlen et al. 1976). Cumulative male catches in traps have
been used to establish action thresholds for insecticide usage (Madsen and Vakenti 1972,
Madsen et al. 1974, Riedl and Croft 1974) and as an indicator of phenology (Riedl et al. 1976,
Beers and Brunner 1992). Moth catch has also been used to evaluate the success of mating
disruption in orchards treated with sex pheromone (Vickers and Rothschild 1991).

The efficacy of a variety of trap types has been evaluated for codling moth in field trials
(earlier work summarized in Riedl et al. 1986, Knodel and Agnello 1990, Vincent et a/. 1990,
Kehat et al. 1994). Traps have typically been constructed with inexpensive and disposable

108 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

cardboard or plastic materials and have had a variety of shapes including cylindrical, delta,
diamond and wing-style. Both disposable, sticky and reusable, non-sticky trap designs have
been tested and compared (Knodel and Agnello 1990, Vincent et a/. 1990). A synthesis of this
work led to the suggestion and partial implementation of a standard protocol for monitoring
codling moth with traps and lures (Riedl er a/. 1986). The use of a wing trap with a notched
bottom liner (Pherocon 1CP) baited with a red rubber septum loaded with 1.0 mg codlemone
has been the mostly widely used monitoring system in the western U.S. during the 1980’s and
1990’s (Ried1 et al. 1986, Gut and Brunner 1998).

The adoption of sex pheromone dispensers for mating disruption of codling moth occurred
relatively rapidly during the 1990’s in apple and pear orchards of Washington, California, and
British Columbia, Canada. A prerequisite for the adoption of this new technology was the need
to develop more intensive monitoring programs. Recommendations for monitoring included
the use of a higher density of traps to detect potential problem areas within orchards and
baiting traps with lures containing higher loads of pheromone to minimize the occurrence of
“false negatives” in moth counts (Gut and Brunner 1996). The increased importance of
monitoring in sex pheromone-treated orchards also led to the use of new trap designs including
a larger delta trap and a new diamond-shaped trap. Unfortunately, the variability in the
physical characteristics of these traps has hindered the implementation of a standardized
protocol for monitoring codling moth and has created uncertainty among pest managers
interpreting moth catches (Knight and Christianson 1999). To date, a comparison of these
traps’ performances for codling moth has not been reported.

Optimizing trap design is vital in developing a useful monitoring system. Slight changes
in trap design can modify the pheromone plume structure and strongly affect moth flight and
landing responses to a trap (Lewis and Macaulay 1976). Typically for most pest species, trap
designs have been compared in a non-systematic, ad hoc approach without regard to
understanding the effect of their individual features on moth behavior (Phillips and Wyatt
1992). Conversely, controlled studies of moth behavior in flight tunnels have proven to be
useful in improving trap designs (Foster and Muggleston 1993, Foster et al. 1995). A study
of codling moth’s response to traps under controlled conditions in a flight tunnel has not been
reported. Here we compare codling moth’s behavioral response to four trap designs. In
addition, the field performances of these traps were compared under low and high moth
densities in trials conducted within apple orchards treated with sex pheromone dispensers for
mating disruption.

MATERIALS AND METHODS

Trap types. Studies were conducted with several paper and plastic trap types
manufactured by Trécé Inc. (Salinas, CA) that are commonly used in tree fruits in the western
United States: the delta trap, Pherocon VI; the diamond trap, Pherocon IIB; and the wing style
traps, Pherocon 1C and Pherocon 1CP. The four traps vary in their overall geometries but have
similar exterior dimensions, except for the smaller IIB diamond trap (Table 1). The two wing
traps differ with regard to their bottom piece. The two pieces of the 1C wing trap are separated
by a 5 cm plastic spacer and are the same size. The bottom wing in the 1CP wing trap is
smaller and fits underneath the upper wing. The primary opening of the 1CP wing trap is a
4.0 x 5.6 cm notch cut in the center edge of the bottom piece. The area of the four traps’
interior surfaces coated with adhesive was not related to a trap’s exterior dimensions. The
smaller IIB diamond trap has the largest area coated with adhesive; however, only 50% of this
treated surface is situated on the bottom of the trap. The 1C wing trap has the largest
horizontal adhesive-treated surface area and the 1CP wing trap has the smallest surface area.
Interestingly, the percentage of the horizontal surface that is effectively covered with adhesive

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 109

varied among traps. The bottom surface of the IB diamond trap is the only trap completely
covered with adhesive. The other three traps have 23 - 48% of their inside bottom surface left
untreated (Table 1). The ratio of nonsticky to sticky surfaces varies among traps, primarily due
to the variability in the exterior size of the traps and because all three inside surfaces of the IIB
diamond trap are treated with adhesive. Both the VI delta and the ITB diamond traps have front
flaps (flap height is about 3.0 cm) that are not treated with adhesive. The area and maximum
height of three of the trap’s openings are similar. However, the opening of the 1CP wing trap
is only half as large as the other traps (Table 1).

Table 1.
Physical characteristics of the Pherocon traps (Trécé Inc., Salinas, CA) evaluated in this study
Tra E

Trap characteristics VI delta IIB diamond 1C wing 1CP wing
Exterior dimensions (cm)

length by width 27.0 x 20.0 17.8 x 16.5 26.0 x 22.0 26.0°x 22.0
Area (cm’) of adhesive-covered

bottom inside surface 420.0 248.7 (497.4) * 409.4 227.3
% inside trap bottom

covered with adhesive 87.3 100.0 64.1 S17
Ratio of non-sticky to

sticky trap surfaces 6.9 Le 5.2 8.5
Height (cm) of front flap 3) 3) 1.5 -— 3.0 - -

Area (cm’) of trap opening 42.8 (8.0)° 48.0 (7.0) 42.7 (5.0) 26012 .0)
“Value in parentheses is the area of all interior surfaces covered with adhesive.
> Value in parentheses is the maximum height of the trap’s opening (cm).

Flight Tunnel Studies. The flight tunnel was constructed from 6 mm acrylic sheeting
(1.66 m long, 0.57 m wide and 0.57 m high). A 12-volt DC blower was used to pull air from
the room (maintained at 22-24 °C and 50-60% RH) into a plenum, through a charcoal filter,
and through a series of screens before passing into the tunnel. Air flow through the tunnel was
maintained at 0.25 m/sec. Exhaust was expelled to the outside of the building. Red lights
installed above the tunnel provided enough light (4.3 lux) to make behavioral observations.
Traps were placed on a ring stand 0.31 m above the tunnel floor and 0.20 m from the entrance
of the tunnel. Traps were baited with a halobutyl gray septum loaded with 0.1 mg codlemone.
Lures were pinned to the middle of the trap bottom and above the adhesive in all traps, except
in the study that evaluated the effect of lure position.

Male moths ( < 36 h old) were obtained from the USDA laboratory colony reared on
artificial diet, and conditioned in constant light for 24 - 48 h at 21 °C and 60% RH. Prior to
testing, moths were placed in complete darkness for 30 min then released from a 30 cm high
platform placed near the air outlet end of the tunnel. Individual moths were flown to traps and
moth behavior was recorded for 6 min or until the moth was caught in the trap. New traps were
used after each replicate.

The first study compared moth’s responses to each trap type. Trap order was randomized
on each day. Five moths (18 replicates) were flown consecutively to each trap type. The
occurrence of wing fanning, upwind anemotactic flight, landing on the trap, entering the trap,
and capture were recorded for each moth for the first six replicates. Data were also recorded
for the position of first moth contact with the trap for the last 12 replicates. The location of
first moth contact with the trap was summarized into three categories: landing on the outside
of the trap, landing on the opening of the trap, or flying inside the trap. Moths landing on the
front flaps of the IIB and VI traps were scored as landing on the front opening.

110 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Two additional studies were conducted in the flight tunnel to evaluate specific features of
the trap / lure system. The first test evaluated the response of males to both the IIB and the VI
traps with and without front flaps. Flaps were removed with a razor blade. Forty moths were
flown to each of these four trap types using the same experimental procedure (the order of
traps was randomized each day and five moths were flown consecutively for 6 min to each
trap). Eight replicates were run with each trap. The occurrences of wing fanning, upwind
anemotactic flight, landing on the trap, entering the trap, capture, and the position of first
contact on the trap were recorded for each moth. The second test evaluated the effect of lure
position within the VI delta trap on capture efficiency. Three lure positions within the trap
were compared: pinned to the top center, pinned to the bottom center, and pinned to the
bottom side. The occurrences of wing fanning, upwind anemotactic flight, landing on the trap,
entering the trap, capture, and the position of first contact on the trap were recorded for each
moth. Forty moths were flown to traps with each lure position using the same experimental
procedure described above.

Field trials. Two field tests were conducted to evaluate the performance of the four trap
types in apple orchards treated with sex pheromone dispensers (1,000 Isomate C+ dispensers
per ha, Pacific Biocontrol, Vancouver, WA). Fifteen traps of each of the four trap types were
randomly spaced 20 m apart in an 18 ha ‘Red Delicious’ orchard near Moxee, WA in test 1.

Trap height was standardized at 3.0 m in the canopy (mean (SE) tree height averaged 4.1 (0.1)
m). The test was conducted from 17 April to 4 May 1998. Test 2 was conducted from 20
August to 9 September 1998 in a nearby 14 ha ‘Red Delicious’ orchard. Trap height was
standardized at 3.0 m in the canopy (mean (SE) tree height averaged 4.2 (0.1) m). Ten
replicates of each trap type were randomized within the orchard and spaced 20 m apart. Five
thousand sterile codling moths (50:50 male: female ratio) obtained from the Sterile Insect
Release Program (Osoyoos, British Columbia) were released into this orchard just prior to the
start of the study and again on 27 August and 3 September. Sterilized moths were exposed to
33 krad of gamma radiation and stored at 2 °C for < 48 h prior to release. Moth catch in each
trap was recorded every two days; however moths were not removed from traps during the test.

Data analysis. A multiple comparison test for proportions (Ryan 1960) was used to test
for significant differences (P = 0.05) among trap types in the behavioral response of moths
(orientation to the trap, trap contact, and moth capture) in the flight tunnel tests. Ryan’s test
was also used to test for differences among traps for the proportion of moths first contacting
a given position on the trap (outside, front opening, and inside) and for each position’s capture
efficiency. Fisher’s exact test (2 x 2 contingency table) was used to compare the proportion
of moths captured in tests evaluating the delta and diamond traps with and without front flaps.
Chi-square analysis was used to compare the frequency distribution of moth contact in delta
traps with lures placed at three positions within traps. All moth counts in field trials were
transformed with square root (x + 0.01) and tested with analysis of variance (PROC GLM,
Hintze 1987). Means were separated in significant ANOVA’s with Fisher’s least significance
difference (Hintze 1987).

RESULTS

Flight Tunnel Studies. No difference in the proportion of moths orienting to or touching
the traps was found among traps (Table 2). However, the proportion of moths touching the
1CP wing trap that were caught was significantly lower than for the 1C wing and IIB diamond
traps. The proportion of moths tested that were trapped was significantly lower with the 1CP
wing versus the IIB diamond trap.

The distribution of moth contacts with traps and the proportion of moths captured varied
among traps (Table 3). A significantly higher proportion of moths first contacted the wing

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 11]

traps on the outside of the traps versus the VI delta and IIB diamond traps. The proportion of
moths landing on the outside of the VI delta trap that was eventually caught in the adhesive
was significantly lower than with the 1C wing trap. A significantly lower proportion of moths
contacting the wing traps landed on the opening of the trap versus the proportion landing on
the flaps of the ITB diamond and VI delta traps. Capture efficiency for these moths was
significantly lower for the 1CP than the other traps. No significant difference was found among
traps for the proportion of moths that flew directly into the trap though nearly 3-fold more
moths flew into the VI delta than the wing traps. The capture rate for moths flying into all four
traps was > 72%. Moths entering the IIB diamond trap avoided the two adhesive-covered
upper sides and were never caught on their surfaces.

Table 2.
Flight tunnel response of codling moth males to traps baited with a grey septa loaded with 0.1
mg codlemone, n = 30.

Proportion of moths
Released that Orienting that Contacting trap that Released that

Tra e oriented to tra contacted tra were caught were caught
VI Delta 0.77a 1.00a 0.82ab 0.62ab
IIB Diamond 0.73a 1.00a 0.96a 0.70a
1C Wing 0.70a 0.95a 0.95a 0.63ab
ICP Wing 0.70a 0.90a 0.74b 0.47b

Column proportions are not significantly different if followed by the same letter, P < 0.05;
Ryan’s (1960) multiple comparison test for proportions.

Table 3.
Distribution of male codling moths' first contact with several trap types baited with 0.1 mg

codlemone and the success of moth capture for each trap location in flight tunnel tests (n=6).

Number
of moths Proportion of moths first contacting’:
contacting Outside of trap Front opening of trap Flying inside trap

Trap type trap __ Landing Captured . Landing Captured Landing Captured
Pherocon VI Delta 50 0.10b 0.10b 0.56a 0.83a 0.34a 0.94a
Pherocon IIB Diamond 41 0.10b 0.25ab 0.63a 0.8la 0.27a 0.73a
Pherocon 1C Wing 34 0.56a 0.58a 0.32b 0.82a 0.12a 0.75a
Pherocon 1CP Wing 36 0.56a 0.30ab 0.31b 0.55b 0.14a 0.80a

Column proportions are not significantly different if followed by the same letter, P < 0. 05;
Ryan’s (1960) multiple comparison test for proportions.

' All moths touching each trap type were scored as having landed on one of three areas
(proportions sum to 1.0). The proportion of moths touching each area that were subsequently
captured is summarized in the table under ‘Captured’.

The presence or absence of a front flap in either the VI delta or IIB diamond trap did not
affect moth capture rates (x =0.56.dt — 1. PP 046 X = 044. di = 1. = 051;
respectively). However, the location of moth contact was significantly different in the VI delta
traps with or without flaps (X* = 8.96, df = 2, P < 0.01) but not with the IIB diamond trap (X’
= 2.74, df= 2, P =0.25). Removal of the flap in the VI delta trap increased the proportion of
moths that flew directly into the trap versus landing on the front of the trap and walking in.

Lure position did not affect the efficiency of moth capture in VI delta traps (X* = 2.19, df
= 2, P= 0.24), however it did affect the distribution of moth contact with the trap (X° = 10.04,
df= 2, P<0.01). When the lure was pinned to the interior top of the trap a majority of moths

1B iy J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

flew into the trap and first landed on the inside top surface before falling down onto the
adhesive. In contrast, a majority of moths first landed on the front flap and walked into traps
when the lure was pinned to the center or side of the interior bottom of the trap.

Field trials. Significant differences in moth catch occurred among traps during both field
tests (Table 4). Test 1 was conducted in the spring during the first flight of codling moth and
few moths were caught in traps. Mean moth catch was significantly lower in the 1CP wing trap
versus the other three traps. Test 2 was conducted later in the season and moth catch was > 10-
fold higher in this test than during the spring trial due to the high number of moths released
into the orchard ( > 90% of moths captured were released sterile moths based on the presence
of a red internal dye). The IIB diamond and 1CP wing traps with the smallest adhesive-treated
surfaces caught significantly fewer moths than the larger VI delta and 1C wing traps in this
test. The rate of catch over time leveled off for each trap due to saturation of the adhesive-
treated surfaces (Fig. 1). Cumulative catch in both the IIB diamond and 1CP wing traps
saturated at about 40 moths per trap (Fig. 1). Cumulative moth catch saturated at a higher level
in the VI delta than the 1C wing trap despite having a nearly 30% smaller adhesive-treated
surface area (Fig. 1, Table 1).

Table 4.
Comparison of male codling moth catch in several trap types baited with 10 mg codlemone
red septa within a sex pheromone-treated apple orchard

Mean (SE) moth catch per trap 7

Trap type Test 1° Test 2°
Pherocon VI Delta 6.3 (0.9)a 93.9 (9.5)b
Pherocon IIB Diamond 6.4 (1.7)a 39.6 (2.2)a
Pherocon 1C Wing 6.6 (1.4)a 77.6 (5.7)b
Pherocon 1CP Wing 3.0 (0.7)b 40.8 (4.6)a
Statistical test F353;=5.15 P= 0.05 F 335=24.7 P<0.001

* This test was conducted from 17 April to 4 May 1998.
> This test was conducted from 20 August to 9 September 1998. The orchard was treated with
three releases of 5,000 sterile moths.

DISCUSSION

Codling moth is a direct pest of pome fruit and typically occurs at low densities in
commercial orchards. For example, the action thresholds established for moth catch in sex
pheromone-baited traps are usually < 5 moths per week (summarized in Ried1 ef al. 1986).
Three of the four traps tested in our field study performed similarly in an orchard with a low
to moderate population density of codling moth. At higher moth densities, the area of a trap’s
adhesive-treated surface was an important factor affecting catch. Riedl (1980) found that a
density of > 0.2 moths per cm’ of adhesive-treated surface reduced subsequent codling moth
captures in sticky traps. Data from our study was consistent with this estimate (Fig. 1, Table
1). However, other factors, such as visual cues can play a role in the capture efficiency of a
trap (Foster et al. 1991). Male E. postvittana flying into traps with moths already captured,
landed closer to the sex pheromone lure than in clean traps. The influence of previous moth
captures within a trap on the orientation and landing behavior of codling moth has not been
addressed.

Saturation of sticky traps with moths is a common problem in monitoring tortricid orchard
pests that occur at high densities, such as tortricid leafrollers (Brown 1984, Knight 2001).
However, our data suggest that saturation is not a factor in any of these trap types when the

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 I 3)

100 - '
90

80 —& 1C wing 7 ;
-—® 1CP wing :
70 -& |IB diamond

-e VV! delta

Mean cumulative catch per trap
16)
@

0 2 4 6 8 10 12 14 16 18 20 22
Time Interval (days)

Figure 1. Cumulative catch of codling moth males from 20 August to 9 September 1998 in
four trap types placed in a 14-ha apple orchard. Five thousand sterilized codling moths were
released in the orchard on 20 August on the first day of the test and again on 27 August and
3 September (release dates indicated by vertical arrows).

cumulative moth catch is < 20 moths. Therefore, current recommendations for codling moth
trap maintenance should be adequate, especially in sex pheromone-treated orchards if trap
liners are replaced frequently (Ried et a/. 1986).

Surprisingly, the Pherocon 1CP wing trap performed poorly in both our flight tunnel and
field tests. Previous field trials have reported that the 1CP wing trap was very effective
(Charmillot et al. 1975) and this trap has been widely used to monitor codling moth in the
western United States (Gut and Brunner 1998). However, in our flight tunnel tests the 1CP
wing trap was the least effective among the four traps tested in capturing moths after they
contacted the trap. In particular, a low proportion of moths landing on the front of the trap
were captured. The low efficiency of the 1CP wing trap was apparently due to the absence of
adhesive on either side of the center notch on the bottom liner. Qualitative physical evaluations
of various 1CP wing traps produced by several manufacturers over the last 15 yr suggest that
traps vary tremendously in the deposition of adhesive. Our data suggest that this variability
would have a significant impact on the relative performance of these traps.

The presence of a front flap in a trap has been suggested to serve as an effective barrier
restricting the ability of moths to exit the trap. Riedl (1986) cited unpublished data that the flap
in a diamond-shaped trap increased catch of codling moth. The inclusion of a front barrier in
the IOBC cylinder trap significantly increased catch of codling moth (Charmillot et a/. 1975).
Foster and Muggleston (1993) in a flight tunnel test with Epiphyas postvittana (Walker) found
that the front flap on a delta trap increased the proportion of moths entering the trap that were
caught. Interestingly, they also found that the height of the flap influenced the moth’s landing
position on the adhesive and the catch efficiency of the trap. Higher flaps caused the moths
to land further upwind and farther from the trap’s exit. Flight tunnel studies with Ctenopseustis
obliquana (Walker) showed that removing the front flap from a delta trap increased the
proportion of moths that entered the trap but also increased the proportion of moths that

114 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

escaped (Foster et al. 1995). The flaps in the diamond and delta traps did not play a significant
role in capturing codling moth in our tests. However, we hypothesize that the presence of the
flap in the VI delta trap may be responsible for retaining a higher number of moths compared
with the IC wing trap in our field tests.

Plume structure and species-specific flight behaviors can influence the effectiveness of trap
designs. Clearly, the responses of codling moth we observed to traps placed in clean air ina
flight tunnel may or may not be consistent with its’ responses to traps placed in an orchard
treated with sex pheromone dispensers. Comparative behavioral studies in a flight tunnel of
the leafrollers, Planotortrix octo (Dugdale) and E. postvittana found that the former species
was more sensitive to its pheromone plume structure. When delta traps were placed at
increasing angles to the wind direction moth orientation and capture of only P. octo declined
(Foster et al. 1991). The wide inter-track reversal distances during anemotactic flight of C.
obliquana reduced the effectiveness of delta traps (Foster et al. 1995). A large proportion of
these moths landed on the outside of the trap and lost contact with the plume. Conversely, we
found that only a low proportion of codling moths landed on the outside of the VI delta trap;
however, a significantly lower proportion of these moths were captured compared with the
other trap types. Foster et a/. (1995) improved the delta trap performance for C. obliquana by
increasing the pheromone dose of the lure, which decreased the flight tracking angles. They
also found that by using a rectangular trap moth capture was improved versus the delta trap
with its narrow apex. A rectangular trap design has not been tested for codling moth nor has
the influence of lure dosage on male anemotactic flight been reported.

Lure placement within a trap can be an important factor affecting moth capture. The

efficiency of the delta trap for FE. postvittana was increased when the lure was placed at the
side of the adhesive-treated bottom surface versus the center or higher in the trap (Foster et al.
1991). However, lure placement did not affect the proportion of moths orienting to the trap.
In comparison, lure placement in the VI delta trap in our study with codling moth did not
affect either capture efficiency or moth orientation. Similarly, McNally and Barnes (1980)
reported that there was no difference in the catch of codling moth in a 1C wing trap whether
the lure was placed high or low in the trap.

Sex pheromone-baited traps play a critical role in monitoring codling moth in orchards
treated with sex pheromone for mating disruption. Trap and lure use have been modified since
1990 when the first pheromone dispensers were registered, to reflect the orchard manager’s
need to assess moth population density in disrupted-orchards more than to measure the level
of disruption in the orchard (Gut and Brunner 1996). Traps are positioned within the orchard
and within the canopy to enhance their ability to capture moths, e.g. orchard borders (Knight
and Christianson 1999), upper canopy (Knight 1995, Barrett 1995), and distant from
pheromone dispensers (Knight et al. 1999). Standardization of these factors, as well as trap
and lure type, will likely improve monitoring of codling moth. Our data suggest that the
currently used delta, diamond, and wing style (1C) traps are equally effective in capturing
codling moth at low to moderate moth densities. Proper maintenance of these traps’ adhesive
surfaces is one factor that can be controlled to improve monitoring of codling moth.

ACKNOWLEDGEMENTS

We would like to thank Bill Lingren (Trécé Inc., Salinas, CA) for providing the traps and
lures and Ken Bloem (Sterile Insect Release Program, Osoyoos, British Columbia) for
providing the sterile codling moths. The paper was strengthened by the helpful comments
made by Eugene Miliczky (USDA, ARS, Wapato, WA), Bruce Barrett (Univ. of Missouri,
Columbia, MO), and Mark Brown (USDA, ARS, Kearneysville, WV). This research was
partially funded by the Washington Tree Fruit Research Commission, Wenatchee, WA.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 115

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Phillips, A.D.G. and T. D. Wyatt. 1992. Beyond origami: using behavioural observations as a strategy to
improve trap design. Entomologia Experimentalis et Applicata 62: 67-74.

Riedl, H. 1980. The importance of pheromone trap density and trap maintenance for the development of
standardized monitoring procedures for the codling moth (Lepidoptera: Tortricidae). The Canadian
Entomologist 112: 529-544.

Riedl, H. And B. A. Croft. 1974. A study of pheromone trap catches in relation to codling moth damage. The

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Canadian Entomologist 106: 525-537.

Riedl, H., B. A. Croft, and A. J. Howitt. 1976. Forecasting codling moth phenology based on pheromone trap
catches and physiological time models. The Canadian Entomologist 108: 449-460.

Riedl, H., J. F. Howell, P. S. McNally, and P. H. Westigard. 1986. Codling moth management: use and
standardization of pheromone trapping systems. University of California Division of Agriculture and
Natural Resources. Bulletin 1918, Oakland.

Roelofs, W., A. Comeau, A. Hill, and G. Milicevic. 1971. Sex attractant of the codling moth: characterization
with electroantennogram technique. Science 174: 297-299.

Ryan, T. A. 1960. Significance tests for multiple comparison of proportions, variances, and other statistics.
Psychology Bulletin 57: 318-328.

Vickers, R. A. and G. H. L. Rothschild. 1991. Use of sex pheromones for control of codling moth, pp. 339-354.
In L. P. S. van der Geest and H. H. Evenhuis [eds.], Tortricoid pests. Elsevier, Amsterdam.

Vincent, C., M. Mailloux, E. A. C. Hagley, W. H. Reissig, W. M. Coli, and T. A. Hosmer. 1990. Monitoring
the codling moth (Lepidoptera: Olethreutidae) and the obliquebanded leafroller (Lepidoptera: Tortricidae)
with sticky and nonsticky traps. Journal Economic Entomology 83: 434-440.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 117

a-Pinene and ethanol: Key host volatiles for
Xylotrechus longitarsis (Coleoptera: Cerambycidae)

W.D. MOREWOOD’, K.E. SIMMONDS

CENTRE FOR ENVIRONMENTAL BIOLOGY, DEPARTMENT OF BIOLOGICAL
SCIENCES, SIMON FRASER UNIVERSITY, 8888 UNIVERSITY DRIVE,
BURNABY, BC V5A 1S6

I.M. WILSON?
PHERO TECH INC., 7572 PROGRESS WAY, DELTA, BC V4G 1E9

J.H. BORDEN, R.L. MCINTOSH°

CENTRE FOR ENVIRONMENTAL BIOLOGY, DEPARTMENT OF BIOLOGICAL
SCIENCES, SIMON FRASER UNIVERSITY, 8888 UNIVERSITY DRIVE,
BURNABY, BC V5A 1S6

ABSTRACT

Xylotrechus longitarsis Casey is a common wood-boring beetle that causes
considerable losses to the softwood lumber industry in British Columbia. Trapping
experiments were conducted in the southern interior of British Columbia to determine
whether the generic bait of apinene plus ethanol is an optimal combination for
attraction of X. Jongitarsis. o-Pinene alone attracted substantial numbers of X.
longitarsis and trap catches were increased significantly with the addition of ethanol
lures but different release rates of ethanol had no significant effect. The blend of pure
a-pinene and ethanol was significantly more attractive than more complete blends of
conifer monoterpenes and ethanol, whether or not a-pinene was a major component of
the blend. Strong attraction to a-pinene reflects the preference of X. longitarsis for
conifers, and particularly Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco, the bark
of which is relatively rich in a-pinene. Increased attraction with the addition of ethanol
lures, irrespective of release rates, to traps baited with a-pinene suggests that X.
longitarsis prefers severely stressed hosts or deteriorating host material but may also
utilize freshly cut or broken material.

Key words: Xylotrechus longitarsis, Cerambycidae, chemical ecology, host volatiles,
woodborers, pest management, trapping

INTRODUCTION

Xylotrechus longitarsis Casey is a coniferophagous wood-boring beetle that is common
and widespread in British Columbia (BC), ranging east into Alberta and south to Colorado
and northern California (Linsley 1964). The flight period is reported as May to August

' Current Address: Department of Entomology, 501 ASI Building, Pennsylvania State
University, University Park, PA 16802 Email: wdmorewood@alumni.uvic.ca

; Current Address: Parks Division, City of Kelowna, 1359 K.L.O. Road, Kelowna, BC
V1W 3N8

: Current Address: Saskatchewan Environment and Resource Management, PO Box 3003,
Prince Albert, SK S6V 6G1

118 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

(Linsley 1964) but adults may be abundant well into September in the southern interior of
BC, where this species is usually the most numerous of the large wood-boring insects
(Cerambycidae, Buprestidae, Siricidae) captured around log storage areas (McIntosh et al.
2001; Morewood ef al. 2002). The relative abundance of X. /ongitarsis suggests that it is a
major contributor to the estimated $43.6 million (US) in annual degrade losses to the
softwood lumber industry caused by large woodborers in the interior of BC (Phero Tech
Inc. 1997).

Wood-boring insects are pests of timber destined for lumber production, attacking and
degrading dead or dying trees and logs after harvest but before processing. Management of
large woodborers has been limited primarily to a strategy of preventing attack through
rapid utilization, peeling, water sprinkling, or storage of logs in water or in compact decks
with maximum shading (Safranyik and Moeck 1995), with operational trapping programs
currently under development. In contrast, prevention of attack by ambrosia beetles
(Coleoptera: Scolytidae) is accomplished in part through well-established trapping
programs using semiochemical-baited traps (Borden 1995). Various combinations of host
monoterpenes and ethanol are attractive to coniferophagous cerambycids (Ikeda et al.
1980; Fatzinger et al. 1987; Phillips et al. 1988), with some evidence that a-pinene is the
most attractive of individual monoterpenes tested (Ikeda et al. 1986; Chénier and
Philogene 1989). Currently, the simple combination of a-pinene and ethanol is used
provisionally as a generic bait for trapping large woodborers in BC. However, this blend
does not replicate the complex mixture of host-associated chemical cues that could be used
by these insects during host selection. Our objective was to determine whether more
complete blends of conifer volatiles, primarily monoterpenes and ethanol, would be more
attractive to X. Jongitarsis than pure a-pinene and ethanol.

MATERIALS AND METHODS

Different combinations of host volatiles were tested as baits in 12-unit multiple funnel
traps (Lindgren 1983) suspended from metal poles with the top funnel about 1.5 m above
ground. Captured insects were frozen until they could be sorted and counted.

To test whether ethanol is important in attracting XY. longitarsis and whether such
attraction is affected by release rate, an experiment was superimposed on operational
trapping programs around the log storage yard at Gorman Bros. Ltd. in Westbank and the
Riverside Forest Products Ltd. 4-Mile dryland log sort and Okanagan Lake log dump, both
on the Bear Forest Service Road west of Kelowna. Traps were baited with a-pinene [96%
(—) enantiomer, > 99% chemical purity, released at 2.2 g/d; Phero Tech Inc., Delta, BC]
alone or combined with either low-release (30-50 mg/d) or high-release (1.8 g/d) ethanol
lures in randomized complete blocks on 2 September 1999. Insects were collected from
nine blocks of traps on 10 September 1999 and 12 blocks of traps on 17 September 1999,
for a total of 21 replicates.

To test whether apinene is an adequate host-recognition cue for X. Jongitarsis,
compared to more complete blends of conifer monoterpenes, two experiments were
conducted in a small clearcut 8.3 km up the Laluwissen Forest Service Road north of
Lytton. The surrounding forest was predominantly Douglas-fir, Pseudotsuga menziesii
(Mirbel) Franco, with some lodgepole pine, Pinus contorta Douglas ex Loudon, and
ponderosa pine, Pinus ponderosa P. & C. Lawson. In each experiment, catches in traps
baited with a blend of host volatiles distilled from lodgepole pine turpentine (H.D. Pierce
Jr., Department of Biological Sciences, Simon Fraser University, unpublished data) were
compared to catches in traps baited with pure a-pinene (as above, release rate 150 mg/d)
and to catches in unbaited traps, deployed in 10 randomized complete blocks. Both host
blends and a-pinene were combined with low-release ethanol lures. In the first experiment,

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 E19

traps were baited with a modified host blend (released at 750 mg/d), with B-phellandrene
enriched and a-pinene nearly eliminated (Table 1), on 1 August 2001 and insects were
collected on 14 August 2001. In the second experiment, traps were baited with a natural
host blend (released at 700 mg/d), distilled without greatly altering the relative proportions
of major components (Table 1), on 28 August 2001. Insects were collected on 11
September 2001 and the treatments were re-randomized to minimize position effects.
Insects were collected again on 23 September 2001 to compensate for the declining
catches later in the season, and the two collections were combined for a total of 10

replicates.

Table 1
Composition of “natural” and “modified” host blends distilled from lodgepole pine
turpentine and used in trapping experiments for Xylotrechus longitarsis.

Compound Enantiomeric composition (%) Percent of host blend
(+) (-) Natural Modified
a-pinene Ad 56 18.4 0.9
B-pinene 0 100 16.6 4.4
3-carene 100 0 23.6 24.8
limonene 42 58 3.8 7.8
8-phellandrene 0 100 28.0 51.0
unidentified minor components __n.a. n.a. 9.6 11.1

Data were transformed by log(x + 1) to correct for non-normality and
heteroscedasticity (Théni 1967; Zar 1999) and then subjected to analysis of variance
(PROC ANOVA) for randomized complete blocks (Kvanli 1988; SAS Institute Inc. 1999).
Means were compared using the Ryan-Einot-Gabriel-Welsch (REGWQ) multiple range
test (Day and Quinn 1989; SAS Institute Inc. 1999).

RESULTS

Traps baited with o-pinene plus ethanol captured significantly greater numbers of X.
longitarsis than traps baited with a-pinene alone (F> 49 = 5.89, P = 0.0057), but trap catches
did not differ significantly between traps with ethanol released at high or low rates (Fig. 1).

Traps baited with a-pinene plus ethanol captured significantly greater numbers of both
sexes of X. Jongitarsis than traps baited with ethanol plus either the modified host blend
(F213 = 16.06, P < 0.0001 for males; F213 = 38.06, P < 0.0001 for females) or the natural
host blend (F213 = 19.82, P < 0.0001 for males; F, 3 = 11.54, P = 0.0006 for females).
Traps baited with either host blend captured significantly greater numbers of males, but
not females, than unbaited traps (Fig. 2).

DISCUSSION

The simple combination of a-pinene and ethanol appears to be particularly well-suited
for attracting X. /ongitarsis, reflecting both host preferences and considerable flexibility
with respect to suitable host material. The only host association published for X.
longitarsis is Douglas-fir (Linsley 1964) and most of the reared specimens in the insect
collection at the Pacific Forestry Centre, Victoria, BC, are from Douglas-fir. a-Pinene is
the dominant monoterpene in Douglas-fir bark (D.S. Pureswaran, Department of
Biological Sciences, Simon Fraser University, unpublished data) and is thought to be the
primary attractant for the Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins
(Coleoptera: Scolytidae) (Heikkenen and Hrutfiord 1965).

120 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

a-pinene
a-pinene + low-release ethanol

a-pinene + high-release ethanol}:

0 10 20 30
NUMBER OF BEETLES CAPTURED (MEAN + SE)

Figure 1. Catches of Xylotrechus longitarsis at three locations in the central Okanagan
Valley, BC, 2-17 September 1999 in traps baited with a-pinene (released at 2.2 g/d) alone
or combined with either low-release (30-50 mg/d) or high-release (1.8 g/d) ethanol lures.
The difference between the bars with the same letter is not statistically significant
(REGWQ, P > 0.05).

1-14 August 2001

unbaited control]

modified host blend + ethanol

a-pinene + ethanol

unbaited control
natural host blend + ethanol
a-pinene + ethanol

9 6 3 0 3 6 9
NUMBER OF BEETLES CAPTURED (MEAN + SE)

Figure 2. Catches of Xylotrechus longitarsis near Lytton, BC, in August and September
2001 in traps baited with a modified (above) or natural (below) host blend (released at 750
mg/d or 700 mg/d, respectively) or a-pinene (released at 150 mg/d), each combined with
ethanol (released at 30-50 mg/d), or unbaited. For a given time period and sex, differences
between bars with the same letter are not statistically significant (REGWQ, P > 0.05).

On the other hand, the geographic range of X. Jongitarsis extends beyond that of
Douglas-fir and other conifer species can serve as hosts. We have reared _X. /ongitarsis in
ponderosa pine bolts and there are specimens in the insect collection at the Pacific Forestry
Centre reared from logs of lodgepole pine, western hemlock, Tsuga heterophylla
(Rafinesque-Schmaltz) Sargent, western larch, Larix occidentalis Nuttall, and white
spruce, Picea glauca (Moench) Voss. Despite this apparent broad acceptability of different
conifer hosts, including lodgepole pine, monoterpene blends derived from lodgepole pine
were significantly less attractive than pure a-pinene (Fig. 2). Furthermore, this lack of
attraction was not simply due to the lack of a-pinene in the modified host blend because
the natural host blend, which contained a substantial amount of a-pinene (Table 1) and
released it at a rate of 129 mg/d (compared to 150 mg/d for the pure compound), was no
more attractive relative to pure a-pinene than the modified host blend (Fig. 2). Coupled gas
chromatographic — electroantennographic detection analyses indicate that the antennae of
X. longitarsis can detect each of the five prominent monoterpenes listed in Table 1 (R.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 121

Gries, Department of Biological Sciences, Simon Fraser University, unpublished data).
These considerations suggest that some component of these natural host blends is either
repellent to X. /ongitarsis or interferes with the response to a-pinene.

Like many cerambycid species that breed in severely stressed hosts (Hanks 1999),
adult X. /ongitarsis are attracted to recently felled trees and cut logs. Ethanol is a primary
attractant for other species that breed in recently felled trees and cut logs, such as the
ambrosia beetles Guathotrichus sulcatus LeConte and Trypodendron lineatum (Olivier)
(Cade et al. 1970; Moeck 1970), and might be considered a general indicator of stress in
trees (Kimmerer and Kozlowski 1982; Kelsey and Joseph 1998). Cut or broken conifers
initially release large amounts of monoterpenes, with gradually increasing amounts of
ethanol as the material deteriorates, and many bark and ambrosia beetles and their
associates are attracted to ratios of a-pinene and ethanol that reflect the condition of host
material to which each species is adapted (Schroeder and Lindeléw 1989). In contrast, X.
longitarsis would appear to accept host material in a broad range of conditions,
considering the similarity of catches in traps with very different release rates of ethanol
and the substantial catches even in traps baited with a-pinene alone (Fig. 1).

For operational trapping, the simple combination of a-pinene and ethanol appears to be
optimal for attracting X. Jongitarsis and also has economic and practical advantages over
more complete monoterpene blends. A lure containing a single compound is likely to be
less expensive than one with a more complex blend and a-pinene is more easily purified
than other conifer monoterpenes (H.D. Pierce Jr., personal communication). In addition,
blends containing large amounts of B-phellandrene are unpleasant to work with because B-
phellandrene is unstable and tends to polymerize on the outside of the lures as it is
released, creating a sticky mess.

ACKNOWLEDGEMENTS

We thank Hugh Norman (BC Chemicals, Prince George, BC) for the generous gift of
lodgepole pine turpentine, Nick Arkle (Gorman Bros. Ltd.), Gary Greyell (Riverside
Forest Products Ltd.), and Ed Senger (BC Ministry of Forests) for access to field sites, and
Bob Duncan for access to the insect collection at the Pacific Forestry Centre. Financial
support was provided by Forest Renewal British Columbia, the Science Council of British
Columbia, the Natural Sciences and Engineering Research Council of Canada, Abitibi
Consolidated Forest Products Inc., Ainsworth Lumber Co. Ltd., British Columbia Hydro
and Power Authority, Bugbusters Pest Management Inc., Canadian Forest Products Ltd.,
Gorman Bros. Ltd., International Forest Products Ltd., Lignum Ltd., Manning Diversified
Forest Products Ltd., Millar Western Forest Products Ltd., Phero Tech Inc., Riverside
Forest Products Ltd., Slocan Forest Products Ltd., Tembec Forest Industries Ltd.,
TimberWest Forest Corp., Tolko Industries Ltd., Weldwood of Canada Ltd., West Fraser
Mills Ltd., Western Forest Products Ltd., and Weyerhaeuser Canada Ltd.

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In: J.A. Armstrong and W.G.H. Ives (Eds.). Forest insect pests in Canada. Natural Resources Canada,
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Cade, S.C., B.F. Hrutfiord and R.I. Gara. 1970. Identification of a primary attractant for Gnathotrichus
sulcatus isolated from western hemlock logs. Journal of Economic Entomology 63: 1014-1015.

Chénier, J.V.R. and B.J.R. Philogéne. 1989. Field responses of certain forest Coleoptera to conifer
monoterpenes and ethanol. Journal of Chemical Ecology 15: 1729-1745.

Day, R.W. and G.P. Quinn. 1989. Comparisons of treatments after an analysis of variance in ecology.
Ecological Monographs 59: 433-463.

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Fatzinger, C.W., B.D. Siegfried, R.C. Wilkinson and J.L. Nation. 1987. Trans-verbenol, turpentine, and
ethanol as trap baits for the black turpentine beetle, Dendroctonus terebrans, and other forest
Coleoptera in north Florida. Journal of Entomological Science 22: 201-209.

Hanks, L.M. 1999. Influence of the larval host plant on reproductive strategies of cerambycid beetles.
Annual Review of Entomology 44: 483-505.

Heikkenen, H.J. and B.F. Hrutfiord. 1965. Dendroctonus pseudotsugae: A hypothesis regarding its primary
attractant. Science 150: 1457-1459.

Ikeda, T., N. Enda, A. Yamane, K. Oda and T. Toyoda. 1980. Attractants for the Japanese pine sawyer,
Monochamus alternatus Hope (Coleoptera: Cerambycidae). Applied Entomology and Zoology 15:
358-361.

Ikeda, T., A. Yamane, N. Enda, K. Oda, H. Makihara, K. Ito and I. Okochi. 1986. Attractiveness of volatile
components of felled pine trees for Monochamus alternatus (Coleoptera: Cerambycidae). Journal of
the Japanese Forestry Society 68: 15-19.

Kelsey, R.G. and G. Joseph. 1998. Ethanol in Douglas-fir with black-stain root disease (Leptographium
wageneri). Canadian Journal of Forest Research 28: 2107-1212.

Kimmerer, T.W. and T.T. Kozlowski. 1982. Ethylene, ethane, acetaldehyde, and ethanol production by
plants under stress. Plant Physiology 69: 840-847.

Kvanli, A.H. 1988. Statistics: A computer integrated approach. West Publishing, St. Paul, Minnesota.

Lindgren, B.S. 1983. A multiple funnel trap for scolytid beetles (Coleoptera). The Canadian Entomologist
115: 299-302.

Linsley, E.G. 1964. The Cerambycidae of North America. Part V. Taxonomy and classification of the
subfamily Cerambycinae, tribes Callichromini through Ancylocerini. University of California
Publications in Entomology, Volume 22. University of California, Berkeley.

McIntosh, R.L., P.J. Katinic, J.D. Allison, J.H. Borden and D.L. Downey. 2001. Comparative efficacy of
five types of trap for woodborers in the Cerambycidae, Buprestidae and Siricidae. Agricultural and
Forest Entomology 3: 113-120.

Moeck, H.A. 1970. Ethanol as the primary attractant for the ambrosia beetle 7rypodendron lineatum
(Coleoptera: Scolytidae). The Canadian Entomologist 102: 985-995.

Morewood, W.D., K.E. Hein, P.J. Katinic and J.H. Borden. 2002. An improved trap for large wood-boring
insects, with special reference to Monochamus scutellatus (Coleoptera: Cerambycidae). Canadian
Journal of Forest Research 32: 519-525.

Phero Tech Inc. 1997. Damage assessment of wood borers in the interior of B.C. Final report, May 1997.
Phero Tech Inc., 7572 Progress Way, Delta, B.C. V4G 1E9. (unpublished)

Phillips, T.W., A.J. Wilkening, T.H. Atkinson, J.L. Nation, R.C. Wilkinson and J.L. Foltz. 1988.
Synergism of turpentine and ethanol as attractants for certain pine-infesting beetles (Coleoptera).
Environmental Entomology 17: 456-462.

Safranyik, L. and H.A. Moeck. 1995. Wood borers. pp. 171-177 Jn: J.A. Armstrong and W.G.H. Ives
(Eds.). Forest insect pests in Canada. Natural Resources Canada, Canadian Forest Service, Ottawa.

SAS Institute Inc. 1999. The SAS system, version eight. SAS Institute Inc., Cary, North Carolina.

Schroeder, L.M. and A. Lindelow. 1989. Attraction of scolytids and associated beetles by different
absolute amounts and proportions of a-pinene and ethanol. Journal of Chemical Ecology 15: 807-817.

Thoéni, H. 1967. Transformations of variables used in the analysis of experimental and observational data.
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J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 123

A Comparison of Gray Halo-butyl Elastomer and Red
Rubber Septa to Monitor Codling Moth (Lepidoptera:
Tortricidae) in Sex Pheromone-Treated Orchards

ALAN L. KNIGHT

YAKIMA AGRICULTURAL RESEARCH LABORATORY, AGRICULTURAL RESEARCH
SERVICE, USDA 5230 KONNOWAC PASS RD. WAPATO, WA 98951

ABSTRACT

The emission rate, isomeric stability, and relative attractiveness of field-aged gray halo-butyl
elastomer and red rubber septa loaded with 4.0 and 10.0 mg of (£,£)-8,10-dodecadien-1-ol
(E8,E10-12:0H, codlemone), the major sex pheromone component for codling moth, Cydia
pomonella L., were evaluated. Initially, field-aged red septa loaded with 10.0 mg had
significantly higher emission rates than gray septa loaded with 4.0 mg codlemone. Emission
rates of codlemone decayed over time from both lures and were similar for lures aged 28-42
d in the field. Isomerization of E8,E10-12:OH occurred rapidly in red septa but did not
occur in the gray septa. Moth capture in traps baited with either lure type aged in the field
did not differ initially (0 and 7 d), but were significantly lower in traps baited with 14, 28,
and 42 d-old red versus gray septa. Significant differences observed in the attractiveness of
these two types of field-aged lures were primarily due to changes in their isomeric purity and
not to differences in their emission rate. Increasing the codlemone load of gray septa up to
20.0 mg did not improve the performance of lures 1n sex pheromone-treated orchards. All
lures were effective for 10 wk. Loading gray septa with 50.0 mg codlemone increased lure
attractiveness and extended its longevity to 16 wk. Proprietary gray septa loaded with a high
rate of pheromone and replaced once per season were more attractive than the standard 10.0
mg loaded red septa replaced three times in a sex pheromone-treated orchard.

Key words: Cydia pomonella, codling moth, sex pheromone, monitoring, lures

INTRODUCTION

Codling moth, Cydia pomonella L., is the major direct pest of pome fruits and walnut
throughout the world (Shel’deshova 1967). Following the identification of the sex pheromone
of codling moth, (£, £)-8,10-dodecadien-1-ol (E8, E10-12:OH, codlemone) (Roelofs ef al.
1971), moth catches in pheromone baited traps have been widely used to establish action
thresholds (Riedl and Croft 1974) and to track the seasonal timing of key population events
(Riedl et al. 1976). Traps baited with red rubber septa impregnated with codlemone have been
the most widely used dispenser system (Riedl et al. 1986). Lure loadings between 9.1 and 1.0
mg codlemone have provided optimal attraction (Maitlen et al. 1976, Culver and Barnes 1977,
McNally and Barnes 1980, Kehat et a/. 1994).

Maitlen et al. (1976) characterized the first-order release characteristics of codlemone from
red rubber septa under controlled laboratory conditions (23 °C). Based on both analytical and
biological studies they determined that a 1.0 mg lure should maintain a maximum level of
attraction for up to four wk. Their calculations suggested that a 5.0 mg loading would remain
effective for >4 months. Ried et a/. (1986) used the half-life calculations of Maitlen et ai.
(1976) to predict that red rubber septa loaded initially with 1.0 mg codlemone should be
effective for up to 11 wk (based on maintaining a residual pheromone content >0.1 mg). This
predicted long field-life of the red rubber septa was demonstrated by McNally and Barnes
(1980). Field-aged lures loaded with 1.0 mg codlemone were equally effective as new lures

124 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

for up to 16 wk. However, Culver and Barnes (1977) found that the attractiveness of 1.0 mg
lures dropped significantly after 2-3 wk in the field. Ried] et a/. (1986) hypothesized that this
large difference observed in the longevity of field-aged red rubber septa might be due to
changes in the manufacturer’s formulation. Subsequent field studies found that both the
emission rate and the attractiveness of red rubber septa declined rapidly after two wk (Kehat
et al. 1994).

Concurrent with the rapid adoption of sex pheromone dispensers for mating disruption of
codling moth during the 1990’s was the use of monitoring lures loaded with higher rates of
codlemone (Charmillot 1990). Red rubber septa loaded with 10.0 mg codlemone have been
the most common lure used in sex pheromone-disrupted orchards in western North America
(Knight 1995a, Judd et al. 1996, Gut and Brunner 1998). Lures with high loads of codlemone
have allowed pest managers to track the population density and phenology of codling moth
within sex pheromone-treated orchards. Current monitoring recommendations of sex
pheromone-treated orchards with 10.0 mg red septa suggest changing lures every 2-3 wk (Gut
and Brunner 1996). The short life of these high load lures was hypothesized to be due to the
depletion of codlemone under high summer temperatures (Gut and Brunner 1995), similar to
the results reported from Israel with standard lures (Kehat et al. 1994). This frequent
replacement schedule for lures combined with a higher density of traps has increased the cost
of monitoring sex pheromone-treated orchards relative to conventional orchards (Knight
1995a).

The chemical instability of many sex pheromones can severely limit the longevity of lures
(House et al. 1998). The chemical sensitivity of codlemone, a conjugated diene alcohol, to
heat, light, and air is particularly acute (Brown and McDonough 1986, Ideses and Shani 1988).

Several biochemical pathways can degrade codlemone, including isomerization and oxidation
to peroxides and furans (Millar 1995). Rapid isomerization of codlemone within rubber septa
is catalyzed by the presence of sulphur and sunlight (Brown and McDonough 1986).
Isomerization of codlemone within sulphur-cured rubber septa begins immediately (Vrko€ et
al. 1988), and the three isomers may account for 20% of the content within seven days in field-
aged lures placed within traps (Brown and McDonough 1986). Chemical protection of
codlemone can be accomplished with the addition of UV stabilizers and antioxidants (Ideses
and Shani 1988, Millar 1995) or by the use of other lure substrates. For example, the
isomerization of codlemone was 4.7 fold slower within phenolic resin-cured, gray halo-butyl
elastomer lures versus the red rubber septa (Brown and McDonough 1986).

The isomeric purity of codlemone is an important factor affecting a lure’s performance.
Various crude mixtures of the three geometrical isomers of codlemone decreased the
attractiveness of codlemone when incorporated together within red rubber septa-baited traps
(Roelofs et al. 1972). A chemical equilibrium blend of the four isomers (61% EE, 5% ZZ, 14%
ZE, and 20% EZ) significantly reduced the flight response of males in flight tunnel tests versus
pure codlemone (McDonough et al. 1993). El-Sayd et al. (1998) found that the single addition
of the Z,E isomer to codlemone synergized the response of codling moth in a flight tunnel, but
moth capture in traps was not affected in a subsequent field study. In contrast, the addition of
20% E,Z isomer strongly depressed male responses both in a flight tunnel and to traps placed
in the field.

The objective of this study was to evaluate the effects of emission rate and isomeric purity
of codlemone on the attractiveness of lures for codling moth. Herein are reported the emission
rate and isomeric purity of field-aged red rubber and gray halo-butyl elastomer septa. The
relative attractiveness of lures was evaluated in a series of field trials within sex pheromone-
treated apple orchards. Subsequent field tests evaluated the optimal loading rate of gray septa
for season-long monitoring of codling moth in sex pheromone-treated orchards.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 Ws)

MATERIALS AND METHODS

Preparation of lures. Gray halo-butyl elastomer septa (No. 1888, size No. 1) and red,
natural rubber septa (No. 1171, size 1F) were used in all tests (West Co., Phoenixville. PA).
Septa were extracted in hexane for 24 h, dichloromethane for another 24 h, and then air-dried
for 48 h prior to loading. E8,E10-12:0H (97% purity, Aldrich Chemical, Minneapolis, MN)
was added to the cup portion of the septa in a 200-1 aliquot of dichloromethane, followed by
another 200 pl of dichloromethane to ensure penetration of the material. Initial studies found
that a maximum of 4.0 mg codlemone could be loaded in gray septa using this technique.
Codlemone crystallized on the surface of these lures at higher loadings. Thus, red and gray
septa were loaded with 10.0 and 4.0 mg E8,E10-12:OH, respectively. Septa were kept frozen
at —15 °C prior to use.

Evaluation of field-aged septa. Prepared septa were field-aged by pinning lures inside
of non-sticky Pherocon IC wing traps (Trécé Inc., Salinas, CA) hung at 2.0 m height in the
canopy of an unsprayed apple orchard (Yakima, WA). Eight septa of one type were placed 4.0
cm apart inside of traps, and pinned 1.5 cm above the traps’ bottom surfaces on 8 July 1995.
Nine lures of each type were randomly collected at weekly intervals (0, 7, 14, 21, 28, 35, 42
and 49 d field exposure) for subsequent field testing and chemical analyses.

The average daily emission rate of lures during each time interval was estimated by
differential residual analysis (difference between the mean sex pheromone content at the
beginning of the time period and the residual values at the end of the time period). Four septa
of each type from each date were extracted with 50 ml dichloromethane by shaking lures in
a flask for 1 h. The extract was further diluted in dichloromethane and heptane and analyzed
with a Hewlett-Packard Model 5880 GC with flame ionization detection (Hewlett Packard,
Mountain View, CA) and equipped with a 60 m by 0.32 mm1.d. Supelcowax capillary column
(Supelco Inc., Bellefonte, PA) with splitless injection. The oven program used for analysis
was: 60 °C for 2 min; ramping at 20 °C/min to 154 °C for 17 min; and a 5-min purge at 210
°C. Recovery rates for E8,E10-12:0H averaged 97 — 102%. The isomeric purity of dispensers
was confirmed with a GC- mass spectrometer (Hewlett-Packard model 5970 GC coupled with
a HP 5970 mass detector).

The attractiveness of the field-aged septa (0, 7, 14, 21, 28, 35 and 42 d) was evaluated in
a commercial apple orchard in July 1996 treated with 1,000 polyethylene dispensers (Isomate-
C+, Pacific Biocontrol, Vancouver, WA) per ha. Polyethylene dispensers were loaded with
182.0 mg of a three-component sex pheromone for codling moth (60:34:6 blend of E8, E10-
12:OH: dodecanol: tetradecanol). Treatments were arranged in a complete randomized block
design. On each date (17, 24 and 31 July and 7 and 14 August) one replicate of each lure type
and age was tested. Traps were hung on trees at a height of 3.0 m and spaced 30 m apart.

Unsexed sterile moths were obtained from the codling moth mass-rearing SIR facility in
Osoyoos, British Columbia. Moths were sterilized with gamma radiation (33 krad) from a
Cobalt” source (dose rate of 1,150-1,320 rad/min) and held at 0 to 2 °C before field release.
The sterilized moths had been marked with a red internal dye during larval mass-rearing. Three
hundred sterilized codling moths were released around each trap at the beginning of each test
by tapping chilled moths out of petri plates onto both the foliage of trees and on the ground
within 10 m of each trap. Traps were checked after six nights and only counts of sterilized
moths were recorded. Treatments were re-randomized prior to the start of the next replicate.

Optimizing the attractiveness of gray septa. Studies were conducted to evaluate the
optimal loading rate of gray halo-butyl septa for use in sex pheromone-treated orchards. Gray
septa were loaded with varying rates of E8, E10-12:OH (4.0, 10.0, 20.0, and 50.0 mg) by
Trécé Inc. (Salinas, CA) personnel using a proprietary technique. Lures were placed in
specialized wire hangers and field-aged within an apple orchard situated near Fresno, CA on

126 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

12 April 1998. Five lures of each loading rate were collected weekly for 16 wk and kept at —
15.0 °C. Field trials to evaluate the attractiveness of these lures were conducted in an apple
orchard in Moxee, WA beginning on 19 August and continuing for five wk. One replicate of
each lure loading and age was placed in a Pherocon IC trap in a completely randomized 30 x
30 m grid and hung at 3.0 m in the canopy. Each week 300 sterilized moths were released
within 20 m of each trap. Traps were checked after six nights and treatments were re-
randomized.

Seasonal comparison of commercial lures. The two proprietary high-load gray septa, CM
Megalure™ and the conventional-load CM L”™, and the standard CM 10X red rubber septa
(Trécé Inc., Salinas, CA) were field tested in an apple orchard treated with 1,000 Isomate C+
dispensers per ha from 18 May to 17 August 1999 near Moxee, WA. Lures were placed in
Pherocon VI delta traps and eight replicates of each lure type were evaluated in a completely
randomized 20 x 20 m grid. Moths were collected and counted, trap liners were replaced, and
traps were rotated one position each week. All three septa types were replaced on 6 July.
Additionally, the10X septa were replaced on 8 June and 26 July.

Statistical analyses. Linear regression analysis was used to fit the emission rate and
changes in the % E,EF isomer as a log decay curve with days aged in the field (Analytical
Software 2000). The effects of field aging and dispenser type on moth catch were tested with
analysis of variance (ANOVA). Moth count data were transformed (square root [x + 0.05])
to remove heterogeneity of variances before analysis. Means were separated in significant
ANOVAs with Fisher’s LSD. Differences in emission rate and isomeric purity on specific ages
for different dispensers were evaluated with Student t-tests. Linear regression of moth catch
on emission rate and % E,£ isomer were also conducted. The effect of dispenser age and their
initial loading rate for gray septa were evaluated with ANOVA. Specific comparisons were
made for 4.0,10.0, and 20.0 mg loads. These data were subsequently grouped and compared
with moth catch by 50.0 mg lures. The attractiveness of three proprietary lures was evaluated
during the season using ANOVA both on individual dates and for each moth generation.

RESULTS

Emission characteristics. The emission rate from new red rubber septa was nearly five-
fold higher than from new gray halo-butyl dispensers (Fig. 1a). The emission rate for each
dispenser fit a log decay curve with number of days aged in the field; gray lure: t = -4.9, df=
26, P < 0.001); and red lure: t = -8.5, df = 26, P < 0.001. Red septa retained a significantly
higher emission rate than gray septa for dispensers aged up to 21 d in the field (¢ values > 3.45,
df = 7, P < 0.05). The emission rate for the two dispensers aged from 28-42 d were not
significantly different (P > 0.05).

The isomeric purity of codlemone loaded in new dispensers was > 96% in both septa types
(Fig. 1b). The percentage of the £,E isomer remained unchanged in gray septa aged in the
field, t= -1.2, df = 26, P = 0.24. In comparison, the percentage of the EF, F isomer declined
over time in the red septa, ¢ = -4.6, df = 26, P< 0.001. The combined percentage of the E,Z
and Z,F isomers increased to nearly 20% after 2 wk and 30% after 4 wk.

Field aging (0 — 42 d) of lures was not a significant factor affecting lure attractiveness (F
= 1.47; df = 6, 56; P=0.21) (Table 1). However, significant differences in the attractiveness
of the two lures were found (F = 9.90; df = 1, 56; P = 0.003). No significant interaction
between lure type and age was detected (P = 0.43). Gray septa were significantly more
attractive than red septa for lures aged for 14 and 42 d (Table 1). Fourteen day-old red lures
had nearly three times the emission rate of gray lures (0.22 vs. 0.07 mg/d) (¢ = -4.86, df= 5,
P =0.003), and a significantly lower isomeric purity (82.1 vs. 96.2% E,E isomer) (t = 3.47,
df = 5, P =0.03). The emission rates of 42 d-old red and gray lures were low (0.1 mg/d) and

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 ly

A. @ Gray 4 mg septa

5 O Red 10mg septa

x rate=0.47exp(-0.07days); r2=0.73
O
8 \ ;

rate=0.08exp(-0.04days); r2 = 0.48

Emission rate (mg/d)

B. %=96.5exp(-0.0003days); r2 = 0.05

%E,E isomer

%=92.8exp(-0.0056days); r2 = 0.45

0) 10 20 30 £40
Number of days aged in field

Figure 1. Emission rate (A) and percentage of the £,F isomer (B) from field-aged gray halo-
butyl elastomer and red rubber septa loaded with 4.0 and 10.0 mg codlemone, respectively.
Some data points overlap, n = 4 per date.

Table 1
Attractiveness of field-aged gray halo-butyl elastomer and red rubber septa loaded with
codlemone tested during July 1996.

Days aged in the Mean no. (SE) moths per trap ___ Statistical analysis
field prior to test Red septa Gray septa t-value P-value

0 6.4 (2.1) 5.8 (2.6) -0.05 0.96

i 8.2 (3.6) Li61(G.5) 0.83 0.43

14 1.6 (0.6) 11.6 (3.8) 7.93 0.02

21 3.0 (1.0) 7.4 (3.4) 0.60 0.56

28 1.0 (0.3) 8.2 (2.5) 2724 0.05

35 2.6 (1.2) S222) 0.42 0.68

42 2.2 (0.5) 7.0 (1.8) 2.39 0.04

Red rubber septa and gray halo-butyl elastomer septa were loaded with 10.0 and 4.0 mg
codlemone, respectively.

did not differ between lure types (¢ = -0.28, df = 6, P = 0.79), but the percentage of the E,E
isomer was again significantly different (76.0 vs. 94.2% for the red and gray lures,
respectively, t = 3.48, df =5, P= 0.03).

The mean number of moths caught per trap with either lure was not linearly correlated to
emission rate (gray: t = -0.22, 7 = 0.01, df= 5, P = 0.83; red: t= 1.49, r = 0.31, df=5, P
=0.20) (Fig. 2a). A significant difference in moth catch was found between lure types when

128 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

0.0 0.1 0.2 G3 0.4
Emission rate (mg/d)

16
number of moths = -23.8 + 0.33(%)

. 2
= 0.66
12 =

Number of moths caught per trap

15 80 85 90 95 100
% EE isomer

Figure 2. Number of moths caught per trap as a function of emission rate (A) and percentage
E,E isomer (B) in gray halo-butyl elastomer and red rubber septa loaded with 4.0 and 10.0 mg
codlemone, respectively.

the data were pooled across similar emission rates (gray septa 0.01 — 0.08 mg/d and red septa
0.01 — 0.13 mg/d), t= 4.52, df= 8, P= 0.004. Within this lure emission range, moth catch in
traps baited with gray versus red septa was more than three-fold higher (Fig. 2a).

A significant linear relationship was found between mean moth catch and the percent E,E
isomer for red septa, t = 3.23, df=5, r = 0.68, P = 0.02. No similar relationship was found
for moth catch and percent F,F isomer for gray septa, t = -1.18, df =5S, r = 0.22, P=0.29.
Combining the data for both lures generated a significant relationship between moth catch (y)
and the percent £,# isomer (x): y = -23.8 + 0.33x, t = 3.76, df= 12, r = 0.66, P = 0.0003.
These data fall into two distinct groups based on the percentage of EF, F isomer (Fig 2b). Lures
containing > 90% E,E isomer caught on average four-fold more moths than lures emitting <
85% E,£ isomer, t= 5.69, df= 12, P= 0.001.

Optimizing the Loading Rate of Gray Halo-butyl Septa. Dispenser age (F = 5.03; df
= 7, 128; P < 0.0001) and pheromone loading (F = 8.69; df = 3, 128; P < 0.0001) of gray
septa were both significant factors affecting moth catch (Table 2). The interaction between lure
age and pheromone load was not significant (P = 0.87). A significant curvilinear decline in
moth catch with lure age was detected with the nontransformed data, F = 4.72,; df= 1, 128 ;
P=0.03. Moth catch among the 4.0, 10.0 and 20.0 mg loads was not significantly different
(F = 0.17; df= 2, 128; P=0.85). Moth catch with the 50.0 mg lure was significantly different
from the other three lure loads (F' = 25.75; df= 1, 128, P< 0.0001) and declined linearly with
age (t = -2.71, df= 37; P =0.01). Moth catch by 50 mg lures aged for 2 wk was significantly
greater than with older lures and a curvilinear relationship was suggested (¢ = 1.96, df = 37;

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 129

P =0.57). Mean moth catch by gray lures loaded with 4.0 — 20.0 mg codlemone aged up to
10 wk did not differ significantly from moth catch by new 10X red rubber septa (Table 2; see
means lacking asterisks). In comparison, moth catch by aged 50.0 mg gray septa did not differ
from 10X red septa over the 16 wk.

Seasonal comparison of lures. The attractiveness of the two proprietary grey septa lures,
CM Megalure™ and CM L”™, and the CM 10X red rubber septa did not differ until the third
wk of the study (Table 3). During the third wk, moth catch was significantly higher in traps
baited with the Megalure™, and the L’™ lure caught more moths than the 10X lure. The 10X
lure was replaced on 8 June and no difference in the attractiveness of lures was found the
following week (17 June) (Table 3). Mean moth catches in traps baited with the 10X lure were
>75% lower than in traps baited with the Megalure™ for wk two and three (23 and 29 June)
though not significantly different (P values = 0.07-0.08). After three wk the 10X lure caught
significantly fewer moths than the Megalure’™™. All lures were replaced on 6 July and moth
catch was again significantly lower in traps baited with the 10X vs. the Megalure'™ after two
wk (19 July). The 10X lure was replaced on 26 July and caught significantly fewer moths than
the Megalure™ after two wk (10 August). The L”™ lure caught an intermediate number of
moths in comparison with the other two lures throughout the season (Table 3).

Table 2
Comparison of mean (SE) captures of codling moth in sticky traps baited with halo-butyl gray
septa loaded with varying amounts of codlemone in a sex pheromone-treated apple orchard
in Moxee, WA.

Pheromone Load (mg)

Lure age (wk) 4.0 10.0 20.0 50.0
2 222.7) 12.2 (4.7) 12:2, 18.0) 32.4 (7.1)
4 HOn 521) 10.6 (6.0) Seo aton) Loe 0e3)
6 6.0 (1.6) D618 3.5) On 2:5) 14.8 (2.2)
8 5765-287) 4.0 (1.9) 7.4 (3.2) 13.2 (4.0)
10 5.0 (1.4) 8.2 (2.0) 6.4 (4.2) 11:6 45:5)
12 OF (0:33) ZS" (2) 3.6* (2.4) 9.2 (4.2)
14 1.6* (0.8) 5.8 (2.0) 3.67 (1.0) 7.6 (3.2)
16 1.8* (0.8) 2.0* (0.7) 0.6* (0.4) 9.2 (4.8)

Mean catches followed by ‘*’ were significantly different (¢-test, df = 8, P < 0.05) from the
mean (SE) catch with new, red rubber septa loaded with 10.0 mg E8,E10-12:OH, 20.2 (6.6)
moths per trap.

DISCUSSION

The chemical instability of conjugated dienes, such as codlemone in sulphur-cured red
rubber septa is well known (Brown and McDonough 1986, Ideses and Shani 1988, Vrko¢ et
al. 1988). Yet, despite this general knowledge, a red rubber septum has been the standard lure
used to monitor codling moth for nearly 30 yr (Riedl et al. 1986). More recently, the use of
a 10.0 mg red septa for monitoring codling moth in orchards treated with sex pheromone
dispensers has been widely reported in North America (Knight 1995a, Judd et al. 1996, Gut
and Brunner 1998) and Europe (Charmillot 1990). Gut and Brunner (1995) found that these
high-load lures require frequent replacement and presumed this was caused by a rapid drop
in their emission rates under sustained warm summer weather. Our data suggest that the 10%
septa are attractive for 1- 2 wk; however, their rapid loss in attractiveness appears to be due
to the degradation of codlemone and not to a reduction in their emission rate. Similar results
emphasizing the importance of the chemical instability of conjugated dienes in rubber

130 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Table 3
Seasonal comparison of three commercial lures (n = 8) rotated weekly within a 12 ha apple
orchard treated with 1,000 Isomate C+ dispensers per ha.

Mean catch (SE)

Date traps Red septa___—Gray septa__ Gray septa _Statistical analysis _
checked 10X MegaLure ON [eee F,>, Pvalue
25 May 1.63 (0.68) 2.88 (0.95) 1.88 (0.58) 0.59 0.56
01 June 0.13 (0.13) 0.63 (0.18) 0.50 (0.27) 1.85 0.18
08 June 0.00* (0.00)a 2.25 (0.67)c 0.88 (0.23)b 11.00 <0.001
17 June 1.00 (0.33) 1.88 (0.40) 0.88 (0.64) 2.08 0.15
23 June 0.50 (0.27) ZAS W067) 0.75 (0.49) 2.94 0.08
29 June 0.13 (0.13) 1.00 (0.38) 0.50 (0.19) 2.98 0.07
06 July 0.13*(0.13)a 1.25*(0.41)b = 0.25* (0.16)a 4.95 0.02
Total 1“ flight 3.52 (0.82)a 12.02 (1.30)b 5.64 (1.03)a 12.81 <0.001
12 July 1.75 (0.75) 1.63 (0.57) 3.00 (0.89) 0.98 0.39
19 July 0.00 (0.00)a 1.63 (073)b 0.38 (0.18)ab 4.69 0.02
26 July 0.13*(0.13)a 4.25 (1.26)b 4.38 (1.84)b 6.02 0.01
03 August 3.25 (0.98) 4.50 (1.69) 5.00 (0.89) 0.60 0.56
10 August 3.13 (1.48)a 16.13 (3.89)b 6.75 (1.51)a 8.74 0.002
17 August 2.38 (1.69)a 10.75 (2.74)b 7.00 (3.70)ab 3.45 0.05

Total 2" flight 10.64 (2.52)a 38.89 (6.17)c¢ 26.51 (4.85)b 4.18 0.03
Traps were placed in the orchard on 5 May in a completely randomized 20 x 20 m grid. Traps
were moved one position within the grid each week. Lures of each type were replaced on dates
designated with a ‘*’. Data were transformed (sqrt(x+0.5) and subjected to ANOVA. Weekly
means followed by a different letter were significantly different, P< 0.05, Fishers LSD.

substrates were found with red rubber septa loaded with (E, E)-8,10-dodecadien-1-yl acetate,
the sex pheromone for the pea moth, Cydia nigricana (F.) (Horak et al. 1989).

The inhibitory effects of the isomeric blend of 8,10-dodecadien-1-ol on the attraction of
codling moth was first shown by Roelofs et al. (1972) and later by McDonough et al. (1993).
El-Sayd et al. (1998) demonstrated that this inhibitory effect was due to the presence of the
E,Z isomer. McDonough et al. (1993) suggested that this reduction in attractiveness would be
amplified in the high load lures used to monitor sex pheromone-treated orchards unless
codlemone was stabilized. Isomerization of codlemone can be minimized by adding
antioxidant or antiultraviolet components (Millar 1995) or by avoiding substrates containing
sulphur (Brown and McDonough 1986). Horak et al. (1989) increase the longevity of lures for
pea moth by using rubber substrates cured with organic peroxides instead of sulphur. Brown
and McDonough (1986) found that isomerization of codlemone in gray halo-butyl elastomer
septa was 4.7-fold slower than in red rubber septa. The gray halo-butyl septa appears to be a
more effective lure than the red rubber septa for monitoring codling moth in sex pheromone-
treated orchards due to this reduced rate of isomerization of codlemone.

The optimal loading rate of gray septa to monitor codling moth 1s difficult to determine
from these studies. Moth catch as a function of lure age was curvilinear across the range of sex
pheromone loads tested, with the highest moth counts for each pheromone load being found
in traps baited with two-wk-old lures. Gray septa loaded with 4.0 — 50.0 mg codlemone all
effectively monitored codling moth for at least 10 wk. The 50 mg lure caught significantly
more moths than the other three lures, and its attractiveness most closely matched the mean
moth catch by new 10X red septa over a 10 wk period. Residual studies of septa aged at 24
°C in the laboratory demonstrated that the half-life of codlemone in gray septa is 2.6-fold
longer than in red septa (unpublished data). The emission rate of alcohol sex pheromones from

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 131

rubber septa has been shown to be proportional to the amount of pheromone present, 1.e., first-
order process (Butler and McDonough 1981). Therefore, one can estimate that the emission
rates from gray septa loaded with 26 mg and a red septa loaded 10 mg codlemone should be
roughly equivalent.

Replacing the standard red rubber septa with the gray halo-butyl lure appears to be a
promising alternative for monitoring codling moth in sex pheromone-treated orchards.
However, this switch may require some changes in the established action thresholds for
supplemental insecticide sprays during both generations. Current recommendations provide
a narrow range (2.5-fold) of cumulative moth catch to trigger the need for supplemental sprays
(Gut and Brunner 1996). In comparison, cumulative catches of codling moth by the
Megalure'™™ were more than three-fold higher than with the 10X red lure during both moth
flights despite frequent lure replacements of the red septa (Table 3). Development of a codling
moth lure that can minimize the occurrence of ‘false negatives’ (absence of moth catch despite
the occurrence of fruit injury) is likely more important than having to change the catch
threshold for applying supplemental controls. Previous studies have examined the effects of
trap placement (Knight 1995b) and the proximity of the monitoring trap to the sex pheromone
dispenser on lure performance (Knight et a/. 1999). Optimizing the use of the gray septa to
provide long-lasting effective monitoring in sex pheromone-treated orchards may require
further refinements. Alternatively, development and testing of new lure substrates and
attractants could further minimize the risks associated with the use of sex pheromones to
manage codling moth.

ACKNOWLEDGEMENTS

I would like to thank Bill Lingren and D. Rajapaske Raja (Trécé Inc., Salinas, CA) for
providing the proprietary lures, Ken Bloem (Sterile Insect Rearing Facility, Osoyoos, British
Columbia) for providing the sterilized codling moths, John Turner and Kathi Johnson (USDA,
ARS, Wapato, WA) for assisting in the field work, and Dave Horton (USDA, ARS, Wapato,
WA) for help with the statistical analyses. This project was partially funded by the Washington
Tree Fruit Research Commission (Wenatchee, WA).

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management in Yakima Valley, Washington. Journal of the Entomological Society of Bntish Columbia 92:
29-38.

Knight, A. L. 1995b. Evaluating pheromone emission rate and blend in disrupting sexual communication of
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Maitlen, J.C., L.M. McDonough, H.R. Moffitt, and D.A. George, 1976. Codling moth sex pheromone: baits for
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McDonough, L.M., H. G. Davis, P. S. Chapman, and C. L. Smithhisler. 1993. Response of male codling moths
(Cydia pomonella) to components of conspecific female sex pheromone glands in flight tunnel tests. Journal
of Chemical Ecology 19: 1737-1748.

McNally, P.S. and M. M. Barnes, 1980. Inherent characteristics of codling moth pheromone traps.
Environmental Entomology 9: 538-541

Millar, J. G. 1995. Degradation and stabilization of E8,E10-dodecadienol, the major component of the sex
pheromone of the codling moth (Lepidoptera: Tortricidae). Journal of Economic Entomology 88: 1425-1432.

Riedl, H. And B. A. Croft. 1974. A study of pheromone trap catches in relation to codling moth damage. The
Canadian Entomologist 106: 525-537.

Riedl, H., B. A. Croft, and A. J. Howitt. 1976. Forecasting codling moth phenology based on pheromone trap
catches and physiological time models. The Canadian Entomologist 108: 449-460.

Riedl, H., J.F. Howell, P.S. McNally, and P.H. Westigard, 1986. Codling moth management, use and
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Roelofs, W.L., A. Comeau, A. Hill, and G. Milicevic, 1971. Sex attractant of the codling moth: characterization
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Roelofs, W.L., R. J. Bartell, A. S. Hill, R. T. Cardé and L. H. Waters. 1972. Codling moth sex attractant — field
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J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 133

A new species of Boreus (Mecoptera: Boreidae)
from Vancouver Island, British Columbia

DAVID C.A. BLADES
ROYAL BC MUSEUM, 675 BELLEVILLE ST., VICTORIA, BC, CANADA V8W 9W2

ABSTRACT

Boreus insulanus Blades (Mecoptera: Boreidae), a new species from Vancouver Island,
British Columbia, Canada, 1s described and distinguished from similar species using
morphological characters and measurements. Similarities of male genitalic structures,
the notched ninth sternum and male wings abruptly narrowed at the middle place this
new species in the brumalis subgroup of the nivoriundus group of Boreus species. The
brumalis subgroup currently includes B. brumalis Fitch, B. nix Carpenter, B. pilosus
Carpenter, and B. bomari Byers. Characters that distinguish Boreus insulanus from
these related species include: fine appressed hairs above the eyes, on the pronotum, and
wings; absence of fine, erect hairs on pronotum, wing and abdominal sclerites; relative
position and number of pronotal bristles (4 anterior; 4 posterior); and dark abdominal
sclerites with a metallic green sheen. The distributional records for this new species
indicate that it may be confined to Vancouver Island's interior mountain range.

INTRODUCTION

Species of Boreus, commonly called snow scorpionflies, inhabit mountainous regions
of Europe, northern Asia, and North America. They are usually associated with mosses
and are unusual in that the adults mate and disperse during the winter months (November to
March) (Penny, 1977; Hagvar, 2001). Specimens are often collected on snow during
sunny, warm days, but recent research on B. hyemalis (L.) shows that activity on the snow
surface is greatest on windless, cloudy days and that most activity during the winter occurs
in air pockets under the snow pack (Hagvar, 2001). Penny's (1977) comprehensive
systematic study of the family provides detailed descriptions and taxonomy of the known
species and the biology of the group. More specific studies on the biology of North
American species include works by Cooper (1974), Shorthouse (1979), and Courtin et al.
(1984). Boreus species described since 1977 include B. jacutensis Plutenko and B.
tardokijanensis Plutenko from Russia (Plutenko, 1984; 1985) , B. jezoensis Hori and
Morimoto from Japan (Hori and Morimoto, 1996) and B. bomari Byers and Shaw from
Wyoming, USA (Byers and Shaw, 1999).

This paper describes a new species of Boreus found in collections carried out by the
author on private property at Camas Hill, Metchosin, on the southern tip of Vancouver
Island, British Columbia (48° 23'57" N 123° 35' 44" W). Specimens were collected using
continuous pitfall trapping over a period of two years. Specimens of this new Boreus
appeared in samples from January to March 2000, and November 2000 to March 2001.
Literature research into the genus and inquiries to various collections in North America
failed to uncover any records of Boreus from Vancouver Island (Penny, 1977; Blades,
unpublished data). Searching of the Royal BC Museum collection uncovered two more
specimens from Vancouver Island. Using Penny's (1977) key and descriptions, original
descriptions (Carpenter, 1935), and literature on recently described species (Hori and
Morimoto, 1996; Byers, 1999), I determined that these specimens represented a new
species. This paper describes this new species of Boreus and provides diagnostic
characters to distinguish it from other nearctic Boreus.

134 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

Specimens were examined in detail and compared using measurements and
characteristics of external morphology. Measurements of overall body length were made
but the obvious distension of the specimens limits the usefulness of this character in
comparisons with other species, a problem also noted by Penny (1977). Instead, only
lengths and ratios of rigid structures such as the head, eye, wing, ovipositor and dististyle
were considered when making comparisons with other species. Characteristics of the male
genitalia in the new species were similar to those of B. pilosus Carpenter and B. nix
Carpenter. This similarity and the notched ninth sternum and male wings abruptly
narrowed at the middle place this new species in the brumalis subgroup of the nivoriundus
group of species as defined by Penny (1977). The brumalis subgroup includes B. brumalis
Fitch, B. nix, B. pilosus, and B. bomari Byers (Penny 1977; Byers 1999). Twelve hundred
and seventy eight specimens of ten related species (including paratypes) were obtained
from various North American collections for additional comparisons.

SPECIES DESCRIPTION

Boreus insulanus n. sp.
(Figs. 1 to 7)

TYPE MATERIAL

Holotype male: Canada. British Columbia: Vancouver Island, Metchosin, summit of
Camas Hill (293 m; 48° 23' 57" N 123° 35' 44" W), sample CH99-18P, yellow pantrap,
1311 to 13 III 2000, D. Blades, C. Reznechenko and L. Rosenblood. Deposited at the
Royal British Columbia Museum (RBCM); specimen number ENT000-0003 14.

Paratypes: Same location as holotype. 2 males, 1 female, 3 I to 13 II 2000; 3 males,
13 I to 13 III 2000; 1 male, 5 to 13 XI 2000 deposited at RBCM; 3 males 13 IJ to 13 III
2000 deposited one at each of: The Snow Museum (RBCM #ENT000-000310), University
of Kansas; the Canadian National Collection (CNC), Ottawa (RBCM #ENT000-000312);
and the California Academy of Sciences, San Francisco (RBCM #ENT0O00-000309).
Remaining specimens (6 males, 1 female; 9 XII 2000 to 8 HII 2001) retained in author's
collection, to be deposited at CNC by December 2004.

ADDITIONAL MATERIAL

Two female specimens with characteristics clearly matching those of type specimens:
one from Goldstream, near Victoria (48° 23' N 123° 33' W; pinned) collected by G.A.
Hardy, 7 III 1927 (RBCM #ENT991-164463); and one from north side of Mount Cokely at
1500 m, near Port Alberni ( 49° 14' 23" N 124° 35' 12" W ; in alcohol) collected by S.G.
Cannings, 23 ITV 1995 (RBCM #ENT998-010684).

ADULTS

General: A large species of Boreus, length of males 3.9 to 6.6 mm (avg 4.7 mm) (Fig.
1) and females 5.4 to 6.0 mm (avg 5.8 mm) preserved in alcohol, 5.0 mm for pinned female
(measured in lateral view from tip of ovipositor/sternum 9 of male to base of antenna).
Abdominal sclerites, wings and occiput dark brown to almost black with a green metallic
sheen. Thorax, legs and rostrum lighter, varying from tan to dark brown; the legs lighter
in colour than thorax. Colours of pinned specimen similar to those in alcohol. Immature
stages unknown. Terms used in this description follow those of Penny (1977) and Byers
(1999); it is important to note that setae and bristles (=strong setae) possess ring-shaped

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 135

6) 4 2mm

Figure 1. Lateral view of male Boreus insulanus indicating setae and bristles; hairs, and
setae of palps, omitted.

basal sockets whereas hairs and spines do not.

Head: Head coloration dark from top of head to just below eyes on sides and down
front of face almost to tip of rostrum. Lateral and posterior areas of rostrum tan to light
brown; an abrupt, horizontal separation of light and dark colours just below eye when
viewed in profile. Eyes reddish. Length of head from vertex to tip of rostrum: 1.57 to 2.00
mm (avg 1.81 mm) in males; 1.88 to 2.07 mm (avg 1.96 mm) in females. Diameter of eye
measured in same direction as head: 0.50 to 0.68 mm (avg 0.62 mm) in males; 0.58 to 0.67
mm (avg 0.63 mm) in females. Ratio of maxillolabial complex to rostrum varies from 0.89
to 0.97 (avg 0.93) in male and from 0.85 to 0.93 (avg 0.88) in female. Occipital region
glabrous, finely wrinkled and without hairs or setae. Area between eyes and occiput with
short, fine, recurved hairs closely appressed to the head. Fine, erect, white hairs dominate
the area between the eyes and below the antennal sockets. A few fine, pale, erect setae
present on base of rostrum (frons) below antennal sockets (about 0.06 mm in length). A
distinct row of 10-12 such setae in furrows (depressions) on either side of the otherwise
bare midline of rostrum. Mentum with 12 to 20 or more setae similar in length, but slightly
thicker, to those in lateral furrows; situated primarily in basal half, with two near distal
margin. Lateral ocelli nearly touching compound eyes with smaller median ocellus present
near bases of antennae. Antennae brown; with 20 to 22 (mode=20) flagellar segments in
males; 19 to 21 (mode=19) in females.

136 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

posterior

anterior

hindwing

}—__0.5mm_

Figures 2 to 7. 2) Lateral view of pronotum of Boreus insulanus showing position of
bristles and appressed hairs. Gray bristle indicates position of occasional intermediate
bristle. 3) Dorsal view of male Boreus insulanus hindwing and forewing. 4) Lateral view
of male Boreus insulanus forewing and hindwing. 5) Lateral view of male Boreus
insulanus genitalia. T=tergum; S=sternum. 6) Dorsal view of male Boreus insulanus
genitalia. 7) Lateral view of Boreus insulanus ovipositor and adjacent segments,
indicating position of setae, hairs and peg-like spines. Illustrations 3, 4, 5 and 6 by D.
Young.

Thorax: Anterior margin of pronotum with two dorsolateral bristles and usually two lateral
bristles; posterior margin with two dorsolateral, two lateral, and rarely, two sublateral
bristles (Fig. 2). Pronotum with two obscure transverse furrows and covered by short
recurved hairs, closely appressed to the surface. Mesonotum, metanotum and lateral
thoracic sclerites without conspicuous bristles or hairs in male. Females with a pair of
short, cruciate bristles near center of mesonotum.

Legs: Colour of coxa same as adjacent lateral sclerites; remaining leg segments lighter

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 137

brown to yellowish. Stout, dark brown, apicofemoral spine present. Anterior faces of
coxae and trochanters with abundant long white hairs. Remaining leg segments covered in
short, fine hairs and armed on ventral surface with two rows of evenly spaced, short, fine,
erect setae. Each tibia with two apical spurs. All tarsi with two simple, terminal claws.
Ratio of lengths of foreleg to midleg to hindleg approximately 1 : 1.3 : 1.7. Hindleg
approximately 1.5 times body length in males; in female about equal to length of body
including ovipositor.

Wings: Male forewings darker brown than thorax; outer margins distinctly narrowed
near middle in dorsal view (Figs. 3 and 4). Length of wing 1.60 to 2.13 mm (avg 1.83
mm). Each forewing bears 17 to 24 (mode=18) strong, black, slightly twisted, outer spines
starting about one-third of length from base and increasing in size from 0.05 mm to 0.11
mm at the wing tip. Inner forewing spines similar in shape, size and position to outer
spines, and numbering between 17 and 26 (mode=19) on each wing. Terminal forewing
spine strongly incurved and about the same length as pronotal bristles (0.16 mm).
Forewing rugulose and covered by fine hairs, recurved and closely appressed to the dorsal
surface. Hindwing light brown and tubular in basal four-fifths and becoming black and
flattened at point of sharp incurving to tip. Middle portion of hindwing bears 7 to 12
(mode=10) short, stout spines with longer fine hairs intermixed. Distal half of hindwing
before curved tip with felt-like pad of short, yellow to orange hairs. Female forewings
short (avg 0.4 mm), oval, brown and covered with fine hairs, recurved and closely
appressed to the surface. Hindwings very small and completely overlapped by the
forewings.

Abdomen: Terga and sterna dark brown to almost black with metallic green sheen and
densely covered in short, fine, semi-erect, posteriorly directed hairs that are clearly visible
when backlit. Intersegmental membranous areas white. All segments with unfused terga
and sterna. Female abdomen clearly larger in diameter than male but otherwise similar for
segments | to 7.

Male genitalia (segments 8 to 10): Segment 8 unmodified, similar in appearance to
segment 7 (Fig. 5). Tergum 9, sternum 9, basistyles and dististyles very dark brown to
black. Sternum 9 long, extending to junction of basistyle and dististyle, and bearing a
shallow, but distinct, notch along the posterior margin. Tergum 9 in profile with low hood
that acts as a receptacle for the tips of the dististyles. In dorsal view the hood appears as a
U-shaped ridge, forming a cup-like invagination, surrounded posterolaterally by the
denticles (Fig. 6). The hood bears long, fine, pale hairs on the inside vertical surfaces (also
visible in lateral view) and a pair of median bare areas, separated by a septum, where the
dististyles touch. Denticles and surrounding cuticle black and difficult to see, especially
when dististyles retracted into hood. Number of denticles ranged from 17 to 32 (mode=20)
on each of the denticular areas of tergum 9. Dististyle with sharp tip, a medial, thumblike
projection and a medial row of 14 to 21 (mode=21) short spines along inner margin.
Tergum and sternum 10 small, oval, brown sclerites each bearing a few fine hairs.
Membranous gonopore situated ventrally to the more prominant anus and usually hidden in
the retracted state.

Female genitalia (segments 8 to 11): Configuration of terminal segments similar to
females of other Boreus species (Fig. 7). Tergum 8 with fine transverse wrinkling and
nearly encompassing segment 8, the sclerite ending just before venter where the valvulae
emerge. Tergum 9 smooth and shorter than preceeding and following segments. Segment
10 about 1 mm in length, equal to the length of the maxillolabial complex, and lacking
hairs or setae on dorsal surface. A few scattered setae present on dorsal surfaces of
segments 8, 9 and 11. Segment 11 and cerci fused into short, triangular segment. Ventral
Ovipositor valves (valvulae of segment 8) with numerous long, fine, white hairs on basal

138 J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002

2/3 to 3/4; replaced by about 30 to 40 evenly spaced, short, peg-like spines in terminal
region. These spines lack emergent terminal hairs evident in other species (Penny, 1977;
Byers, 1999) but longer hairs are occasionally present between the peg-like spines.

DIAGNOSIS

Presence of pronotal spines, notched sternum 9 of male, male wings narrowed near
middle and configuration of male genitalia place this species in the brumalis subgroup with
B. nix, pilosus, brumalis, and bomari. It is distinguished from these other species by the
following combination of characteristics: head and abdomen dark brown to black with a
distinct metallic green sheen, wings darker than brown thorax, legs light brown to
yellowish and clearly lighter than thorax; fine hairs above eyes, on pronotum, and wings,
recurved and appressed to surface; pronotal margins with two or four anterior and four
posterior bristles, these bristles strong, dark and slightly curved; pronotum lacking erect,
pale, intermediate length setae or hairs; distinct, erect hairs also absent from wings,
occipital area, and sclerites of thorax and abdominal segments 1 — 8; 17 to 24 outer
forewing spines; 17 to 26 inner forewing spines; 17 to 32 denticles; 14 to 21 dististyle
spines; antenna with 19 to 22 flagellar segments; ovipositor equal in length to maxillolabial
complex; tergum 10 of female without dorsal hairs or setae; terminal peg-like spines of
valvulae without emergent hairs.

ETYMOLOGY

The latin word for “islander”, insulanus, was chosen to indicate the apparent restriction of
the species’ range to Vancouver Island.

DISTRIBUTION AND HABITAT

This species is apparently confined to Vancouver Island and is currently the only
Boreus species known from the island (Penny 1977 ; Blades unpublished data). Collection
locales are situated along the interior mountain range of Vancouver Island at elevations
from 300 to 1500 m. Like other Boreus species, it is believed to be associated with
mosses.

ACKNOWLEDGEMENTS

I would like to thank Catherine Reznechenko and Lorne Rosenblood for hosting my
study of insect diversity on their property at Camas Hill, Metchosin. I would also like to
thank David Young for his contributions to the illustrations in this paper and Robert
Cannings for his helpful comments on the manuscript and his moral support. Thanks are
also extended to the many people who responded to my requests for distributional records
and specimens from their collections — full acknowledgements to these people and their
institutions will appear in an upcoming paper on Boreus distributions in western North
America. I would also like to thank Richard Zack for supplying specimens of another
undescribed species of Boreus during preparation of this description.

REFERENCES

Byers, G.W. & S.R. Shaw 1999. A new species of Boreus (Mecoptera: Boreidae) from Wyoming.
Journal of the Kansas Entomological Society 72(3): 322-326.

Carpenter, F.M. 1935. New nearctic Mecoptera with notes on other species. Psyche 42: 105-121.

Cooper, K.W. 1974. Sexual biology, chromosomes, development, life histories and parasites of Boreus,

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 139

especially of B. notopterates. A southern California Boreus. Part II. (Mecoptera: Boreidae). Psyche
81: 84-120.

Courtin, G.M., J.D. Shorthouse, & R.J. West 1984. Energy relations of the snow scorpionfly, Boreus
brumalis (Mecoptera) on the surface of the snow. Oikos 43: 241-245.

Hagvar, S. 2001. Occurance and migration on snow, and phenology of egg-laying in the winter-active
insect Boreus sp. (Mecoptera). Norwegian Journal of Entomology 48: 51-60.

Hori, S. & Morimoto, K. 1996. Discovery of the Family Boreidae (Mecoptera) from Japan, with
description of a new species. Japan Journal of Entomology 64(1):75-81.

Penny, N.D. 1977. A systematic study of the family Boreidae (Mecoptera). University of Kansas Science
Bulletin 51(5): 141-217.

Plutenko, A.V. 1984. A new species of the genus Boreus (Mecoptera: Boreidae) from the Soviet Far
East. Zoologicheski1 Zhurnal 63: 778-781. (In Russian)

Plutenko, A.V. 1985. New and little-known species of Mecoptera from the Soviet Far East.
Entomologicheskoe Obozrenie 64: 171-176. (In Russian) [Entomological Review 64: 113-119]

Shorthouse, J.D. 1979. Observations on the snow scorpionfly, Boreus brumalis Fitch (Boreidae:
Mecoptera) in Sudbury, Ontario. Quaestiones Entomologicae 15: 341-344.

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 141

Erratum

Insect population ecology in British Columbia

J.H. MYERS

DEPARTMENT OF ZOOLOGY, UNIVERSITY OF BRITISH COLUMBIA
6270 UNIVERSITY BLVD, VANCOUVER, BC, CANADA V6T 1Z4

D.A. RAWORTH

AGRICULTURE AND AGRI-FOOD CANADA,
PACIFIC AGRI-FOOD RESEARCH CENTRE, AGASSIZ, BC, CANADA VOM 1A0

In Volume 98, pp. 107-108, the historical account relating to the work of Neil Gilbert
and collaborators was not correct. Neil wrote the following response:

‘After cabbage aphid and Masonaphis, I thought it might be possible to introduce some
limited generalization into population dynamics by constructing a Universal Aphid of
which every aphid species would be a particular case. But pea aphid put paid to that. In
the first two species, predation could be represented by a simple formula because it only
took surplus production of aphids and was more or less compensated by density-dependent
reproduction. In pea aphid, the coccinellids drove aphid numbers down low, so we either
had to study predation in detail or admit defeat. It was all very messy but proved possible
to predict predation rates, but not the numbers of beetles entering the field. That would be
possible only if you knew the dynamics of the whole local ecosystem, an impossible task.
In other words, the aphid population couid not be isolated from the rest, even to a first
approximation. The whole thing was a failure as far as I was concerned, although a very
instructive one.’

J. ENTOMOL. SOC. BRIT. COLUMBIA 99, DECEMBER 2002 143

Erratum

Behavioural and chemical ecology in British Columbia

B. ROITBERG AND G. GRIES

CENTRE FOR ENVIRONMENTAL BIOLOGY,
DEPARTMENT OF BIOLOGICAL SCIENCES, SIMON FRASER UNIVERSITY,
8888 UNIVERSITY DRIVE, BURNABY, BC, CANADA VSA 1S6

Volume 98, p 113, paragraph 5, lines 1-5 should read:

The second major development was the formation of the Pestology Centre at Simon Fraser
University (SFU) in 1967. Several members of the centre focused on behavior including,
Bert Turnbull (predators), John Borden (host and mate-seeking behaviors), Peter Belton
(acoustic and oviposition behaviors in mosquitoes), Manfred Mackauer (behavior of
parasitoids) and Bryan Beirne (behavior of biocontrol agents).

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Journal
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of British Columbia
Volume 99 Issued December 2002 ISSN #0071-0733

Directors of the Entomological Society of British Columbia 2000-2001...................:::cceee 2

Hamilton, K.G.A. Homoptera (Insecta) in Pacific Northwest grasslands. Part 1 — New and
revised taxa of leafhoppers and planthoppers (Cicadellidae and Delphacidae) ............ 3

Hamilton, K.G.A. Homoptera (Insecta) in Pacific Northwest grasslands. Part 2 — Pleistocene
refugia and postglacial dispersal of Cicadellidae, Delphacidae and Caliscelidae ....... 33

Raworth, D.A. and M.C. Robertson. Occurrence of the spider mite predator Stethorus
punctillum (Coleoptera: Coccinellidae) in the Pacific Northwest ................ (NOTE) 81

Furniss, M.M., E.H. Holsten, and M.E. Schultz. Species of Alaska Scolytidae: Distribution,
hosts, and historical TEVICW .i:..cc..:2ccseneorue sovcn onc ovodovavcdenccer ssauceseaece ae geeete a eee 83

McGregor, R.R., R.P. Prasad, and D.E. Henderson. Searching behaviour of Trichogramma
wasps (Hymenoptera: Trichogrammatidae) on tomato and pepper leaves................... 93

Kenner, R.D. Stylops shannoni (Stylopidae, Strepsiptera): A New species for Canada, with
commments on XeNOS, DECK ..:.i.00ccvec Bc cwcetetnses a teeceetne cee ae ee ee i oo

Miller, D.R. Short-range horizontal disruption by verbenone in attraction of mountain pine
beetle (Coleoptera: Scolytidae) to pheromone-baited funnel traps in stands of lodgepole
tae ee cee rec ee cay hae See doe nce cece (NOTE) 103

Knight, A.L., D. Larson, and B. Christianson. Flight Tunnel and Field Evaluations of Sticky
Traps for Monitoring Codling Moth (Lepidoptera: Tortricidae) in Sex Pheromone-treated
Orchards ...:scecccsesen cece loses sacescogdeet dented denea deits soe Un euieyaleeteesaae HONGO Oh Reels Sass eae 107

Morewood, W.D. and K.E. Simmonds. a-Pinene and ethanol: Key host volatiles for
Xylotrechus longitarsis (Coleoptera: Cerambycidae) ..........22..c...c0:0:-ecceesseceeeeesesee = 117

Knight, A.L. A Comparison of Gray Halo-butyl Elastomer and Red Rubber Septa to Monitor
Codling Moth (Lepidoptera: Tortricidae) in Sex Pheromone-Treated Orchards ...... 123

Blades, D.C.A. A new species of Boreus (Mecoptera: Boreidae) from Vancouver Island,
British Columbia ooo. cce.c.. cc seewwesseadbadueeeisaerspees eee beatae yee eee eee eee ee

Erratum: Insect population ecology in British Columbia ..................ceeecceeeeeceeeeeneeeeeeeees

Erratum: Behavioral and chemical ecology in British:Columbia::.:).).3. 3503 aes

NOTICE TO CONTRIBUTORS ooo ooo i ee ee

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