Table Of ContentPROC. ENTOMOL. SOC. WASH.
108(4), 2006, pp. 964-976
DIVERSITY OF SYRPHIDAE (DIPTERA) IN CENTRAL
APPALACHIAN FORESTS
CYNTHIA J. FRITZLER, CRYSTAL B. MAYLE, JOHN S. STRAZANAC, AND LINDA BUTLER
Plant and Soil Sciences4Entomology, P.O. Box 6108, West Virginia University,
Morgantown, WV 26506-6108, U.S.A. (e-mail: [email protected])
Abstract.4Species richness and abundance of Syrphidae in the central Appala-
chian forests was sampled with Malaise traps from 1995 through 2001. Syrphids
were identified from the George Washington National Forest, Virginia, and the
Monongahela National Forest, West Virginia. Of the 131 species identified,
Eristalinae were the most species rich subfamily making up 61% of the total
richness (80); Syrphinae were the most abundant making up 66% (5,138) of the total
sampled abundance; Microdontinae with 3% of the total richness (4) represented
8% (587) of the total sampled abundance.
Key Words: Syrphidae, central Appalachian forests, species richness, diversity
Syrphidae are one of the more diverse A project was initiated in 1995 to
families of Diptera with nearly 6,000 study the effects of biopesticide control
species described worldwide; approxi- of gypsy moth (Lymantria dispar (L.))
mately 870 species in 88 genera occur in (Strazanac et al. 2003a, Strazanac and
the Nearctic Region (Vockeroth and Butler 2005) on nontarget organisms.
Thompson 1987). Adult syrphids rank Pollinators were a part of the original
second only to bees (Hymenoptera: study plan which included a defoliation
Apoidea) as important flower-visiting treatment caused by gypsy moth. While
and flower-pollinating insects (Larson gypsy moth was effectively removed due
et al. 2001). Syrphid larvae have highly a pathogenic fungus, limiting defoliation,
diverse feeding habits (Duffield 1981, important pollinators, including 4 syr-
Maier 1982, Vockeroth and Thompson phids, continued to be tallied. From the
1987, Rotheray 1993); Syrphinae include sampling of adult syrphids we deter-
important predators of Homoptera and mined their richness and abundance
other soft bodied insects such as young within the broadleaf-pine forests in the
caterpillars, Eristalinae are primarily adjacent George Washington National
saprophagous, with some exceptions 1n- Forest (GWNF) and Monongahela
cluding phytophagy, and predation with- National Forest (MNF) in the central
in the Pipizini, and Microdontinae are Appalachians.
predators on ant broods. In America
MATERIALS AND METHODS
north of Mexico, species recognition
relies primarily on the more conspicuous Eighteen 200-ha_ study plots were
adults, approximately 7% of which located in the GWNF and the MNF
have their immature stages associated (Strazanac 8et aly 2003a)" (Fig! 1) Phe
(Thompson 1990). GWNEF study plots were along the
VOLUME 108, NUMBER 4 965
west with the collecting head up slope.
Fifteen weekly samples were taken early
May through mid-August from 1995
through 2001. Malaise sampled Syrphi-
dae were sorted, tallied, and stored in
70% ethanol. Material was later pinned
and processed through increasing con-
centrations of ethanol that was replaced
with ethyl acetate before air drying.
Syrphids from 1995 through 1999
(7,763) were fully processed and identi-
fied. Syrphidae from 2000 and 2001
(3,993) were tallied only.
Generic determinations were made by
CJF and CBM using Vockeroth and
Thompson (1987). Species-level identifi-
cations were facilitated for the authors
(CJF, CBM) by F. C. Thompson and his
compilation of keys to most eastern
Syrphidae (unpublished). In addition,
Fig. 1. Appalachian 4 broadleaf-pine forests the works of the following authors were
(after Keys et al. 1995) with study plot locations
regularly referenced: Eristalinae, Coo-
in the George Washington National Forest, Vir-
vert (1996), Coovert and Thompson
ginia, and in the Monongahela National Forest,
West Virginia. (1977), Curran (1921); Microdontinae,
Thompson (1981); and Syrphinae, Knut-
southeast slope of Great North Moun- son (1973), Vockeroth (1980, 1983,
tain in Augusta Co., VA, centered at 1986a, b, 1990, 1992). Species confirma-
38°07'30"N, 79°22'309"W, with elevations tions were made using the collection
of 561-744 m. The MNF study plots at the National Museum of Natural
were in Pocahontas Co., WV, in groups History, Smithsonian Institution, Wash-
of three on three different mountains, ington, DC with the guidance of F. C.
centered at 38°159N, 80°009W, with Thompson.
elevations of 80541,232 m. Oaks (Quer- Voucher specimens are deposited in
cus spp.) dominate both forest, but the West Virginia University Arthropod
species of hickories (Carya spp.), pines Collection, Morgantown, WV, and the
(Pinus spp.) and maples (Acer spp.) are National Museum of Natural History,
also abundant (Keys et al. 1995). A Smithsonian Institution, Washington,
highly diverse understory of herbaceous DC.
plants (Strazanac et al. 2003a) dominates
the forest floor in the spring until the RESULTS AND DISCUSSION
deciduous canopy fully leafs out. From 1995 through 2001, a total of
A randomly placed 30-ha subplot was 11,756 Syrphidae were sampled in the
established on each study plot. Two GWNE and MNF. The relative trend in
Townes-style Malaise traps (Townes year to year abundance (Fig. 2) for the
1972) were set up at different elevations two forests were similar with the excep-
on each subplot usually with one on tion of years 1999 and 2001 when the
a ridge and the other in a valley. Traps sampled abundance was much greater in
were placed on southerly-facing slopes or the MNF. In 1999 there were relatively
ridges, and the trap length oriented east4 large increases in abundance of Syrphus
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
GWNF MNF
Abundance
1395 1996 Ago 1998 1999 2000 2001
Fig. 2. Total sampled abundance of Syrphidae from 199542001 in the George Washington National
Forest (GWNF), Virginia, and in the Monongahela National Forest (MNF), West Virginia.
species (Syrphinae) in both forests and reached relatively high numbers in the
Microdon craigheadi Walton (Microdon- GWNEF only.
tinae) in the GWNEF (Fig. 3). Syrphidae Eristalinae was the most species-rich
for 2001 are unsorted so the relative subfamily in the GWNF and MNF com-
increase of the group or groups involved bined, making up 61% of total richness.
are not known. Total abundance from The more abundant Syrphinae made up
1995 through 1999 of Syrphinae in the 36% of the total species richness. Mate-
GWNEF and MNF combined was 66% rial identified through 1999 represents
compared with 26% for Eristalinae. a total of 131 species: species richness
Representatives from 15 tribes (Vock- was greater in the MNF with 118 species
eroth and Thompson 1987) were found collected; 106 species were collected in
in each forest (Table 1). The species-rich the GWNEF (Fig. 4). Distinct differences
Syrphini (35), Milesiini (36), and Bra- between study plots, such as, elevation,
chyopini (23) shared the greatest number rainfall, and vegetation may influence
of species between forests, 26, 22, and 17 the sampled richness. Other major taxa
respectively, while the remaining tribes also had greater species diversity in the
are represented by | to 7 species most MNF compared to the GWNF plots.
of which were shared between forests. Five years (1995-1999) of Malaise sam-
Species richness was not a predictor of pled Symphyta (8,884 adults) produced
the relative sampled abundance within 155 species (Strazanac et al. 2003b,
each tribe. Abundance (1,155) of Tox- Braud et al. 2003) with 127 species from
omerini with three species was exceeded the MNF and 104 from the GWNF.
only by Syrphini abundance (3,602) Similar species richness trends were also
made up of 35 species. Milesiini abun- noted during this long-term study within
dance (618) was third followed by 4 the Lepidoptera (Butler et al. 2001) and
species of Microdontini with a sampled Carabidae (Carrington 2002).
abundance of 587. While Toxomerini All sample years began with high
and Microdontini have similar species abundance; species richness and _ total
richness, they represented different dis- abundance generally peaked by the third
tributional patterns. Toxomerini species week of sampling in the GWNEF and the
were widely distributed in both forest fourth week in the MNF. Most species
and in all the plots while Microdontini were first collected in May (Table 2) and
VOLUME 108, NUMBER 4 967
600
G. Washington N. F.
GM Eristalinae [Hl Microdontinae [J Syrphinae
Ww( ep)( ep)
Abundance
2000
| Monongahela N. F. Re. |
500
400
(ee)( =)( ep)
Abundance
1995 1996 1997 1998 1999
Fig. 3. Total sampled abundance of Syrphidae by subfamily from 1995-1999 in the George
Washington National Forest, Virginia, and in the Monongahela National Forest, West Virginia.
those shared between forests typically overlapping adult flight periods. Micro-
first appeared in the GWNF one week don tristis was collected early in the
earlier than in the MNF. Declines in season while M. craigheadi was most
counts of individual species started abundant during the last third of our
earlier in the GWNF, for example, sampling period. Within more abundant
Syrphus torvus Osten Sacken (Syrphinae) species, peaks in male abundance often
quickly declined in mid-May from the occurred before peaks in female abun-
GWNYF, then early June in the MNF. dance (e.g., M. craigheadi, Epistrophe
Seasonal occurrence and abundance for nitidicollis Meigen (Syrphinae), Syrphus
some species, including 7oxomerus gemi- knabi Shannon (Syrphinae)). Males were
natus (Say) (Syrphinae), Toxomerus mar- also typically collected during a shorter
ginatus (Say), Copestylum vesicularium period as was especially evident in
(Curran) (Eristalinae), and Chrysotoxum Chalcosyrphus libo (Walker) (Eristali-
derivatum Walker (Syrphinae), were nae), Dasysyrphus venustus (Meigen)
fairly consistent throughout the 15-week (Syrphinae), and Epistrophe emarginata
sampling period. The two more abun- (Say) (Syrphinae).
dant Microdon species (Microdontinae) Tables | and 2 are conservative in pre-
in the GWNF, M#. craigheadi Walton senting species richness. Six specimens
and M. tristis Loew, had mostly non- tentatively identified as Eupeodes cana-
968 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Sampled richness and abundance from 1995-1999 of Syrphidae by tribe in the George
Washington National Forest (GWNEF), Virginia, and in the Monongahela National Forest (MNF),
West Virginia.
Species Richness Abundance
Subfamily Tribe GWNF MNF Total Shared GWNEF MNF Total
Eristalinae
Brachyopini 17 23} 23 17 87 238 325
Callicerini | 1 1 1 12 10 22
Cerioidini ] ] 1 l 16 3 19
Eristalini 2 3 3 2 10 20 30
Merodontini l | 1 1 22 4 26
Milesiini 29 29 36 22 TTS 343 618
Pipizini 6 6 6 6 106 269 aS
Rhingiini 6 5 6 5 186 165 85)
Sericomyiini | ] ] ] 9 51 60
Volucellini 2 1 Dy; l 183 29 PW
Microdontinae
Microdontini 3 2 4 ] 579 8 587
Syrphinae
Bacchini 4 fl 7 4 48 266 314
Paragini 2 2 D D, 49 18 67
Syrphini 28 34 35 26 995 2,607 3,602
Toxomerini 3 2 3 D 525) 630 SS
densis (Curran), Eupeodes confertus weeks ending in mid-May, approximately
(Fluke) and Eupeodes luniger (Meigen) the time our sampling started. Criorhina
are tallied as Eupeodes sp. (Syrphinae). was not sampled in our survey.
Characters used in species determina- Based on the five years of identified
tions for this group may be influenced by material, species accumulation may be
temperature differences experienced dur- approaching an asymptote in both the
ing the pupal stage (DuSek and Laska GWNEF and MNF (Fig. 5). If true, the
1974). The following groups are taxono- MNF does have a richer fauna with 118
mically difficult, species differentiation is species compared to GWNF9s 106 spe-
based on male genitalia and secondary cies. Chao estimators (Colwell 1997)
sexual attributes: Heringia (Eristalinae), suggest the two forests may ultimately
Paragus (Paragus) (Syrphinae), and Pla- prove to be more similar and species rich.
tycheirus (Syrphinae). These genera were Chao 1 and Chao 2 predict GWNF to
identified as far as current literature
permitted. 150
Species counts peaking in the early Py
weeks of our annual sampling period tee a Total
90 Wh 92 » Shared
may indicate that some taxa may be
under-represented or missed. Sampling
60
29 April through 27 September 19914
1993, Cicero and Barrows (2000) collected SRpiecchinee3ss0 s
Criorhina nigriventris Walton and C. 0
verbosa Walker (Eristalinae: Milesiini) in . Te ees 8
Malaise traps in a similar Appalachian aR. Specs SOs smpiigae from
1995-1999 in the George Washington National
habitat at a more northern latitude. These 4 Forest (GWNF). Virginia, and in the Mononga-
species were sampled during the first few hela National Forest (MNF), West Virginia.
V
8Ioquinu = N IqQeL <7 PpoyRsuoucy apsjdueg qsuA[RUONRN aijrr ouaedsy qns ssouyosu 18°04 pur <(NIA) ISOA\ souvpunge Jo <PIUIsITA WS seprydsdg 1yS4 PolSlaOd PWor porod N soyep 666[-S66| oie uoqsUl OuNise] r y<Is4 WeoY) nSy Avp PoJo llesod[ Bd109H PUulaAep-7 .f N Surdures UOI;UIYSeAA 1314 8sporsiod P[PUONPN olSoeung [ RI = 189104 N 8oung [ng PpeyO= N<(4NMD) R s<u41o 4u301p8A[ng¢ 4y POLl<PIUISITA sny esdoy P= UulPUP ay NUl snsny oy) OLUME 108, N
ALSVTNAI IV U
M
rurdoAyorig B
E
R
pdodyonilga yvapp uosuyor APIN 974-E], 7 API 6C-07 S ABW 9T-LI ih APIN LT-81 b
pdodyopig suarsaan]{ uouueys API 07-8 9 APY ung4QOQ]] 8 Avy ung4zTLI] T ZI APP unf4¬OTQ ] 8 4
vdodyovvaigp jou uajsQ uayorg uneZT I APL UNf4O-EO ] S APY IngS4$- O] 74
pdodyovigp xajdiad ueling Avpy ung4p-lg ] v Avy ung4p[l ¢ 74
NdodYyoNDIADGA udIsO udyoRSg ABWO T I AryQ T I
AAISDBOSAASYNDA YIMUY IOY[VM APIO T I KEWL Z I KEWL T E AvP] ung4¬sT [ Ik
DIdIIODAAs IU MOOT Avy ung49g7] 8 une 6I-ZI T
VV DIdaIOA 1IDI1814M18O OT APIN 6I-sS Z APIN 97-Z1 9 ABWO T I
daDji odw Sadiapa MOOT Av ungL4 -Lz T API [ng9- ZI TI ARN6 7 I ALI UNg4O¢L¬z 9
vIISpOAN VSOGO]s (1DY[VM) ABWL I I
DIISPOANT 221//D1AWU (UOIST]IIMA) ABW SI-O1 6
DAAAUOYVIPAICI U (UURWUSPIOAA) AP 97 I APIN 67-61 ¬
puisayds sisualyapjJpIdDdAyO OD ARIO T I API UNL46-cL Z ¢
vuisayds Vipjnuunigy ool[ey, UNL 6C-CT Z me IZ I une 6c I
puisaydy 4aisps[dNyHo pag ARIS T I AP SI-OL G API 97-9 lI
Dulsayds YIVJNUYdiUYu os}Iaqoy AVP Ing¢§- zz ¬ Av Ing-srgly, Ol Av [ng¬4 sz L APIN Inf4-6017 T GG
puisayds vupuiany{ yore ung Ss, I ung [oeZ- 6| S APIN 6C-ST of Ing snyO-lL 9
vuisaydsy Vipjnovuoaynoyofy [eW, une Q[ I API 6c74-S 9I Av ung6- ZI G
DUuIsayds VUVIUIAY UOISTI[IAA ung g] I Avpy ung4pgl y G Avy yngZL4 s SI
DUIsaYydVsI DGO] M907 APW6 c I
puisayds VAafijngo] YOol[R] APIS T I AR¢I I Av un¢g 4Zz 72
yusaydgy vinjoied noy[InboD Avy ung49Tg] ung Inf-L9Iz API Ing49ccz CZ AVW Ime-I8Z1 6C
Dulsayds SLJUaALM[NOAO T Ae/Iw] I APIA 97-8] Z Are¬ ] I
I[UeLSI DOI]
DAIIIIDI VIIDAMI (1OY[VMA) Ar QI-s ZI KEW L7-S 9 API 6c-Ol P
TULprloleD)
pula MoIsypIM( YRS) API ST4-OT APPZ ung-egc ] OI L7KEW OT AeFW unf-9gT]e a
eee n4_e4_4_ nn
IqI<e¢L 8penunuo)
9
7
0
qtA[ tureyqns sa1eW <D wsuuro yesOeNem <4 ceoen ewsS iPkpeyo Ne s<u4o uopy sojeuieg
sa1oadg 1314 poled N 143114 poued N Ws poued N 4 ysl poted N
TUT[VISITY
S1]DISIAT SMS4AASUDA] UUPLUDPIIAA AV [ur¬4- ¢E] ¬
P
snpiydojays npiospf{ JoyeA Arg v6 ARI O78 G AR ¬1-8 16 R
eSIALAOAT9(PpDTl1 7n4i WIPP e0 e)m vI]- g s 6-szs otd O
C
E
IUNUOPOIDy E
D
v1ojisd siuuadipiavjyfa enboewyy AVIO Q, I API 6c4-S \I76 APIL I I API 67-8 ¬ IN
G
TUTISSTTIA S
ZTU(sp7j soN4iau-v gFepIn boery])
O
DAAIT PIPPG (A94TRM) Ary ungL 4gy ¬ AP UNL4OZ¬ Z UNceE I any IepL- op 9 F
naalg sadijaid Qosig) Avy ung4¬lZ ] 4 T
psalg SUPsquin (UOISTTTIM) APIZ I APNS t-7T OZ API 97-81 APNESt T-7 z HE
sndjpdScNyAoDvOa g (1OATVAA) ARI 3-9 ¬ Aryl ungg 4s 6 Ari QI-S S Avy ungZ4 9g 8 E
snyd4sdsorSjPpaysDy juD (1OyJVMA) uneg I APIL I I N
T
Snyd4sdsoasjNpzYDDI sLAD (uOsuYyOoL) ungZL I O
M
snydidsozjpSysnDa gAdAj)p ys (uURWapaIA RFL I I Ine 87-L G ARI6 7 I O
sSnnyyddstOddGssYoo oojjppsyYmDDyp Fo(uY1t)9i p9uTlP M(1)g }UN AArvyP unugng4Zp4T9l ¢ E] ¬6 AABRWI 6C¬-IZnIg -9 IGZi AALeIW uuNngg4-¬L{SgT S~T AAVPIIVA mgU-NIsnZg 4O-E[ E ¬9 7 LOG
I
snydsdsoapwynys ow(4a)u [ne snZyT- 9zZ 6 C
A
snydid4saos8oijdp( y4y) APL BnY¢y 4-O| 8 AvP Ung4Oz¢Z¬] ¬ API snYZyI4] gI LZ ung 3sny94]- 9| ¬ L
snydsdsvoisaajjpdy y (uvlIInD) UNE 97-rI ¬
snydsdsso4ooIjJpaya D Ud}1SO) (UdYIVS API!L I I AvL e I SO
ET WOApvYi d Snypdan (AOTVAA) API 6C-OI ob ARPI9N 7 I CI
DIOlaT vasan (M20J) Ae 07-9 Ae ungg -¢ OIZ APN t7-S API6c -8 07 E
T
PIS] sisuaMs.Ai(AA NIC) une sny4bl- 7z g Ing 3nyI-l L v INE 6c I Y
SAISV]JDAII SNILIDAOMYO]O T ime OL I ung [ng-L¬IT ¢ = unF 7Tsny-O¢ G O
F
vinwos v4so2ap yenbory, Av ung-T7IZ z S API 67-01 Ve Ae ung4LI6 z 7
W
DIAWOJaY2y1d1S1A 1) (UURLTSpsI AWW SI-ZI Z Avy ung-TZI I S AP unLp 46z iG A
DIAWOIdSS nUMIjD (19ATVAA) ung 97 I S
H
vIAUOTvIIdSSn f MOOT [of¬ I Ime Te-¬1 S [ne 3nyI-T s 61 I
N
DULOISOULUSAU DUsIIIMD3 07T ung jngZL- Z II une Ing-IzZZ 6 G
J DULO]SOUSHW4Ad ]DG (19x[VAMA) API unf4¬76 z CG APW Ling-~z APOi ngcS- sz O6T ARI mg-I8zi 6 T
O
N
V
O
tuizidig 8ponunuoy <7 9IqeL XD(M907 DIVINIDULApONh YyjojAXy (ueIIND) 1auly DIOJAX uouurys vsnfuod pIojJAX eUBIIND Magupg piulojsouuea J, yIDMzeA_sJJI9yT,UjD e0 . LSo 7 A i)ApT pDDIOJAX T jIs opvOd8ijJI)s)AAs uyXAyreueyqng uie pDslaquy uaUSs10/001g zlieLapyouO p ajJieUseoSZoso1sadg ueO0uAn (Y UMOOT wleSnyaWagIio AS JsneNU aueaIuAie 8(WDtA sdiuAsIdaNMNoosIYyIfO( 1pSyiYIJONo d s8)TB(swdY UMa dfaDaOs{u iOsaT]0) aU OnS }IOQGOYLo6c-61 UNE L ne Oe] Ung ge AAAAPI vveeplap1sy yt [4e u3nuUNf4-4OT spgnong4lQy4SlL|gcoe]97 -I4 T Id o9e TP W <5 Na oe s6g woUUuulnNAnnAAAAsfGreuevv vv rOWy<Wf s T esONnyOc f ¬u]-3np uUEnpsfnenco]y8g4wnT8wgfc4La4¢g a7 y4 TsI-y- d oI4z Og ¢c] ] NI 9Z¬S¬vOI9I I 9c [NE I¢ 1ae LZ KEW [ unAv1e-¬1 AeW uuuAP nnnALP AeeeP PW sCr[Ii n-Ig6c-O1]--TsUNL4LzZ UIz p cZNo flSe4OE OOdOE R] ypcuN4 oO o s6Cvu O s eyepP NUUuONnnAAAA Eee4vPP e< eIP1 3p4II [-N8nILfUfn8g p4n-Rfos--9Ougp4lInI6l7T64]eEeZazLy sO-7d yI] o 4g] g] o]] N OGO¢ccvlbl SeLUME 108, NUMBER 4 sSse
8dds pisuiay pvpzzziiidddiiiggg VyjaSnydVp SsOjoiwduiMaUaOOfsI OiSTuT [U[OIIASAT T[IAA gA PI ABIN 7-8 it IngA vy OcO-I9- 9 G SAg4g y IR-IO 1A v [e -g el APIN 9 OZ6 API unfgO-lo T I6 9
LULISUTY y I DvpIIiSssOooPjpIliAaappYyySiiDDDIdd SSwwa8ADoodd]IsssyD d jdIopPoMyoIyu9oddMoosO DMv9iiTIssO0A i Oy7d T y Iay[eVeM IneAA RBIICC SI] 8nyL-I0 Z CO 6, une UnAAfe 4FWepI o O cu ngL -g ifSh ineAA PBIW 6s1Sn-Iy8-O- OI|1 ¬ 8 ne unAPIN APN API eTI 07 67-9 Ingung-,] ¬- 6T¬c I 81 ¬ ~ (6
p8diss oplayD une pl Ing 8nyI-l ¢ v1 [ne sny-TI¢ ¬ OT
DIBULvYaYp DuIvIuSIDAUp BsPSaI yD IONG (M907) AAeeC T BnyQ4T¢ o il oy Ae snyL-I¢ SEI AAveP UsNnngLy64 4s s IOZST AARRP4TW snsLyn ¢-y -¢6 1 i9l9
IUTTAWLOOLIAS ) 8¢ INL
vidwuooias sapioxojosdayyeon boryy Avy ung4b9c 6 Avy un[g 4¢] APINI¬ng 4-8gc Sr
9
SSS e7] e
N 9 I] 3ny-g APO OI mf-st APN 6 TI sny-T ung 7 pT INf4-61 API M20] Sadiasnf vapiq
O
T +SuMTQuAQLKASA(ssOIhrS n7nEnnuEvPv]- l 7 -yIgga gyLy s8dN44s8n i sgIud aasJad) A sog
G
N 9 67-8 API I g AR (UOFSTT[IAA) smyixnod snydidstspg
I I LI] ung I ¬ une Z TZ unf4OT AvP 8ds winxojostay)
H
S ZI ZI Sny-p7hPI4N 9. TI SnYV-LIARPIN OO LI Bny4-s AeFCW SOO. 9| sny-9 Aepy JOY[eM wNIPAlsap wunxojoskayD
A
W v 6c uUnf4-Oc APN auc gung4_] Ae =Z TI (mg4-T une (Avg) pnbyqo vidvssojp y
F rurydaXg
O
IQAI¢38()sd T snsr s ns neyvs ipvagp d
Y
T ¬1 IL 3sny-O1[ARPW Oo LI ung4g Avy TE LIT sny-[[ ARN OI Ol 8ny4ST ARI UdBIII, SNOYALOWIADY SNSDAD T
ENTOMOLOGICAL SOCIE 9¢ GE I II ¢ sny49 sny4s 6c Ae Arp API tT aye ZS Ol I I QO] LI Pc ung4g] 8ny4s [ng4g 974-S LT ¢ KEW Av Ae API AvP une ¬ L II sny-s] LI-9 AeYL= une 8 i ¬ SI ZT pc sny-OQ] ung4pz [ng4g Ae Av API (JPOISI9}19Z) (1989R1S) YIOIBYIO (Aes) (J) (Aes) SNIDULBADUIWL A, snasogsaddy wnuyjaiu SNINJIADAU snipaponb snanosgo pulojsounjapy SNALAYIAID] snsélayIAID] snslayIAIY] SNALBYIAID] snaayrAIv] TuryI gq J g J Il 97 Inf-O1 APIN IZ L 3ny-o01 KEW OL 9 Inf-el ARP OO ¬ [ng49 Ar (A) Pipsuoja DYyII0g stS OT sny4s Ar Il I¬ [ng-g AvP 8dds snuayrtv]doeITIMSSUIs e1eg g nIUNI e OBO NIHdYAS ADT{I IDITS NBUD
E V
TH A
9 L Ing4-vz PW II gc ung-7z APIN ¢ unf467 Ave MOOT SUSIAT UOPOLP Y
F
O G O¢¬-6I UNL MOUS 49JSVBO]DBIUW UOPOAILY
S +6 TI sny-s Ing Lor LI 8ny-6z une UOYLM MpPVaYsSIVAI UOPOANPY
NG Z pe Ine M207 snsaidolpg Uoposrly
I IU UOPOIIPAY
D
E
E AVNILNOGO SIN
C
O i. 9 [ng4-[TE API I pl ung (J) supjpdquiog vyaonjo,
R
P LI TI sny-ZI une ZI ZT SNYW-ST ARI 6cl pl sny4gI Aew Or pl sny-sgI Ae (UBIIND) wintDnsisaa wnjAjsadoy
Tureon[OA .
N poled 1st4 N poled 1WsI4 N poled 1Wyst4 N powed 1yst4 sarsadg
soyRulaf SoTRIW so[PUlo 4 sole equ
<dN bpeyRsuou0p] <JN uo wurysem <OD Ajrureyqns
2 8ponunuoy <¢ eIqeL
7
9
V
LVQIA 7< younunop8 O
L
U
M
Ayrueyqns <DH sUIUYoy sOENM <4 Ppeyes<ucduNc py E
equ sar soyPRulaf IA So[P soyPulaf
sosog SypnbapM OOT AeW= Ing4SsIt t API7 7 I 108,
N
aydoasidy vinuisin(iAuetsa) Ae OI-9 g Ary] sn¢y¬ 49 9¢ APIA OT-Ol 9 Avy sny4<9T] SE U
M
aydosjsidy av1avjnssos(sU dBIOY) snlyL I [ug pc74-S ¬ ine sny-Q4I LI Ing 3nVp-l6 I ¢ B
eayyddoo.assiiddyyY jpsiojjoarui p(iaiyun p(4Ua)S IBIA]) API OI-s OO AARvP4y|W uunngcgg-4 p¢gl 6 APOL7E-9 LZ AAPvN Ingu-Ing|cg 46z GLa l ER
aYydoslsidy VUOJsoyjUvxX( UOFISTT[IAA) Ae OI-8 (6 Avy ung[ 4o], t ABW 6C-II S Avy ung4QOg]] LI 4
Sapoadnyz SNUDAIAI)M ALUD (UUPLUOPaI Ary Ung46g 07 Avy SnyQ4]s QP Avy unLg 49 PET Ary 8snyL-I s ¬¬l
sapoadny snxajdiad (uinqsQ) AVIO T I API PT-LI G
Sapoadny IpuDjaUlA (UBIIND) Ae SI-38 7 Avi snpyb 4g 87 Arp Unf4QOg, Il Ary snyO4L9 cP
sapo8adsd ny ARIO L Z Av OI-g SCNG a v
DUOZOINAT SsaplojojAx (uOsUYyOL) UNL 97-6| G
DUASUDLAWP4Wa Yyslf (u0][BM) Av[ ¢ I
DudAsuYjayy vugoylydoisn] Qpeisi9197) ABT¬ [ I Kegl I AE L7-9 9S
DUASUDIAPY vaafinsupié] ({poiss19je7Z) AvP Ung-TOI l ¬T Avy ung4LIg y ¬l
DAavISyaPY jJAIIULI (QQpoisi99Z) APgI I Avy [ng4OgL S ALI UNg46sc Lv Avy snyp4l9 +9
snupidt9¢sQn ipiso(sA es) snpy I
snumjidkIE S7uuAad)i aspf (UUPLUSpaI Ine Oc I
snumidta¢Q siuuadi(oAsensf) API 8nyLI4]s Z g snyQ O] I une snyb-lS z 9
snydsdspsSvYgY nuas (UOFSTI[IAA) AVIN 77-8 9
snydidsvivgM apunjaiu (URIIND) APIS I I
visoydosapvyndssiyj uos yienboryy APP ung4Osg ¬ Av uNngT4Tg Ol Aveypy ung[ 4c{T - AvP unf4¬OTT 8
piuo8ydsd osapyds snY¢y I AeS T I
snydidgi qpuy uouurys Avy [ng4O9L SI Avy 3snyLI- 9 68 Ar [ng4QsZ ra AFI 8nyL4I- O| GS
snydidgs Njoa4 UdISO UdYIVG Avy [In¢g 49 Lvl APY UNg46sc 691 Ary snyI4] g ¬¬Pr Avy sny94]g MLE
snydidgu saqia( J) Ae unf4b-lo T ae OVINE CGS SZ API 61-8 LT APIA Unf46gc os
snydidgys nAso] UdISO udyIeVS APL LI-s Lv AeWw LI-S OT Ary unpg4 ¢g S87 Ary UN4TsT SIZ
punupssoySJadUidaXn pf (M907) API UNs466c] 7 API 8nWL-I Zz OE API 8snLVIy -Zz 97
|, TULIDULOXO
J sn4aUOXoO SniIpU(AAsBSS ) Av 8nyL4Is 66l Avy snyL4Is 187 Avy 8nyL4Is LEI Ae snyL4Is bor
J SNAIUOXO SNIDUIB(AADBUSL) API snyp4l- Ol tl API sn¢y 48] 6% Ae IngQ4 s IZ AvY sny94]9 gE
J snsauiosxnoj jod (Aes) jal© I me T¢¬ I
e9 e
7
3
