USDA 

United States 

Department Insects and Related Arthropods Associated with 

of Agriculture 

Greenleaf Manzanita in Montane Chaparral Forest Service 

Pacific Southwest Communities of Northeastern California 

Research Station 

General Technical Report Michael A. Valenti George T. Ferrell Alan A. Berryman 

PSW-GTR- 167 
Publisher: 

Albany, California 

Mailing address: 

PO Box 245, Berkeley CA 

94701 -0245 

September 1997 

Pacific Southwest Research Station 

Forest Service 

U.S. Department of Agriculture 

Abstract 

Valenti, Michael A.; Ferrell, George T.; Berryman, Alan A. 1997. Insects and related 

arthropods associated with greenleaf manzanita in montane chaparral 

communities of northeastern California. Gen. Tech. Rep. PSW-GTR-167. Albany, 

CA: Pacific Southwest Research Station, Forest Service, U.S. Dept. Agriculture; 26 p. 

Specimens representing 19 orders and 169 arthropod families (mostly insects) were 

collected from greenleaf manzanita brushfields in northeastern California and identified 

to species whenever possible. More than500 taxa below the family level wereinventoried, 

and each listing includes relative frequency of encounter, life stages collected, and 

dominant role in the greenleaf manzanita community. Specific host relationships are 

included for some predators and parasitoids. Herbivores, predators, and parasitoids 

comprised the majority (80 percent) of identified insects and related taxa. 

Retrieval Terms: Arctostaphylos patula, arthropods, California, insects, manzanita 

The Authors 

Michael A. Valenti is Forest Health Specialist, Delaware Department of Agriculture, 

2320 S. DuPont Hwy, Dover, DE 19901-5515. George T. Ferrell is a retired Research 

Entomologist, Pacific Southwest Research Station, 2400 Washington Ave., Redding, 

CA 96001. Alan A. Berryman is Professor of Entomology, Washington State University, 

Pullman, WA 99164-6382. 

All photographs were taken by Michael A. Valenti, except for Figure 2, which was taken 

by Amy H. Valenti. 
Insects and Related Arthropods Associated with 

Greenleaf Manzanita in Montane Chaparral 

Communities of Northeastern California 

Michael A .Valenti George T.Ferrell Alan A .Berryman 

Contents: 

Introduction .......................................................... 1 Pacific Southwest 


Research Station 

StudyArea ........................................................... 2 


LoganLake ....................................................... 2 USDA Forest Service 


Tamarack Swale ................................................... 2 General Technical 


Bearwallow ...................................................... 2 Report 


Hat Creek ........................................................2 PSW-GTR- 167 


Materials and Methods ................................................ 2 September 1997 


Results and Discussion ................................................ 3 


Fungivores ........................................................ 3 


Pollinators ........................................................4 


Herbivores ........................................................ 4 


Xylophagous forms (wood-feeders) ................................4 


Rhizophagous forms (root-feeders) ................................ 4 


Foliovores (foliage-feeders) ........................................ 15 


Leaf rollers/ tiers ................................................ 15 


Leafminers .................................................... 15 


Gall formers .................................................... 15 


Piercing/sucking insects .........................................15 


Defoliators ..................................................... 17 


Predators ........................................................ 20 


Parasitoids ....................................................... 21 


Scavengers ....................................................... 22 


Conclusions ......................................................... 23 


Acknowledgments ...................................................24 


References ..........................................................25 

Figure I-Greenleaf manzanita in Shasta County, California, several days after budbreak. 

Introduction 

Greenleaf manzanita (Arctostaphylos patula E. Greene) is a broadleaf 

evergreen shrub that commonly grows in the Sierra Nevada of 

California (Hiclunan 1993) (fig.1). This species grows in mesic habitats 

associated with montane forests generally above 1,000 m in elevation 

(Ball and others 1983). After a natural or human-caused disturbance, 

such as a fire or clearcutting, greenleaf manzanita shrubs often become 

established and quickly dominate a site preventing or prolonging the 

successional sequence or inhibiting growth of planted conifer seedlings 

(McDonald and Fiddler 1990, Oliver 1990). This type of domination, in 

which shrubs prevent the establishment of important conifers (e.g., 

ponderosa pine [Pinus ponderosa Dougl. ex Laws. var. ponderosa]), inhibit 

conifer growth, and reduce conifer survival (Conard and Radosevich 

1982, Lanini and Radosevich 1986, Radosevich 1984), has been termed 

"inhibition succession" (Connell and Slatyer 1977). 

Current shrub control methods (e.g., mechanical removal, 

prescribed burning, and herbicide treatments [Bentley and Graham 

1976, Paley and Radosevich 19841) can have drawbacks such as high 

costs, poor efficacy (Lanini and Radosevich 1982), damage to conifer 

seedlings (Newton 1963), and environmental risks such as adverse 

effects on non-target organisms. Past restrictions on controlling 

undesirable vegetation have significantly increased the total area of 

competing vegetation on Federal lands (Walstad and others 1987). 

The controversy over current direct control techniques and the 

increase in demand for more environmentally sound control 

methods has increased the need for alternatives such as using native 

herbivorous insects to control greenleaf manzanita in northeastern 

California (Valenti 1994). 

In this study, we compiled a list of insects known to be associated 

with greenleaf manzanita. An extensive literature search yielded only a 

few general books (e.g., Fumiss and Carolin 1977, Johnson and Lyon 

1988, Keen 1952) and several research publications (e.g., Duckworth 

1964, Fumiss and Barr 1975, Heinrich 1923) that mention insects, mostly 

lepidopteran larvae known to feed on manzanita species. A more 

intensive study by Focarile (1975) reported 111 insect taxa from the 

cosmopolitan manzanita species A. uva-ursi (L.) Spr. in Italy's Aosta 

Valley. In addition, the North American Hymenoptera catalog 

(Krombein and others 1979) listed 27 pollinating hymenopteran species 

that visit greenleaf manzanita flowers. However, these species accounts 

were inadequate for our investigation because information concerning 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
herbivorous insects feeding specifically on greenleaf manzanita or 

surveys of insects occurring in brushfield ecosystems (e.g., montane 

chaparral communities) are scant in the literature (Force 1990, Haws and 

others 1988). Therefore, we conducted an intensive survey to identify the 

insects associated with greenleaf manzanita and to determine their 

respective roles in the manzanita community. 

This paper summarizes the results of a detailed entomofaunal survey 

of arthropods that use greenleaf manzanita for food and shelter. The 

inventory can be used in an effort to accelerate succession of greenleaf 

manzanita-dominated sites to conifers on sites historically dominated by 

the latter. This paper also includes information on the characterisitc 

damage caused by major groups of insect herbivores. As the problems of 

manipulating native competing species become more pressing, these 

lists may contain critical information about insect communities (e.g., 

faunal associations) that can be a valuable asset to biologists and 

ecologists in the future. 

Study Area 

The four study areas in northern California were Logan Lake, 

Tamarack Swale, Bear Wallow, and Hat Creek; plant 

identifications were based on descriptions given by Munz (1973). 

Logan Lake 

The Logan Lake site (elevation 1,512 m [4,960 ft]) was located 3.2 km 

northwest of California Route 89 near Old Station and 1.6 km west of 

Logan Lake, about 24 km south of Hat Creek (T32N R4E S2&3). The 

50+ ha site was on an east-facing slope and was mechanically cleared 

of all standing vegetation in 1976 and planted to ponderosa pine the 

following year. Dominant vegetation (measured as percent canopy 

coverage) consisted of greenleaf manzanita (74 percent), tobacco 

brush (Ceanothus velutinus Dougl. ex Hook.) (11 percent), and 

planted ponderosa pine (9 percent). Greenleaf manzanita shrubs 

averaged 1.5 to 2.0 m in height. 

Tamarack Swale 

The site was located in a valley 6.4 km southwest of California Route 

89 about 5 krn south of Hat Creek (T33N R4E S4&9) (elevation 1,646 

m [5,400 ftl). In 1974 this 25+ ha site was mechanically cleared and 

ponderosa pine seedlings were planted in 1975. Greenleaf manzanita 

shrubs were removed again in 1984 during a second mechanical 

clearing treatment; however, shrubs have subsequently become 

reestablished and average about 0.75 m high. Tobacco brush (16 

percent) and planted ponderosa pine (8 percent) were mixed in with 

the most abundant plant species, greenleaf manzanita (61 percent). 

Bear WaHow 

This 20+ ha old-growth greenleaf manzanita brushfield (elevation 1,524 

m [5,000 ft]) was located 3 km (by dirt road) from California Route 89 in 

Old Station (about 6.6 km south- southwest of the Logan Lake site) 

(T32N R4E S10). Greenleaf manzanita comprised about 85 percent of the 

total canopy closure and averaged about 1.5 to 2.0 m high. 

Hat Creek 

Located adjacent to the USDA Forest Service work center at Hat 

Creek (T34N R4E S16) (elevation 1,018 m [3,340 ft]), this site consisted 

of a variety of woody plant species. Greenleaf manzanita accounted 

for less than 30 percent of the total canopy closure. Other apparent 

plant species included sagebrush (Artemisia tridentata Nutt.), curlleaf mountain-mahogany (Cercocarpus ledifolius Nutt.), birch-leaf 

mountain-mahogany (C. betuloides Torr.), antelope bitterbrush 

(Purshia tridentata [Pursh] de Cand.), California black oak (Quercus 

kelloggii Newb.), and a mixture of conifers (30+ years old) including 

ponderosa pine, sugar pine (Pinus lambertiana Doug.), California 

white fir (Abies concolor [Gord.] var. lowiana Lemm.), Douglas-fir 

(Pseudotsuga menziesii [Mirb.] Franco), incense-cedar Calocedrus 

decurrens Torr.), and western juniper (Juniperus occidentalis Hook.). 

This site was also characterized by extensive areas of exposed lava. 

Materials and Methods 

Arthropods (mostly insects) were sampled from all four study sites 

by using a variety of collecting techniques. In descending order of 

relative importance, or frequency of use, these techniques included 

beating branches and foliage over a muslin cloth or bag (fig. 2), handpicking individuals from plants, sweeping foliage with a canvas net, 

using an aerial net for flying insects, pitfall-trapping, attracting adults 

to black light, using a Berlese funnel, and Malaise-trapping (fig. 3). 

Collecting occurred at a minimum of 1-week intervals (and often 

more frequently) and generally began in May and ended in August 

from 1989 to 1994. Specimens were collected in the field and 

transported to the USDA Forest Service's Forest Insect Laboratory, 

Hat Creek, California, for preparation. All adult specimens were 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Figure 2-Insects were collected at the Logan Lake site in 1989 by beating greenleaf rnanzanita 

branches over a muslin bag. 

Figure 3-Malaise trap used to collect flying insects at the Logan Lake site. Insects fly into and land 

on the sides of the trap and crawl upwards becoming trapped in a container at the top. 

either pinned or preserved in alcohol and labelled with complete 

collecting information. Specimens were sent to various taxonomic 

experts for positive identification. The ecological role of each taxon 

(i.e., how it functions in the manzanita community) was determined 

by direct observations and rearings, or inferred from related species 

by consulting general entomological references (e.g., Borror and 

others 1989, Essig 1958, Powell and Hogue 1979). Unless otherwise 

indicated, voucher specimens are now stored in the Maurice T. 

James Entomological Museum, Department of Entomology, 

Washington State University, Pullman. 

We attempted to rear herbivorous immature insects (mainly 

Lepidoptera) collected from greenleaf manzanita plants to 

adulthood because larval forms usually cannot be identified to 

species. Individual larvae were placed in small plastic containers or 

screen cages with host material and allowed to develop and pupate. 

Many pupae required a cold treatment (e.g., 90 days at 4 OC) for 

adults to emerge. From these larval rearings, parasitic flies (Diptera) 

and wasps (Hymenoptera) were also collected. 

Arthropods collected in greenleaf manzanita brushfields were 

grouped by their dominant role in the community. Sampling was 

biased towards these groups because the primary goal of this study 

was to identify important greenleaf manzanita herbivores and their 

associated predators and parasitoids. Other groups of arthropods, 

such as fungivores and pollinators, were generally not actively sought, 

and their presence should be considered mostly a random encounter 

during the collection of herbivores and their natural enemies. 

Results and Discussion 

Specimens representing 19 orders and 169 arthropod families 

(mostly insects) were collected from greenleaf manzanita 

brushfields and identified to species whenever possible (table 1). 

More than 500 taxa below the family level were inventoried and 

each listing includes relative frequency of encounter, life stages 

collected, and dominant role in the greenleaf manzanita community. 

Specific host relationships are included for some predators and 

parasitoids. Herbivores, predators, and parasitoids comprised the 

majority (80 percent) of identified insects and related taxa (fig4). 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Fungivores 

Eleven arthropod taxa the it feed on fungi were collected in this survey 

including a mite (Acari: Eupodidae), a true bug (Hemiptera: 

Aradidae), a beetle (Coleoptera: Erotylidae), four fungus gnats 

(Diptera: Mycetophilidae), three dark-winged fungus gnats (Diptera: 

Sciaridae), and an ant (Hymenoptera: Formicidae). 

Pollinators 

To Krombein's (1979) list we added 10 more pollinator taxa (all in 

the order Hymenoptera) associated with greenleaf manzanita 

flowers: three solitary bees (Halictidae [21 and Andrenidae [I]), two 

long-tongued bees (Anthophoridae), and one honey bee and four 

bumble bees (Apidae). One additional solitary bee (Andrena 

cerasifolii Cockerel1 [Andrenidael) was also encountered and is 

included in the list by Krombein and others (1979). 

Herbivores 

A total of 184 herbivorous insect species representing 7 orders and 55 

families were collected. This group includes insects that feed on any 

part of a living greenleaf manzanita plant (i.e., wood, roots, and leaves). 

Xylophagous forms (wood-feeders) 

Adult beetles (Coleoptera) of species suspected of using greenleaf 

manzanita as a host (in the larval form) were collected from foliage. 

These species include six metallic wood-boring beetles 

(Buprestidae), five long-horned beetles (Cerambycidae), and three 

powderpost beetles (Anobiidae). Small (3mm) D-shaped holes were 

frequently observed in dead and dying stems of greenleaf 

manzanita. Such adult exit holes are typical of beetles belonging to 

the genus Agrilus (Buprestidae). 

Rhizophagous forms (root-feeders) 

Cicada nymphs (Homoptera: Cicadidae) and larvae of certain 

scarab beetles (Coleoptera: Scarabaeidae), elater beetles 

(Coleoptera: Elateridae), weevils (Coleoptera: Curculionidae), and 

moths (Lepidoptera: Hepialidae) are all known to feed on plant 

roots. Representatives of all groups were collected, as adults, in 

association with greenleaf manzanita and we suspect the immature 

forms feed on the roots. However, because we focused on 

foliovores and detailed studies were not made on root-feeders, we 

cannot be absolutely sure of direct association with greenleaf 

manzanita plants. 

Role in manzanita community 

Figure 4-Number of arthropod taxa (below family level) grouped by their dominant role in a 

greenleaf manzanita brushfield community. All specimens were collected in Shasta County, 

California from 1989 to 1994. Fu, fungivore; Po, pollinator; He, herbivore (including wood 

boring forms); Pr, predator; Pa, parasitoid (including hematophagous forms); Sc, scavenger 

(including detritivores and coprophagous forms); X, no apparent direct association with manzanita. 

*Does not include 50+ species of unidentified spiders. 

USDA Forest Service Gen.Tech. Rep. PSW-GTR-167. 1997. 
Table 1-Arthropods associated with greenleaf manzanita brushfields in Shasta 

County, California 

Relative Stages Role in 

Faxon frequency1 encountered2 community3 

ARANEAE 

Thomisidae 

Misumenops celer (Hentz) R Pr 

host: Synaxis cervinaria 

(Packard) (Lepidoptera: Geometridae) 

Xysticus sp. R Pr 

host: Synaxis cervinaria 

(Packard) (Lepidoptera: Geometridae) 

Salticidae 

Metaphidippus sp. R Pr 

host: Synaxis cervinaria 

(Packard) (Lepidoptera: Geometridae) 

families and species (50+) undet. C Pr 

ACARI 

Ascidae 

Proctolaelaps sp. 0 Pr* 

Ixodidae 

species undet. R Hm 

Bdellidae 

Spinibdella sp. 0 Prt 

Eupodidae 

Eupodes sp. R Fut 

found in leaf galls formed by Tamalia coweni 

(Cockerell) (Homoptera: Aphididae) 

Erythraeidae 

Balaustium sp. 0 Pr* 

Erythraeus sp. R Pr* 

associated with Tamalia coweni 

(Cockerell) (Homoptera: Aphididae) 

Phytoseiidae 

Typhalodrominsp. R Pr* 

associated with Tamalia coweni 

(Cockerell) (Homoptera: Aphididae) 

Johnstonianidae 

Lassenia sp., prob. n. sp. R Prt 

associated with Tamalia coweni 

(Cockerell) (Homoptera: Aphididae) 

PSEUDOSCOwIoNES 

family and species undet. Pr 

DIPLOPODA 

family and species undet. sc 

IHILOPODA 

family and species undet. Pr 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 

Table 1, continued 

Relative Stages Role in 

Faxon frequency1 encountered2 community3 

ZOLLEMBOL A 

Isotomidae 

species undet. De 

Entomobryidae 

species undet. De 

Sminthuridae 

species undet. De 

MICROCORWHIA 

Machilidae 

species undet. De 

ODONATA 

Aeshnidae 

Aeschna palmata Hagen Pr 

Anax junius (Drury) Pr 

Cordulegasteridae 

Cordulegaster dorsalis Hagen Pr 

Libellulidae 

Libellula sp. Pr 

Pantala hymenea (Say) Pr 

Tarnetrum corrupturn (Hagen) Pr 

Lestidae 

species undet. Pr 

Coenagrionidae 

species undet. Pr 

ORTHOPTERA 

Acrididae 

Camnula pellucida (Scudder) He 

Melanoplus aridus (Scudder) He 

M. sanguinipes (F.) He 

Tettigoniidae 

Capnobotes attenuatus Rentz & Birchum He 

Scudderia furcata Brunner von Wattenwyl He 

Gryllacrididae 

Ceuthophilus sp. sc 

Stenopelmatus sp. sc 

Gryllidae 

Oecanthus sp. He 

BLATTARIA 

Blattellidae 

Parcoblatta americana (Scudder) sc 

continued 

5 
Table 1, continued Table 1, continued 

Relative Stages Role in Relative Stages Role in 

axon frequency1 encountered2 community3 axon frequency1 encountered2 

-

community3 

-

'SOCOPTERA 3EMIPTERA (HETEROPTERA), continued 

Psocidae Aradidae 

species undet. Sc Mezira sp. 

Tingidae 

WYSANOPTERA Garaphia iridescens Champion 

Aeolothripidae 

Aeolothrips sp., nr. oregonus Hood 

A. sp. 

Thripidae 

Chirothrips aculeatus Bagnall 

Frankliniella occidentalis (Pergande) 

Thrips tabaci Lindeman 

Phlaeothripidae 

Leptothrips mcconelli (Crawford) 

R 

R 

R 

C 

R 

R 

Pr* 

Pr 

He 

He 

He 

Pr 

Phymatidae 

Phymata sp. 

Reduviidae 

Apiomerus sp. 

Pselliopus spinicollis Champion 

Sines sp. 

Zelus tetracanthus StSl 

Nabidae 

Nabis alternutus Parshley 

Anthocoridae 

-1EMIPTERA (HETEROPTERA) 

Pentatomidae 

Anthocoris whitei Reuter 

host: Tamalia coweni (Cockerell) 

Banasa dimiata (Say) 0 

B. sp. R 

Chlorochroa ligata (Say) C 

parasitoid reared from eggs: 

Telenomus sp. (Hymenoptera: Scelionidae) 

Euschistus sp. R 

Holcostethus abbreviatus Uhler R 

He 

He* 

He 

He 

He 

(Hornoptera: Aphididae) 

Onus tristicolor (White) 

Miridae 

Chlamydatus manzanitae Knight 

Deraeocoris brevis (Uhler) 

D. fulgidus (Van Duzee) 

D. vanduzeei Knight 

Thyanta pallidovirens (Stil) 

Scutelleridae 

C He Ectopiocerus anthracinus Uhler 

Irbisia sp. 

Homaemus bijugis Uhler 

Vanduzeeina californicus Van Duzee 

R 

R 

He 

He 

Lygus Hesperus Van Duzee 

Macrotyloides sp. 

Cydnidae 

Macroporus repetitus Uhler 

Coreidae 

R He 

Parthenicus sp. 

Phymatopsallus sp. 

Phytocoris hettenshawi Bliven 

Leptoglossus clypealis Heidemann I x Pilophorus sp. 

Alydidae 

Alydus eurinus (Say) 0 

Tollius quadratus Van Duzee 0 

Rhopalidae 

Arhyssus sp. R 

Lygaeidae 

Ashlockaria sobria (Uhler) C 

Eremocoris sp., prob. semicinctus Van Duzee R 

Geocoris pallens StSl 0 

Lygaeus kalmii St51 R 

Malezonotus grossul Van Duzee C 

Thylochromus nitidulus Barber R 

Berytidae 

Jalysus wickhami Van Duzee R 

Neides muticus (Say) R 

He 

He 

He 

He 

He 

Pr 

Pr 

He 

He 

He 

He 

HEMIPTERA (HOMOPTERA) 

Cicadidae 

Okanagana ferrugornaculata Davis 

Platypedia putnami (Uhler) 

Membracidae 

Telamona woodruffi Ball 

Tortistilus wickhami (Van Duzee) 

Cercopidae 

species undet. 

Clastopteridae 

Clastoptera sp. 

Cicadellidae 

Aceratagallia californica (Baker) 

A. longula (Van Duzee) 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Table 1, continued Table 1. continued 

Relative Stages Role in Relative Stages Role in 

frequency1 encountered2 community3 Taxon frequency1 encountered2 community3 

HEMIPTERA (HOMOPTERA), HEMIPTERA (HOMOPTERA), 

Cicadellidae, continued Aphididae, continued 

A. lyrata (Baker) 0 Tamalia coweni (Cockerell) 

Aligia sp. C Wahlgreniella nervata (Gillette) 

Balclutha incisa (Matsumura) R Diaspididae 

B. neglecta (DeLong & Davidson) 0 Lepidosaphes sp. 

Ballana aperta DeLong 0 Pseudococcidae 

B. dim DeLong C Puto arctostaphyli Ferris 

B. latula DeLong C 

B. titusi (Ball) 0 NEUROPTERA 

Colladonus geminatus (Van Duzee) C Raphidiidae 

Dikraneura variata Hardy 0 Raphidia assimilis Albarda 

Empoasca filaments DeLong 0 R. bicolor (Albarda) 

E. pallida Gillette C R. sp., prob. bifurca Banks 

Exitianus exitiosus (Uhler) 0 R. herbsti Esben-Peterson 

Gyponana sp. R R. sp., prob. neglecta Carpenter 

Liburnia sp. 0 R. unicolor (Carpenter) 

Momoria turgida (Beamer & Lawson) C Inocelliidae 

Neocoelidia lineata Baker C Inocellia inflata Hagen 

Osbornellus borealis DeLong & Mohr R Coniopterygidae 

0.curvatus (Beamer) R Coniopteyx sp. 

Paraphlepsius apertinus (Osborn & Lathrop) R Conwentzia sp. 

Delphacidae Hemerobiidae 

Delphacodes sp. Hemerobius sp., prob. ovalis Carpenter 

Cixiidae H. sp., prob. pacificus Banks 

Oeclidius sp. Micromus posticus Hagen 

Dictyopharidae M. variolosus Hagen 

Scolops abnormis Ball Sympherobius sp. 

Achilidae Chrysopidae 

Epiptera henshawi (Van Duzee) Ch ysopa sp. 

Synecdoche albicosta (Van Duzee) Chrysoperla carnea (Stephens) 

S. rubella (Van Duzee) C. oculata Say 

Nogodinidae Eremochysa fraterna Banks 

Dictyssa clathrata Melicher Berothidae 

D. marginepunctata Melicher Lomamyia sp., prob. latipennis 

Psyllidae carpenter or nr. 

Euglyptoneura robusta (Crawford) Myrmeleontidae 

Euphalerus vermiculosus Crawford Brachynemurus sp. 

Neophyllura arctostaphyli (Schwarz) Mantispidae 

N. arctostaphyli var. bifasciata (Crawford) Climaciella brunnea (Say) 

N. bicolor (Martin) 

N. separata (Tuthill) COLEOPTERA 

Aleyrodidae Cicindelidae 

Trialeurodes vaporariorum? (Westwood) Cicindela purpurea lauta Casey 

Aphididae Carabidae 

Cinara sp. Calosoma luxatum Say 

Dactynotus sp. 

continued 

USDA Forest Service GemTech. Rep. PSW-GTR- 167. 1997. 7 
Table 1, continued Table 1, continued 

Relative Stages Role in Relative Stages Role in 

Faxon frequency1 encountered2 community3 Faxon frequency1 encountered2 community3 

ZOLEOPTERA, ZOLEOPTERA, 

Carabidae, continued Elateridae, continued 

Lebia perita Casey Ctenicera pallidipes (Brown) 

Pterostichus inanis? Horn C. tigrina (Fall) 

P. lama Mknetries Dalopius sp. 

P. melanarius Illiger Hemicrepidius dilaticollis (Motschulsky) 

P. planctus? LeConte Limonius bicolor Van Dyke 

P. protractus? LeConte L. ornatulus LeConte 

Staphylinidae Megapenthes aterrimus (Motschulsky) 

Staphylininae Neopristilophus maurus (LeConte) 

Bolitobius sp. Selatosomus edwardsi (Horn) 

Eusphalerum sp. Phengodidae 

Medon sp. Zarhipis sp. 

Quedius sp. Cantharidae 

Sepedophilus sp. Cantharis sp. 

Stictolinus sp. Silis sp. 

Tachyporus sp. Anobiidae 

Histeridae Oligomerus sp., prob. undesc. sp. 

Xerosaprinus sp. Xylentinus fucatus LeConte 

Scydmaenidae X. sp., nr. rotundicollis White 

Lophioderus sp. Ptinidae 

Stenichnus (Brachycepis) fuchsi? Brendel Ptinus fallax Fall 

Lucanidae Bostrichidae 

Platyceroides latus (Fall) Scobicia declivis (LeConte) 

Scarabaeidae Cleridae 

Aphodius opacus LeConte Phyllobaenus funebris (Chevrolat) 

Canthon simplex LeConte P. scaber (LeConte) 

Cremastocheilus armatus Walker Melyridae 

Dichelonyx backi (Kirby) Amecocerus sp. 

D. valida (LeConte) Hapalorhinus sp. 

Serica anthracina LeConte Nitidulidae 

S. curvata LeConte Thalycra sp. 

Buprestidae Erotylidae 

Acmeodera mariposa mariposa Horn Dacne californica Horn 

A. plagiaticauda plagiaticauda Horn Coccinellidae 

A. vandykei Fall Adalia bipunctata (L.) 

Agrilus arbuti Fisher Anatis rathvoni LeConte 

Anambodera gemina (Horn) Cycloneda polita Casey 

Anthaxia deleta deleta LeConte Exochomus californicus Casey 

Chysobothris dentipes (Germar) Hippodamia apicalis LeConte 

Ch ysophana placida (LeConte) H. convergens Guerin 

Elateridae Hyperaspis sp. 

Athous imitans Fall Mulsantina picta (Randell) 

A. nigropilis (Motschulsky) Psyllobora borealis Casey 

Cardiophorus edwardsi Horn Scymnus (Pullus) mendocino Casey 

C. sp., nr. tenebrosus LeConte host: Tamalia coweni (Cockerell) 

C. tumidicollis LeConte (Homoptera: Aphididae) 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Table 1, continued Table 1, continued 

Relative Stages Role in Relative Stages Role in 

frequency1 encountered2 community3 

COLEOPTERA, continued ZOLEOPTERA, continued 

Lathridiidae Curculionidae 

Corticarina fuscul (Gyllenhal) Anthonomus sp., prob. signatus 

Tenebrionidae Say spp. complex 

Coelocnemis californica Mannerheim Gymnetron tetrum (F.) 

C. rugulosa Doyen Leposoma segnis (LeConte) 

Coniontis subpubescens? LeConte L. sp., nr. but not tanneri Van Dyke 

Eleodes hybridus? Blaisdell Lepidophorus sp. 

E. scabricula LeConte Nemocestes montanus Van Dyke 

Helops simulator Blaisdell Pachyrhinus cinereus (Casey) 

Scotobaenus sp., prob. parallelus LeConte Panscopus (Dolichontus) sp. 

Alleculidae P. (Nomidus) sp. 

Isomira variabilis Horn Peritelinus oregonus Van Dyke 

Mycetochara pubipennis (LeConte) Pselaphorhynchites aureus (LeConte) 

Pseudocistela or Telacis sv. Scolytidae 

Stenochidus yanescens LeConte Hylastes nigrinus (Mannerheim) 

Melandryidae 

Anaspis? sp. 3IPTERA 

Mordellidae Tipulidae 

Mordella albosuturalis Liljeblad species undet. DeW 

Meloidae Mycetophilidae 

Lytta sp. 

Anthicidae 

Anthicus punctulatus LeConte 

lschyropalpus nitidulus (LeConte) 

Cerambycidae 

Acomoptera sp. 

Allodia sp. 

Megophthalmidea occidentalis Johannsen 

Mycetophila sp. 

Sciaridae 

Fu(L) 

Fu(L) 

Fu(L) 

Fu(L) 

Centrodera spurca (LeConte) 

Leptura obliterata (Haldeman) 

Bradysia spp. (3) 

Cecidomyiidae 

Fu(L) 

Necydalis sp., prob. laevicollis LeConte 

Neoclytus baltaetus (LeConte) 

N. resplendens Linsley 

Ulochaetes leoninus LeConte 

Xylosteus ornatus (LeConte) 

Asphondylia sp. 

Lestodiplosis sp. 

Porricondyla sp. 

Winnertzia sp. 

Simuliidae 

He(L) 

He(L) 

He(L) 

He(L) 

Xylotrechus longitarsis Casey Simulium (Psilozia) vittatum Zetterstedt Hm 

Bruchidae S. (Simulium) decorum Walker Hm 

Acanthoscelides aureolus (Horn) Chironomidae 

Chrysomelidae species undet. Hm 

Altica bimarginata Say Tabanidae 

Colaspidea sp., prob. grata Fall Ch ysops coloradensis Bigot Hm 

Cyptocephalus sanguinicollis C. excitans Walker Hm 

sanzuinicollis Suffrian " Tabanus aegrotus Osten Sacken Hm 

Glyptoscelis juniperi Blake or nr. T. similis Macquart Hm 

Pachybrachis sp., nr. m-nigrum (Melsheimer) C Rhagionidae 

P. relictus Fall R ~~m~horom~ia sp. Pr 

Saxinis saucia LeConte C Therevidae 

Typophorus nigritus (FJ R species undet. ? 

continued 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 9 
Table 1, continued Table 1, continued 

1 Taxon 

Relative 

frequency" 

Stages 

encountered2 

Role in 

community3 Taxon 

Relative 

frequency1 

Stages 

encountered2 

Role in 

community3 

DIPTERA, continued DIPTERA, 

Vermileonidae Chamaemyiidae, continued 

Vermileo sp. Pachyneuron sp., either P. eros Girault 

Asilidae or P. mucronatum Girault 

Dioctria sp. (Hymenoptera: Pteromalidae) 

Asilus sp. Melanips sp. (Hymenoptera: Figitidae) 

Bombyliidae undet. sp. (Hymenoptera: Eulophidae) 

species (6)undet. Agromyzidae 

Empididae species (2)undet. 0 

Apalocnemis sp., nr.hirsuta Melander Heleomyzidae 

Rhamphomyia sp. Pseudolaria sp. R 

Dolichopodidae Suillia barberi (Darlington) C 

Medetera sp. S. hemorum (Meigen) C 

Phoridae S. limbata (Thomson) C 

species undet. Trixoscelididae 

Syrphidae Trixoscelis frontalis Fallen R 

Blera humeralis (Williston) T. tumida (Melander) R 

B. scitula (Williston) Chloropidae 

Chysotoxum fasciatum Miiller Liohippelates robertsoni (Sabrosky) R 

C. flavifrons Macquart Thaumatomyia bistriata (Walker) R 

Eupodes fumipennis (Thompson) T. rubida (Coquillett) R 

E. lapponicus (Zetterstedt) Anthomyiidae 

E. volucris Osten Sacken Delia platura (Meigen) R 

Helophilus latifrons Loew Muscidae 

Heringia sp. Caricia sp. R 

host: Tamalia coweni (Cockerell) Coenosia sp. 0 

(Homoptera: Aphididae) Fannia sp. C 

Ocyptamus diversifasciatus (Knab) Helina troene (Walker) C 

0.lemur (Osten Sacken) Limnophora sp. R 

Paragus haemorrhous Meigen Lispe neouliginosa Snyder R 

P. variabilis Vockeroth Neomyia cornicina (FJ R 

Scaeva pyrastri (L.) Pentacricia aldrichi Stein R 

Toxomerus marginatus (Say) Phaonia spp. (5) 0 

Tephritidae Schoenomyza dorsalis Loew 0 

Paracantha gentilis Hering Calliphoridae 

Tephritis sp. Eucalliphora latifrons (Hough) C 

Trupanea californica Malloch Phaenicia sericata (Meigen) R 

T.femoralis (Thomson) Phormia regina (Meigen) R 

Sepsidae Sarcophagidae 

Sepsis sp. Amobia aurifrons (Townsend) R 

Lauxaniidae Blaesoxipha sp. R 

Lauxania sp. Hilarella hilarella (Zetterstedt) 0 

Chamaemyiidae Metopia argyrocephala (Meigen) 0 

Leucopis (Leucopis) sp. Phrosinella pilosifrons Allen R 

host: Tamalia coweni (Cockerell) Ravinia lherminieri (Robineau-Desvoidy) 0 

(Homoptera: Aphididae) R. planifrons (Aldrich) R 

parasitoids reared from puparia: Senotainia trilineata (Wulp) C 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Table 1, continued Table 1, continued 

Relative Stages Role in Relative Stages 

Taxon frequency1 encountered2 community3 Taxon frequency1 encountered2 

DIPTERA, DIPTERA, 

Sarcophagidae, continued Tachinidae, continued 

Sphenometopa tergata (Coquillett) R Rindge (Lepidoptera: Geometridae) 

Taxigramma heteroneura (Meigen) R Siphona sp., nr. pacifica O'Hara R 

Tachinidae Stronggaster robustus (Townsend) 0 

Allophorocera sp., nr. occidentalis (Coquillett)0 Tachina sp. R 

Aplomya theclarum (Scudder) R Voria sp., nr. ruralis (Fallen) R 

Campylochaetasp. 0 

host: Synaxis cervinaria (Packard) LEPIDOPTERA 

(Lepidoptera: Geometridae) Hepialidae 

Cyptomeigenia sp. R Hepialis hectoides (Boisduval) 0 

Cylindromyia fumipennis (Bigot) R Tineidae 

Cyzenis sp. R species (2) undet. R 

Deopalpus sp., nr. contiguus (Reinhard) R Psychidae 

host: Hesperumia fumosaria impensa Hyaloscotes fumosa Butler 0 

Rindge (Lepidoptera: Geometridae) Gracillariidae 

Enrogalia morigera? Reinhard R species undet. (serpentine mine) C 

Eynnia tortricis (Coquillett) 0 species undet. (blotch mine) C 

hosts: Choristoneura rosaceana complex Coleophoridae 

(Lepidoptera: Tortricidae) Coleophora glaucella Walsingham 0 

Pseudochelaria manzanitae (Keifer) parasitoids reared from larvae: 

(Lepidoptera: Gelechiidae) Agathis sp. (Hymenoptera: Braconidae) 

Euclytia flava (Townsend) R Chelonus sp. (Hymenoptera: Braconidae) 

Eumea caesar (Aldrich) 0 Gelechiidae 

host: Choristoneura rosaceana complex Gelechia sp., nr. panella Busck 0 

(Lepidoptera: Tortricidae) Pseudochelaria manzanitae (Keifer) 0 

Eutrixa sp. 0 parasitoid reared from larvae: 

Lespesia sp. R Eynnia tortricis (Coquillett) (Diptera: Tachinidae) 

host: Saturnia mendocino Behrens Tortricidae 

(Lepidoptera: Saturniidae) Amorbia cuneana (Walsingham) R 

Leucostoma simplex (Fallen) R Choristoneura sp., rosaceana (Harris) 

Madremyia saundersii (Williston) R spp. complex C 

host: Hesperumia fumosaria impensa parasitoids reared from larvae: 

Rindge (Lepidoptera: Geometridae) Eynnia tortricis (Coquillett) (Diptera: Tachinidae) 

Microchaetina sp. R Eumea caesar (Aldrich) (Diptera: Tachinidae) 

Myatelemus? sp. R Nilea or Lespesia sp. (Diptera: Tachinidae) 

Nilea or Lespesia sp. R Epinotia arctostaphylana (Kearfott) C 

host: Choristoneura rosaceana complex E. miscana (Kearfott) C 

(Lepidoptera: Tortricidae) E. subplicana (Walsingham) C 

Periscepsia helymus (Walker) 0 E. terracoctana (Walsingham) C 

hosts: Apharetra californiae Pyralidae 

McDunnough (Lepidoptera: Noctuidae) Tulsa oregonella? (Barnes & McDunnough) R 

Aseptis ethnica Smith Hesperiidae 

(Lepidoptera: Noctuidae) Hesperia sp. I 

Phantasiomyia sp., prob. n. sp. R Lycaenidae 

Ph yxe pecosensis (Townsend) R Celastrina ladon (Cramer) R 

host: Hesperumia fumosaria impensa 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Table 1, continued Table 1, continued 

Relative Stages Role in Relative Stages Role in 

Faxon frequency1 encountered2 community3 Faxon frequency1 encountered2 community3 

.EPIDOPTERA, .EPIDOPTERA, 

Lycaenidae, continued Lasiocampidae, continued 

Incisalia augustus iroides Boisduval 0 L,P,rA M. constrictum (Henry Edwards) 

Nymphalidae M. disstria Hiibner 

Nymphalis californica (Boisduval) C P,rA Pyllodesma americana (Harris) 

Geometridae Lymantriidae 

Aethaloida packardaria (Hulst) R E,L,P,rA Orsyia cana Henry Edwards 

Anacamptodes clivinaria clivinaria (Guenee) 0 L,P,rA Saturniidae 

Apodrepanulatrix litaria (Hulst) R A Hemileuca eglanterina shastaensis (Grote) 

Chlorosea nevadaria Packard R A Hyalophora euyalus (Boisduval) 

Cyclophora dataria (Hulst) R A Saturnia mendocino Behrens 

Drepanulatrix rectifascia (Hulst) R A parasitoid reared from pupae: 

D. unicalcararia (Guenee) R A Lespesia sp. (Diptera: Tachinidae) 

Eupithecia sp. R L,P,rA Sphingidae 

Hesperumia fumosaria impensa Rindge 0 E,L,P,rA Hyles lineata (F.) 

parasitoids reared from larvae: Paonias myops (J. E. Smith) 

Deopalpus sp., nr. contiguus (Reinhard) (Diptera: Tachinidae) Sphinx vashti Strecker 

Madremyia saundersii (Williston) (Diptera: Tachinidae) Arctiidae 

Phyxe pecosensis (Townsend) (Diptera: Tachinidae) Spilosoma vestalis Packard 

Aleiodes nolophanae (Ashmead) (Hymenoptera: Braconidae) Noctuidae 

Itame guenearia (Packard) R A Acronicta perdita Grote 

I. quadrilinearia (Packard) 0 A Apharetra californiae McDunnough 

Nemoria darwiniata (Dyar) 0 A parasitoid reared from larvae: 

N. glaucomarginaria 0 E,L,P,rA Periscepsia helymus (Walker) (Diptera: Tachinidae) 

(Barnes& McDunnough) Aseptis ethnica Smith C L,P,rA 

Sabulodes edwardsata (Hulst) R A parasitoid reared from larvae: 

Semiothisa signaria dispuncta (Walker) R A Periscepsia helymus (Walker) (Diptera: Tachinidae) 

Sericosema juturnaria (Guenee) R A 

S. wilsonensis (Cassino & Swett) R A 3YMENOPTERA 

Synaxis cervinaria (Packard) 0 E,L,P,rA Stephanidae 

parasitoids reared from eggs: Schlettererius cinctipes (Cresson) R A 

Trichogramma sp. (Hymenoptera: Trichogrammatidae) Aulacidae 

Telenomus alsophilae Viereck (Hymenoptera: Scelionidae) Pristaulacus montanus (Cresson) R A 

parasitoids reared from larvae: Braconidae 

Campylochaeta sp. (Diptera: Tachinidae) Aleiodes nolophanae (Ashmead) R L,P,rA 

Aleiodes n. sp. (Hymenoptera: Braconidae) host: Hesperumia fumosaria impensa 

Meteorus rubens (Nees) (Hymenoptera: Braconidae) Rindge (Lepidoptera: Geometridae) 

Dusona nigritibialis (viereik) (~imeno~tera: Ichneumonidae) A. n. sp. R A 

Euplectrus sp., poss. plathypenae Howard (Hymenoptera: Eulophidae) host: Synaxis cervinaria (Packard) 

parasitoid observed attacking larval instar 111: (Lepidoptera: Geometridae) 

Goniozus gracilicornis (Kieffer) (Hymenoptera: Bethylidae) Agathis sp. R A 

predators observed attacking larvae: host: Coleophora glaucella Walsingham 

Misumenops celer (Hentz) (Araneae: Thomisidae) (Lepidoptera: Coleophoridae) 

Xysticus sp. (Araneae: Thomisidae) Apanteles sp. R A 

Metaphidippus sp. (Araneae: Salticidae) Bassus sp. R A 

Lasiocampidae Chelonus sp. R A 

Malacosoma californicum (Packard) 0 L,P,rA He(L) host: Coleophora glaucella Walsingham 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Table 1, continued Table 1, continued 

Relative Stages Role in Relative Stages Role in 

frequency1 encountered2 community3 Taxon frequency1 encountered2 community3 

HYMENOPTERA, HYMENOPTERA, 

Braconidae, continued Pteromalidae, continued 

(Lepidoptera: Coleophoridae) 

Cotesia sp. R 

Zatropis sp. 

host: Tamalia coweni 

R Pa&) 

Dolichogenidea cacoeciae (Riley) 

Macrocentrus gravitarsis Muesebeck 

host: genus and species undet. 

(Lepidoptera: Lymantriidae) 

Meteorus rubens (Nees) 

R 

R 

C 

(Cockerell) (Homoptera: Aphididae) 

Perilampidae 

Perilampus sp. 

Eurytomidae 

Sycophila sp. 

R 

R 

Pa(L) 

PaU 

host: Synaxis cervinaria (Packard) Figitidae 

(Lepidoptera: Geometridae) 

Trioxys sp. 

Urosigalphus sp. 

Ichneumonidae 

R 

R 

Melanips sp. 

host: Leucopis sp. 

(Diptera: Chamaemyiidae) 

Scelionidae 

R Pa&) 

Dusona nigritibialis (Viereck) 

host: Synaxis cervinaria (Packard) 

0 Telenomus alsophilae Viereck 

host: eggs of Synaxis cervinaria 

0 Pa(L) 

(Lepidoptera: Geometridae) 

species (7) undet. 

Trichogrammatidae 

Trichogramma sp. 

0 

0 

(Packard) (Lepidoptera: Geometridae) 

T. sp. 0 

host: eggs of Chlorochroa ligata 

(Say) (Heteroptera: Pentatomidae) 

Padl 

host: eggs of Synaxis cervinaria Chrysididae 

(Packard) (Lepidoptera: Geometridae) 

Eulophidae 

Aprostocetus sp., nr. hibus Burks 0 

Chrysis sp. 

Omalus glomeratus (Buysson) 

Encyrtidae 

R 

0 

P4L) 

Pa(L) 

host: Tamalia coweni 

(Cockerell) (Homoptera: Aphididae) 

Enyrtus sp. 

Bethylidae 

R Pa(L) 

Euplectrus sp., poss. plathypenae Howard 0 

host: Synaxis cervinaria 

(Packard) (Lepidoptera: Geometridae) 

Tetrastichus ch ysopae (Crawford) R 

Zagrammosoma sp. R 

genus and species undet. 0 

Goniozus columbianus Ashmead R 

G. gracilicornis (Kieffer) R 

host: Synaxis cervinaria 

(Packard) (Lepidoptera: Geometridae) 

Epyris clarimontensis Kieffer R 

Dryinidae 

Pad) 

Pat) 

Pa(L) 

host: Leucopis sp. 

(Diptera: Chamaemyiidae) 

Gonatopus sp. 

Sphecidae 

R Pad). 

Eupelmidae 

species undet. 

Torymidae 

Pseudotorymus sp. 

0 

R 

Ammophila azteca Cameron 

A. regina Menke 

Ancistromma capax (Fox) 

Aphilanthops subfrigidus Dunning 

R 

R 

C 

R 

Pr 

Pr 

Pr 

Pr 

Toymus tubicola (Osten Sacken) 0 Astata sp. 0 Pr 

Pteromalidae Eucerceris similis Cresson 0 Pr 

Catolaccus aineoviridis (Girault) R Go ytes sp. 0 Pr 

Gastrancistrussp. 

Pachyneuron sp., either P. eras Girault or 

R Harpactus rugulosus (Bohart) 

Isodontia elegans (Smith) 

R 

R 

Pr 

Pr 

P. mucronatum Girault R Mimesa serrana Finnamore R Pr 

host: Leucopis sp. 

(Diptera: Chamaemyiidae) 

Sceliphron caementarium (Drury) 

Steniolia tibialis Handlirsch 

R 

R 

Pr 

Pr 

Pteromalus sp. R Tnjpoxylon tridentatum Packard R Pr 

continued 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 13 
Table 1, continued Table 1, continued 

Relative Stages Role in Relative Stages Role in 

Taxon frequency' encountered2 community3 Taxon frequency1 encountered2 community3 

HYMENOPTERA, continued 

Halictidae HYMENOPTERA, continued 

Dialictus sp. 

Eylaeus sp. 

Andrenidae 

Andrena cerasifolii Cockerel1 

Nomadopsis sp. 

Anthophoridae 

Anthophora urbana (Cresson) 

Epeolus sp. 

Apidae 

Apis mellifera L. 

Bombus edwardsii Cresson 

B. vandykei (Prison) 

B. vosnesenskii Radowszkowski 

Psithyrus insularis (Smith) 

Mutillidae 

Dasymutilla aureola Cresson 

D. sackenii (Cresson) 

Sphaerophthalma sp. 

C 

R 

R 

R 

R 

R 

C 

R 

R 

C 

R 

0 

0 

R 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

Po 

Po* 

Po 

Po* 

Po 

Po* 

Po 

Po 

Po 

Po 

Po 

Pr 

Pr 

Pr 

Vespidae 

Ancistrocerus sp. 

Euodynerus sp. 

Stenodynerus sp. 

Vespula pensylvanica (Saussure) 

Formicidae 

Aphaenogaster occidentalis (Emery) 

Camponotus essigi M. R. Smith 

C. laevigatus (F.Smith) 

C. vicinus Mayr 

Crematogaster coarctata Mayr 

Formica haemorrhoidalis Emery 

F. podzolica Francoeur 

F. subpolita Mayr 

Leptothorax rugatulus Emery 

Myrmica sp. 

Solenopsis molesta (Say) 

Tapinoma sessile (Say) 

R 

R 

0 

C 

0 

C 

C 

C 

R 

C 

C 

C 

0 

C 

C 

C 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

A 

Pr 

Pr 

Pr 

Pr 

Fu 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pompilidae 

Ageniella coronata Banks 

A. euphorbiae (Viereck) 

Anoplius insolens (Banks) 

Aporinellus completus Banks 

Aporus luxus Banks 

Pompilus angularis Banks 

P. fumipennis eureka (Banks) 

P. solonis (Banks) 

Priocnemis notha navajo Banks 

R 

R 

R 

R 

R 

R 

C 

R 

R 

A 

A 

A 

A 

A 

A 

A 

A 

A 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Pr 

Relative frequency: I, incidental (transient species); R, rare (only a few individuals 

ncountered during five field seasons); 0,occasional (individuals encountered fairly 

requently); C, common (individuals locally abundant and encountered very fre- 

[uently). 

'Stages encountered: E, egg; I, immature; J, juvenile; N, nymph; L, larva; P, pupa or 

uparium; A, adult; rA, adult reared from immature form. 

'Dominant role in community (stage performing role if other than stage encounÂ¥red)Fu, fungivore; PO, pollinator; Wb, wood borer; He, herbivore; Pr, predator; Pa, 

~arasitoid; Hm, hematophage; Sc, scavenger; De, detritivore; Co, coprophage; X, no 

pparent direct association with manzanita. 

Voucher specimens retained by the USDA Agricultural Research Center, Beltsville, 

Maryland. 

+Voucher specimens retained by Crop Protection Division, Agriculture Canada, 

Ittawa, Ontario. 

Woucher specimens retained by the Florida State Arthropod Collection, Gainesville, 

lorida. 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Figure 5-Greenleaf manzanita leaves tied together with silk from an unidentified caterpillar in 

the family Gelechiidae or Tortricidae (Lepidoptera). 

Figure 6-Choristoneura st>., rosaceana (Harris) complex (Lepidoptera: Tortricidae) larva in an 

unraveled leaf bundle. This species generally completes its larval development before new leaves 

expand in the late spring and pupation occurs within a loose silken cocoon inside the tied leaf 

bundle. Damage can be locally severe. 

USDA Forest Service Gen.Tech. Rep. PSW-GTR-167. 1997. 

Foliovores (foliage-feeders) 

Leaf rollersltiers 

As soon as leaf buds break in the late spring or early summer, injury 

due to leaf rollers and tiers becomes evident (figs. 5-6). This type of 

damage is caused by a guild of caterpillars (Lepidoptera: 

Gelechiidae and Tortricidae). Numbers of tied leaf bundles fluctuate 

from year to year but the bundles are usually quite obvious. Larvae 

use silk to tie leaves together. The resulting bundle of leaves affords 

protection from natural enemies and provides a source of food as 

larvae feed on leaf tissue inside the bundle. After larvae fully 

mature, individuals of most species drop to the leaf litter to pupate 

leaving behind a loosely tied bundle of leaves containing frass or 

droppings. In time, the silken threads wear away, allowing leaves to 

expand and produce photosynthates. Although damage by leaf 

rollers and tiers often appears severe, greenleaf manzanita plants 

are usually able to withstand repeated yearly attacks by these 

foliage-feeders. 

Leaf miners 

Three types of leaf mines were frequently encountered throughout 

this study: serpentine, blotch, and hollow (figs. 7-9). Larvae causing 

mining damage feed between the outer two layers of a leaf and 

"mine" the inner layers leaving behind a characteristic damage 

pattern. Damage to greenleaf manzanita plants is typically minor 

because only a portion of the leaf is usually damaged. 

Gall formers 

The manzanita leaf gall aphid, Tarnalia coweni (Cockerell) 

(Homoptera: Aphididae), causes the edge of a newly expanding 

leaf to curl, thus causing the formation of a leaf gall. Damage to 

greenleaf manzanita plants can be quite obvious but not usually 

serious (fig. 10).A number of arthropods were found in association 

with this leaf gall aphid, including predators, scavengers, and 

opportunistic shelter-seekers (Valenti and others 1996).One other 

type of gall, formed within a newly expanding stem, was 

occasionally found but we were not able to identify the arthropod 

reponsible for its formation. 

Piercinglsucking insects 

Insects with piercing/sucking mouthparts (e.g., thrips [Thysanopteral, 

true bugs [Hemiptera], and homopterans ([Homopteral) suck plant 
Figure 7-Serpentine mine caused by an undetermined species (Lepidoptera: Gracillariidae). Figure 8-Blotch mine caused by an undetermined species (Lepidoptera: Gracillariidae). 

Figure 9Ã‘Characteristi leaf mine caused by Epinotia miscana (Kearfott) (Lepidoptera: Figure IO-Greenleaf manzanita plant infested with leaf galls caused by the manzanita leaf gall 

Tortricidae) (note adult exit hole near base of leaf). aphid, Tamolio coweni (Cockerell) (Hornoptera: Aphididae). Emerging alate forms are sexual 

produce eggs that overwinter. 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Figure 1 1-Non-gall forming aphids (Homoptera: Aphididae) congregated on a leaf (also note 

two leaf galls in the photograph). 

sap, usually from leaves. This type of feeding produces a necrotic 

stipling effect and, if severe enough, can cause serious damage to leaf 

tissue. Aphids (Homoptera: Aphididae) and leafhoppers (Homoptera: 

Cicadellidae) were common on greenleaf manzanita foliage during the 

growing season (fig. 11). Cicada adults (Homoptera: Cicadidae) 

periodically were locally abundant (fig. 12). Females oviposited in 

greenleaf manzanita stems but feeding was not observed. 

Defoliators 

Perhaps the most important natural enemies of greenleaf manzanita 

are the defoliators, defined here as those insects that possess chewing 

mouthparts. Damage can be severe because entire leaves can potentially 

be removed in a relatively short time period by a single insect. Greenleaf 

manzanita defoliators belong to one of three insect orders; Orthoptera, 

Coleoptera, or Lepidoptera. Grasshoppers (fig. 13) and katydids 

(Orthoptera) were often common as were members of several beetle 

(Coleoptera) families including scarabs (Scarabaeidae) (fig. 14), leaf 

beetles (Chrysomelidae), and weevils (Curculionidae). Lepidoptera 

(moths and butterflies), however, were the most common defoliators 

encountered. This group of insects is represented by numerous families 

and species (Valenti and Zack 1995) (figs. 15-24). 

Figure 12-Adult cicada, Platypedia putnami (Uhler) (Homoptera: Cicadidae), resting on a Figure 13-Grasshoppers, such as this immature Melanoplus sp. (Orthoptera: Acrididae), are 

greenleaf rnanzanita branch. In 1993 adult females were commonly encountered at the Hat quite common in greenleaf manzanita brushfields. 

Creek site notching and depositing eggs in greenleaf rnanzanita branches. 

USDA Forest Service GemTech. Rep, PSW-GTR- 167. 1997. 
Figure 14-Two color variants of the ubiquitous scarab Dicheloryx bad4 (Kirby) (Coleoptera: Figure 15-A case bearer, probably Coleophora glaucella Walsingham (Lepidoptera: 

Scarabaeidae). Adults were commonly encountered feeding on greenleaf manzanita leaves Coleophoridae), and its charactersitic damage to a greenleaf manzanita leaf. 

making characteristic small notches on leaf edges. 

Figure 1 &The slug-like caterpillar of lncisalia augustus iroides Boisduval (Lepidoptera: Lycaenidae). Figure 17-Eady I d of Packordoria (Hulst) (Lepidoptera: Geometridae) in a 

This larva grazes the surfaces of leaves and fruits. typical resting postureon &a Â¥dg <rf a leaf. 

USDA Forest Service Gen.Tech. Rep. PSW-GTR-167. 1997. 
Figure I(Kareen larvalcolor variant ofHesperumio furnosaria impensa Rindge (Lepidoptem Geometridae) Figure 19-An unusual greenleaf manzanita inflorescence mimic, Nemoria glaucomarginaria 

that mimics newly expanding manzanita shoots. A second crimson larval color variant of this species (Barnes & McDunnough) (Lepidoptera: Geometridae). 

occurs concurrently and mimics mature manzanita stems. Larvae feed on previous years' leaves and 

complete development by late spring generally before new leaves mature. Pupation occurs in the leaf 

litter and adults are active during June and July. Eggs deposited in litter overwinter. 

Figure 20-Synaxis cervinaria (Packard) (Lepidoptera: Geometridae) fourth larval instar. Larvae Figure 21-Mature larva of the ceanothus silk moth, Hyalaphora euryalus (Boisduval) 

mimic dead and dying manzanita twigs (see Valenti and others [I997 for details on the biology (Lepidoptera: Saturniidae). 

and ecology of this species). 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Figure 22-Adult male ceanothus silk moth, Hyalophora euryalus (Boisduval) (Lepidoptera: 

Saturniidae), on top of its cocoon soon after emergence. 

Figure 24-An unusual noctuid larva, Acmnicta perdfta Grote (Lepidoptera: Noctuidae). 

Figure 23-A hawk moth larva. Sphinx vashti Strecker (Lepidoptera: Sphingidae). 

Predators 

Predators generally consume all or part of their prey soon after 

capture. During the life of the predator more than one prey individual 

is killed and often dozens or even hundreds are consumed. A total of 

161 invertebrate predator taxa was collected in greenleaf manzanita 

communities representing 11 orders and 51 families. Snakeflies 

(Neuroptera: Raphidiidae) were common during the greenleaf 

manzanita growing season (fig. 25). These insects feed on soft-bodied 

arthropods such as aphids and leafhoppers. One individual was 

observed feeding on a small lepidopteran caterpillar. Another quite 

common predator group related to the snakefly includes green 

lacewings (Neuroptera: Chrysopidae) (fig. 26). Green lacewing eggs, 

larvae, and adults can be found easily with a minimal amount of 

searching during the summer months. These insects, like the 

snakeflies, feed on soft-bodied arthropods. Dragonflies and 

damselflies (Odonata) capture their prey on the wing and can be 

abundant at certain times of the year during hatches (mass adult 

emergences). These "mosquito hawks" were particularly abundant 

at the Logan Lake site (fig. 27). 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997 20 
I 

Figure 25-A male snakefly, Raphidia unicolor (Carpenter) (Neuroptera: Raphidiidae). Snakeflies 

feed on soft-bodied arthropods and are extremely common during the summer months. 

Figure 26-Green lacewing adult (Neuroptera: Chrysopidae). 

The most important invertebrate predators were ants 

(Hymenoptera: Formicidae). Ants were commonly encountered at all 

study sites, both on the ground and on greenleaf manzanita plants. 

Most ant species are generalist predators and numerous studies have 

shown them to be very important in regulating many insect 

populations (Finnegan 1971, Laine and Niemela 1980, Youngs 1983, 

Valenti and others 1998). 

Spiders (Araneae) are probably the second most important 

invertebrate predators in greenleaf manzanita communities. A variety 

of spiders, including orb builders (fig. 28), crab spiders (fig. 29), and 

jumping spiders (fig. 30), were encountered. More than .50 species 

were collected but have not yet been identified. 

Parasitoids 

The term parasitoid is used to describe a parasite that kills its host 

(usually only one), in contrast to a true parasite that feeds without 

killing its host, such as blood-feeding insects. Eight blood-feeders 

Figure 27-Damselfly (Odonata) adult resting on a greenleaf manzanita leaf. 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Figure 28-Unidentified orb weaver spider (Araneae). Figure 29-Unidentified crab spider (Araneae: Thomisidae) feeding on a larva of Synaxis 

cervinaria (Packard) (Lepidoptera: Geometridae). 

were encountered in greenleaf manzanita brushfields. These included 

a tick (Acari: Ixodidae), two black flies (Diptera: Simuliidae), a midge 

(Diptera: Chironornidae), and four deer flies (Diptera: Tabanidae). 

Seventy-nine parasitoids were collected from two orders and 19 

families, many of which we reared from known hosts (table 1). 

Adult tachinid flies (Diptera: Tachinidae) were commonly 

encountered on and around greenleaf manzanita foliage (fig. 31). 

Female flies locate a suitable host and deposit eggs or larvae on the 

host's body. Larvae eventually work their way into the host, feed, 

develop, and remain there until pupariation occurs. The host is kept 

alive for nearly the entire time it takes for the fly to develop. This is 

accomplished by the fly larva first feeding on non-vital host tissues. 

A number of parasitic Hymenoptera were also collected by rearing 

them from their lepidopteran hosts (figs.32-33). 

Scavengers 

Sixty-nine scavengers, or organisms that feed on waste material, 

Figure 30-Unidentified jumping spider (Araneae: Salticidae). were captured (8orders and 29 families). Many were collected in 

pitfall traps. Included in this group are detritivores and 

USDA Forest Service Gen.Tech. Rep. PSW-GTR-167. 1997. 
Figure 3 1-Adult tachinid fly (Diptera: Tachinidae) resting on manzanita leaf. 

Figure 33-Fifth larval instar of Synaxis cewinaria (Packard) (Lepidoptera: Geometridae) being 

parasitized by Euplectrus sp., possibly plathypenae Howard (Hymenoptera: Eulophidae). 

Figure 32-Parasitoid cocoons (Hymenoptera: Braconidae) on a mature ceanothus silk moth 

larva, Hyalofihora euryalus (Boisduval) (Lepidoptera: Satumiidae). 

coprophagous forms. Detritivores feed on dead organic material 

and were common. Representatives of three families of springtails 

(Colembola: Isotomidae, Entomobryidae, and Sminthuridae), one 

family of jumping bristletails (Microcoryphia: Machilidae), a stag 

beetle (Coleoptera: Lucanidae), crane flies (Diptera: Tipulidae), and 

a lauxaniid fly (Diptera: Lauxaniidae) were encountered. 

Coprophagous forms feed on dung and are also decomposers and 

are represented by two scarab beetles (Coleoptera: Scarabaeidae). 

The majority of scavengers collected, such as rnuscid flies (fig. 341, 

feed on a variety of waste materials and all play an important role in 

decomposition and mineral recycling. 

Conclusions 

Northeastern California montane chaparral communities dominated 

by greenleaf manzanita harbor a rich diversity of insect and related 

arthropod fauna. Because of the need for alternatives to control 

competing forest vegetation in the future, arthropod diversity 

studies will become increasingly important. This inventory provides 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
Acknowledgments 


This study was done under Cooperative Agreement No. PSW-89-0017CA, between 

the Department of Entomology, Washington State University, Pullman, and the Pacific 

Southwest Research Station, USDA Forest Service, Redding, Calif. 

We thank the many cooperating taxonomists whose efforts and expertise are 

Figure 34-Adult rnuscid fly (Diptera: Muscidae) (larvae are scavengers). 

basic biological information about insect fauna associated with 

greenleaf manzanita that can be used in future studies. For example, 

a portion of the herbivorous insects encountered in this study have 

the potential to cause serious damage to greenleaf manzanita plants 

after they are released from their natural regulating agents. If this 

release can be accomplished on a large scale, it represents an 

ecologically sound method of altering vegetation composition to 

favor more desirable plant species such as high-value conifers. 

gratefully appreciated: 

The Burke Museum, University of Washington, Seattle: 

California Academy of Sciences, San Francisco: 

Centre for Land and Biological Resources Research, 

Agriculture Canada, Ottawa, Ontario, Canada: 

Florida Agricultural and Mechanical University, Tallahassee: 

Florida State Department of Agriculture, Gainesville: 

Houston, Texas: 

International Institute of Entomology, London, England: 

Liverpool Polytechnic, Liverpool, England: 

The Ohio State University, Columbus: 

Oregon State Department of Agriculture, Salem: 

Texas A & M University, College Station: 

University of Alberta, Edmonton, Alberta, Canada: 

University of California at Berkeley: 

University of Connecticut, Storrs: 

University of Florida, Gainesville: 

R. Crawford 

R. Robertson 

K.B. Bolte 

G. Gibson 

K.G.A.Hamilton 

J. Huber 

J.-F. Landry 

L. Masner 

J. Read 

M. Sharkey 

J.R. Vockeroth 

C.W. O'Brien 

L.B. O'Brien 

W.C. Welbourn 

D. Heffern 

J. LaSalle 

I.D. Hodkinson 

C.A. Triplehorn 

R.L. Wescott 

J. Jackson 

G.E. Ball 

K.S. Hagen 

S. O'Keefe 

J.A. Powell 

D.L. Wagner 

D.H. Habeck 

P. Skelley 

USDA Forest Service Gen.Tech. Rep. PSW-GTR- 167. 1997. 
University of Idaho, Moscow: 

University of Mississippi, University: 

University of New Hampshire, Durham: 

University of Wisconsin, Madison: 

U.S. Department of Agriculture, Olympia, Wash.: 

U.S. Department of Agriculture Systematic Entomology Laboratory, 

Beltsville, Maryland, and cooperating institutions: 

Special thanks are extended to R.S. Zack, M.T. James Entomological Museum, 

Washington State University, Pullman, for his assistance, guidance, and genuine 

interest in the identification and curation of this assemblage of species. 

W.F. Barr (retired) 

J.B.Johnson 

P.K. Lago 

D.S. Chandler 

P.J. Johnson 

D. Suomi 

C.H. Dietrich 

R.J. Gagnk 

T.J. Henry 

A.S. Menke 

J. Pakaluk 

M.E. Schauff 

F.C. Thompson 

N.E. Woodley 

R.S. Hanley 

T.D. Miller 

J. Shepard 

W.J. Turner 

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USDA Forest Service Gen.Tech. Rep. PSW-GTR-167. 1997. 
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United States Insects and Related Arthropods Associated with 

Department Greenleaf Manzanita in Montane Chaparral of Agriculture 

Communities of Northeastern California .. 

Forest Service 

Pacific Southwest 

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General Technical Report 

PSW-GTR- 167