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Home , Acalypta, Aphodius fossor, Boreus, Buprestis, Buprestis aurulenta, Chionea

AMER. ZOOL., 33:578-587 (1993)

Arthropod Diversity and Conservation in Old-Growth mon et al., 1990; Hz Northwest complex litter layer 1973; Lattin, 1990; JOHN D. LATTIN and other features Systematic Entomology Laboratory, Department of Entomology, State University, tural diversity of th Corvallis, Oregon 97331-2907 is reflected by the 14 found there (Lawtt SYNOPSIS. Old-growth forests of the extend along the 1990; Parsons et a. e coastal region from southern Alaska to northern and are com- While these old posed largely of rather than . Many of these ity over time and achieve great age (500-1,000 yr). Natural succession that follows product of sever: stand destruction normally takes over 100 years to reach the young through successioi mature forest stage. This succession may continue on into old-growth for (Lattin, 1990). Fire centuries. The changing structural complexity of the forest over time, and diseases, are combined with the many different species that characterize succes- bances. The prolot sion, results in an array of habitats. It is estimated that 6,000 a continually char arthropod species may be found in such forests—over 3,400 different ments and habitat species are known from a single 6,400 ha site in Oregon. Our knowledge (Southwood, 1977 of these species is still rudimentary and much additional work is needed Lawton, 1983). throughout this vast region. Many of these species play critical roles in have lx the dynamics of forest ecosystems. They are important in nutrient cycling, old-growth site, tt as herbivores, as natural predators and parasites of other arthropod spe- mental Forest (HJ cies. This faunal diversity reflects the diversity of the environment and old-growth Dougi the arthropod complex provides a sensitive barometer of the conditions lamette National of the forest. Conservation efforts for forest arthropods are limited at Oregon. The HJA present and controlled largely by land-use policies. For example, an effort by the U.S. Fore is being made to include arthropods in conservation efforts for the North- consequences of ern and arthropods will be included in the Forest Health ulation. It was a Monitoring program now underway by the U.S. Forest Service. Evidence western Coniferot: from other parts of the world suggest that arthropods that depend upon International Bio large pieces of dead may be particularly threatened by forest man- 1960s, was nun( agement practices. Much remains to be done in the conservation of forest logical Reserve ii arthropods. one of the first Na INTRODUCTION carpus Blume, L., Quercus L., and funded, Long-Tc Sites in 1980. Toe The old-growth forests of the Pacific Salix L.) (Franklin and Dyrness, 1971). The large trees develop extensive canopies char- on the site and c( Northwest form a distinctive biotic region tigations relate( that extends along the Pacific Coast from acterized more by their height than breadth. These conifer species achieve great age. Included in these southeast Alaska to northern California. many involving Waring and Franklin (1979) characterized Many of the oldest individuals of each spe- cies are found in these forests (e.g., Sitka lated informatior these forests as unique in their size and in paper. the longevity of the individuals of the dom- [ (Bong.) Carr.] >500 inant tree species, resulting in very large yr, Douglas- [ menziesii ARTHROPODS 0 accumulations of biomass. The 25 domi- (Mirb.) Franco] >750 yr, Alaska yellow- cedar [Chamaecyparis nootkatensis (D. Don) were fi nant tree species are rather than Northwest betwt although there are hardwood Spach] > 1,000 yr, western red cedar [ Donn ex D. Don] > 1,000 yr, coast John Kirk Town taxa found in the forests (e.g., Acer L., Alnus Philadelphia (H Mill., Castanopsis (D. Don) Spach, Litho- redwood [ (D. Don) Endl.] >1,250 yr) (Waring and Franklin, he collected spt 1979). Other related attributes of the forests growth forest i From the Symposium The Crisis in Invertebrate are the presence of massive amounts of dejeani, describe Conservation presented at the Annual Meeting of the (CWD) as snags or on ist, Louis Reich( American Society of Zoologists and the Canadian Soci- less, nocturnal ti ety of Zoologists. 27-30 December 1992, at Vancou- the ground (Maser and Trappe, 1984; Har- ver, . mon et al., 1986; Maser et al., 1988; Har- of Omus, collect 578 V le

OLD-GROWTH FOREST ARTHROPOD CONSERVATION 579

mon et al., 1990; Harmon, 1992) and a deep, also was described by Reiche. These were complex litter layer (Franklin and Dyrness, the first insects to be recorded in the sci- 1973; Lattin, 1990; Moldenke, 1990). These entific literature from the Pacific Northwest and other features contribute to the struc- (Hatch, 1949). mversity, tural diversity of the forests—diversity that Other collectors came to the region in the is reflected by the diversity of arthropods years that followed. M. H. Hatch (1949) found there (Lawton, 1983; Asquith et al., provided an interesting account of the his- long the 1990; Parsons et al., 1991). tory of entomology in the Pacific Northwest. re corn- While these old forests have great stabil- One collector/sportsman was Lord Wal- of these ity over time and space, the forests are the singham from England who traveled through ollows product of several centuries of change Oregon and California in 1871 and 1872 young through succession following collecting moths while on a hunting expe- h for (Lattin, 1990). Fire and wind, besides insects dition. This trip resulted in a slim volume time, and diseases, are the main natural distur- on the plume moths of California and Ore- !succes- bances. The prolonged succession results in gon, complete with colored plates of most t 6,000 a continually changing mosaic of environ- species (Walsingham, 1880). At the time, ifferent ments and habitats available to arthropods European specialists wrote most of the sci- owledge (Southwood, 1977; Southwood et al., 1979; entific literature based upon these early col- r -eded Lawton, 1983). Over 3,400 species of lections. As the area became populated and is in arthropods have been documented from one institutions established, there was a gradual ling, old-growth site, the H. J. Andrews Experi- accumulation of knowledge on the arthro- spe- mental Forest (HJA). This site is a 6,400 ha pods of the region. The number of species tent and old-growth Douglas-fir forest in the Wil- involved, and the relatively few scientists nditions lamette National Forest east of Eugene, present, made progress slow. Even today, mited at Oregon. The HJA was established in 1948 the problem of numbers of species and the an effort by the U.S. Forest Service to examine the shortage of specialists continues. There are e North- consequences of long-term habitat manip- still many taxa largely unstudied and ,t Health ulation. It was a major component of the unknown (Stanton and Lattin, 1989).By way Evidence western Coniferous Forest Biome during the of example, the collection of the Systematic .nd upon International Biological Programme in the Entomology Laboratory, Department of •est man- 1960s, was named an Experimental Eco- Entomology, Oregon State University, was of forest logical Reserve in the 1970s, and became initiated over 100 years ago. Today, the col- one of the first National Science Foundation lection contains over 2,500,000 specimens Quercus L., and funded, Long-Term Ecological Research with heavy emphasis on the Pacific North- rrcss, 1971). The Sites in 1980. Today, over 50 scientists work west. It contains thousands of unstudied inopies char- on the site and conduct a variety of inves- specimens because there are no specialists than breadth. tigations related to forest ecosystems. for many groups. :eve great age. Included in these and past investigations are Our knowledge of insects in forest eco- luals of each spe- many involving arthropods. This accumu- systems grew in several ways. As interest in 3rests (e.g., Sitka lated information provides the basis for this these productive forests increased, there was ong.) Carr.] > 500 paper. increased interest in the arthropods of the , menziesii forests, chiefly insects. Emphasis was on ARTHROPODS OF OLD-GROWTH FORESTS Alaska yellow- those species that were considered pests for tkatensis (D. Don) Insects were first collected in the Pacific one reason or another. Eventually this red cedar [Thuja Northwest between 1834 and 1836 by Dr. knowledge was synthesized by Keen (1952) > 1,000 yr, coast John Kirk Townsend, an ornithologist from and, more recently, by Furniss and Carolin rvirens (D. Don) Philadelphia (Hatch, 1949). Remarkably, (1977). The latter volume contains a vast ng and Franklin, he collected specimens of a typical old- amount of information on western forest mtes of the forests growth forest , the , Omus insects, including many that occur beyond ;sive amounts of dejeani, described by the French coleopter- the region considered here. Central to the "D) as snags or on ist, Louis Reiche, in 1838. This is a flight- acquisition of such information were the rappe, 1984; Har- less, nocturnal . A second species Canadian and government- et a!., 1988; Har- of Omus, collected during the same period based forest services. Other major sources JOHN D. LATTIN of information were the many taxonomic work expanded, many additional specimens ■ A substantial nn revisions of major taxa found in forests (e.g., were added (Lewis and Maser, 1981; Ander- ciated with the wood boring , , loopers, and son et al., 1982; Lightfoot, 1986). Emphasis debris (Schow spiders). These studies provided the back- was placed on aquatic taxa, the insects asso- (1984), Warre drop to studies focussing upon arthropods ciated with different plant species, and Peterken (1992 found in old-growth forests in western North extensive collections from the litter and soil ments of this fa America. Without the previous work of lit- habitats (Moldenke and Fichter, 1988; Lat- upon adequate erally hundreds of individuals, studies on tin and Moldenke, 1990; McIver et al., 1990; and are likely old-growth forests would have been Moldenke and Lattin, 1990a, b). Canopy (Table 1). extremely difficult or impossible. studies were conducted 70 m overhead and A small but dis resulted in a surprising number of species enous species The international biological programme (Voegtlin, 1982; Moldenke et al., 1987; est.This contra The International Biological Programme Schowalter, 1989, 1990; Schowalter et al., non-indigenou (IBP), started in the 1960s, increased inter- 1986). There were studies of the fauna of 1). est in ecosystem structure and function. One early successional trees as well (Schowalter While these sub-unit of the IBP in was et aL, 1986). The role of arthropods in the appear rather the Western Coniferous Biome. Within this long-term decomposition of wood received tance, they col group, attention was given to the structure special attention and many species resulted Approximatel and function of coniferous forests, including from these studies (Deyrup, 1975; Carpen- terflies knowr old-growth forests along the Pacific Coast. ter et al., 1988; Moldenke and Fichter, 1988; found on the Many reports and publications resulted from Schowalter et al., 1992). After a number of (: these efforts, including some on arthropods draft editions, the annotated list of over of 659 are fou (e.g., Deyrup, 1975; Mispagel and Rose, 3,400 species from the Andrews Forest was the 260 found 1978). As investigations proceeded, the rec- published (Parsons et al., 1991). This doc- ognition of the role arthropods play in such ument, combined with many other papers CONSERVATI ecosystems increased. Initial emphasis was and reports, forms the basis for our present Ow-C more on functional group than on the spe- knowledge. We owe a debt of gratitude to cies composition. As taxonomic and bio- the many people who contributed to that Our knowled[ logical information increased, the role of volume. old-growth fore some species in the systems became clari- During the completion of the annotated we know such fied. The variety of studies carried out pro- list of arthropods for the H. J. Andrews For- able and essenti vided vast numbers of specimens of many est, many interesting details emerged. A few servation stratel taxa that accumulated at different localities. of these are cited below. have been mad One such site was the H. J. Andrews Exper- we are forced tc imental Forest in western Oregon. Although they are considered quite mesic few sites where forests, a small, but distinct xeric arthro- Additional studi THE H. J. ANDREWS ARTHROPOD pod fauna is present. More common are some undisturl COLLECTION warm-dry adapted species found chiefly son, 1988; Latt This collection had its origins in the Inter- to the south, less common are the cold- national Biological Programme of the 1960s. dry adapted species whose distribution is The Problem The H. J. Andrews Experimental Forest was northern. Monitoring the populations of Much of our established by the U.S. Forest Service in warm-dry species should be of value in arthropods of of 1948 and when the IBP was organized, the detecting response to Global Climate the informatio site became an important part of the West- Change (Table 1). Andrews Expe ern Coniferous Biome program. Some of the A rich and diverse litter/soil arthropod Oregon (Mispa, forest ecosystem research at that time fauna, regularly containing 250 species per et al., 1990; Mc involved arthropods and the specimens from m2. This fauna plays an important role Moldenke, 19( that era formed the basis for the present in nutrient cycling (Moldenke, 1990). 1990 a, b; Pars collection (Mispagel and Rose, 1978). A very distinct fauna of flightless species, haps 50-60 per I joined the H. J. Andrews Research chiefly beetles but including some of the There is, of cou Group in 1976 and began to organize and true bugs. These are species of low vagil- on taxa found ii build the collections with the cooperation ity and certain to be impacted by forest Furniss and C of over 100 individuals. As the arthropod fragmentation (Lattin, 1990) (Table 1). Bright, 1987; ar,

OW—GROWTH FOREST ARTHROPOD CONSERVATION 581 itional specimens A substantial number of arthropods asso- most such information is not site-specific ser, 1981; Ander- ciated with the abundant coarse woody enough to know whether it applies to old- 1986). Emphasis debris (Schowalter et a!., 1992). New growth forests. Since it is unrealistic to , the insects asso- (1984), Warren and Key (1991), and assume that there will be massive efforts to lilt species, and Peterken (1992) indicated that some ele- acquire such information for all taxa, it the litter and soil ments of this fauna are entirely dependent seems more reasonable to identify repre- ichter, 1988; Lat- upon adequate amounts of large, old wood sentatives of key groups that belong to dif- [diver et al., 1990; and are likely targets for conservation ferent functional guilds or occur in habitats ,90a, b). Canopy (Table 1). unique to these forests (e.g., coarse woody m overhead and A small but distinct number of non-indig- debris) for study, collection, and sampling umber of species enous species (56 of 3,400) on the for- (Goldsmith, 1991). Some of the important ke et al., 1987; est.This contrasts to 41 of 480 species of terrestrial functional groups include: polli- ichowalter et a!., non-indigenous on the site (Table nators, foliage feeders, xylophages, litter and s of the fauna of 1). fungal comminutors, root feeders, preda- well (Schowalter While these old-growth forests may tors, and parasitoids. There are appropriate arthropods in the appear rather homogeneous from a dis- aquatic counter groups. Some examples of of wood received tance, they contain a mosaic of habitats. proposed efforts follow: y species resulted Approximately half of the species of but- ,p, 1975; Carpen- terflies known to occur in Oregon are Arthropod diversity and forest health hter, 1988; found on the H.J.A. For the true bugs A national program has been established if, number of (Hemiptera: ), 206 species out to monitor the health of forests under the list of over of 659 are found there and 134 genera of direction of the United States Forest Service ,drews Forest was the 260 found in the state. and in cooperation with the Environmental 1991). This doc- Protection Agency. The goals of the Forest any other papers CONSERVATION OF ARTHROPODS OF Health Monitoring Program are "to detect ;is for our present OLD—GROWTH FORESTS changes and trends in forest conditions and bt of gratitude to to report and interpret these changes at a ,ntributed to that Our knowledge of arthropod diversity in multistate level." Fourteen states have or old-growth forests is limited even though are developing forest health monitoring of the annotated we know such information is highly desir- programs. A plan for Oregon is being devel- 1. J. Andrews For- able and essential if we are to develop con- oped while that for California is nearing Is emerged. A few servation strategies. So few detailed studies completion. This nation-wide program is have been made at different localities that superimposed upon the EPA-EMAP grid we are forced to extrapolate from the very established several years ago. Approxi- dered quite mesic few sites where we have some knowledge. mately 4,500 of the 12,600 plots established tinct xeric arthro- Additional studies are urgently needed while are considered forested. While species lore common are some undisturbed forests remain (Morri- will receive special attention, we feel that it.s found chiefly son, 1988; Lattin and Moldenke, 1992). the inclusion of representatives of different an are the cold- functional groups will provide valuable ose distribution is The Problem information about the health of forested he populations of Much of our specific knowledge about the ecosystems. At the same time, more wide- rid be of value in arthropods of old-growth forests comes from spread knowledge about the distribution and Global Climate the information gathered on the H. J. abundance of forest arthropods will result, Andrews Experimental Forest in western making conservation efforts more effective. er/soil arthropod Oregon (Mispagel and Rose, 1978; Asquith We are developing sampling protocols to ing 250 species per et a!., 1990; McIver et al., 1990; Lattin and deal with selected taxa found in forested in important role Moldenke, 1990; Moldenke and Lattin, ecosystems (Table 2). The numbers of spe- ,ldenke, 1990). 1990 a, b; Parsons et al., 1991) where per- cies and individuals of each species are high - flightless species, haps 50-60 per cent of the fauna is known. for arthropods. These very numbers create iding some of the There is, of course, a considerable literature logistical problems when large-scale sam- .,cies of low vagil- on taxa found in forests (e.g., Prentice, 1962; pling efforts occur. Still, the fine resolution npacted by forest Furniss and Carolin, 1977; Danks, 1979; provided by these numbers makes the effort 1990) (Table 1). Bright, 1987; Danks and Foottit, 1989) but worthwhile. 582 JOHN D. LATTIN C

TABLE 1. Examples of old-growth forest arthropods. Warm xeric insects Coleoptera Hemiptera: Heteroptera : 0601 Berytidae: Acanthophysa echinata Uhler MO, Lygaeidae: Malezonotus obrieni Ashlock - SitOt Thylochromus nitidulus Barber : Aphoc : Macrotylus essigi Ban Duzee Macrotylus multipunctatus Van Duzee Cold xeric insects The National Park Hemiptera: Heteroptera Inventory and mot : Vanduzeeina borealis californica Van Duzee : Acalypta lillianus Torre-Bueno While not direc Flightless insects National Park Seri Orthoptera tory and monitor Acrididae: Boonacris alticola Rehn and Randall. Patterson, 1992). Gryllacrididae: Pristoceutophilus spp. to a 1988 Manage Tropidischia xanthostoma (Scudder) "assemble baselin Hemiptera: Heteroptera ing the natural res Lygaeidae: Plinthisus longisetosus Barber ship and will mo Tingidae: Acalypta saundersi (Downes) to detect or predit Coleoptera will generate info] Carabidae: Cychrus tuberculatus Harris tebrate conservz Metrius contract us Eschscholtz included in such Promecognathus laevissimus Dejean see that indeed th Zacotus matthewsi LeConte Cicindelidae: Omus dejeani Reiche Curculionidae: Lobosoma horridum Mannerheim TABLE 2. Exa Boreidae: Boreus sp. Organism Diptera Tipulidae: Chionea sp. Acarina Oribatid mites Coarse woody debris arthropods Isopoda Diplopoda Ligidiidae: Ligidium gracile Dang Insecta Collembola Isoptera Springtails Hodotermitidae: Zootermopsis angusticollis (Hagen) Coleoptera : aurulenta Linnaeus Insecta Cerambycidae: Leptura obliterata Haldeman Hemiptera: Heter Lucanidae: Playcerus oregonensis Westwood Platypodidae: wilsoni Swaine Seed bugs (Lygl Scolytidae: Dendroctonus pseutotsugae Hopkins Trogositidae: Ostoma ferruginea (Linnaeus) Plant bugs (Mil Formidicae: Camponotus spp. Coleoptera Siricidae: Urocerus albicornis (Fabricius) ! Diptera beetles (5_ : cinctus DeGeer Non-indigenous insects Hymenoptera Hemiptera: Heteroptera Sawflies (Symi Lygaeidae: Megalonotus sabulicolus (Thomson) sabulosus (Schilling) Caterpillars Miridae: Megaloceroea recticornis (Geoffroy) binotatus (Fabricius) Adults OLD-GROWTH FOREST ARTHROPOD CONSERVATION 583

TABLE 1. Continued. Coleoptera Curculionidae: Otiorhynchus ovatus (Linnaeus) Otiorhynchus sulcatus (Fabricius) (Linnaeus) Scarabaeidae: fossor Linnaeus

The National Park Service: (1992) has reviewed the state of inventories Inventory and monitoring plan in southwestern U.S. National Parks and While not directed at forests alone, the included both aquatic and terrestrial inver- National Park Service is initiating an inven- tebrates. The present level of knowledge tory and monitoring program (Rugh and about the invertebrates was quite low. We Patterson, 1992). This effort is in response initiated a similar idea with Acadia National to a 1988 Management Policy directive to Park a few years ago and it now appears it "assemble baseline inventory data describ- may become a reality. ing the natural resources under its steward- ship and will monitor those resources .. . Northern spotted owl conservation to detect or predict changes." These efforts There have been extensive investigations will generate information of value to inver- on the northern spotted owl as an indicator tebrate conservation provided they are of old-growth forests. The threatened status included in such efforts. We are working to of the bird throughout much of its range has see that indeed they are included. Stolhgren resulted in the proposal to establish Habitat

TABLE 2. Examples of arthropod taxa for monitoring with functional roles and sampling methods.

Functional Sampling Organism role method Acarina Oribatid mites fungal, litter high-gradient feeders, predators extractors Diplopoda Millipedes litter communitors pit-fall traps Collembola Springtails litter/soil Berlese funnels, organic matter high-gradient feeders extractors Insecta Hemiptera: Heteroptera Seed bugs (Lygaeidae) seed feeders pit-fall traps on ground Plant bugs (Miridae) foliage feeders, beating foliage predators Coleoptera Ground beetles (Carabidae) Predators, pit-fall traps seed feeders Bark beetles (Scolytidae) xylophagous pheromone, intercept traps Hymenoptera Sawflies (Symphyta) foliage feeders malaise traps Lepidoptera Caterpillars foliage feeders beating, branch clipping Adults nectar feeders blacklight traps JOHN D. LATTIN

Conservation Areas (HCAs) for its conser- the location of such should be vation. We have provided information on selected with care. potentially sensitive arthropod species likely to benefit from the establishment of these Importation of raw logs Diplopoda Xestodermidae: HCAs (Lattin and Moldenke, 1992) (Table The decline in the availability of logs for haydeni. 3). As mentioned several times, these rec- mills in the west has increased the interest ommendations represented extrapolation in importing raw logs from other countries. Insecta from information from the H. J. Andrews It has not stopped the export of raw logs Orthoptera Acrididae: Experimental Forest. We will feel more from our region, however. Over the past two Boonacris altica comfortable when sources from other local- years, there has been intense interest in Gryllacrididae: ities are available. bringing in raw logs from Siberia, New Zea- Pristoceutophilu. land, and Chile (Anonymous, 1991, 1992). Pristoceutophilu The Ministry of , The present lack of adequate quarantine Pristoceutophilu British Columbia regulations for large-scale log importations Tropidischia xa We supplied information to the Ministry has intensified the efforts to bring in such Hemiptera: Heterop of Forestry, Province of British Columbia material. Only now are comprehensive reg- Enicocephalidae: on selected species of arthropods that are ulations under public review that will pro- Boreostolus am, Lygaeidae: likely to be of conservation value (Lattin vide some protection against the uninten- Plinthisus longi and Moldenke, 1992) (Table 3). This was tional introduction of insects and pathogens. Thylochromus in response to a request for information that The risks are very high. This has been doc- Miridae: could be considered in decisions about the umented extensively for the Siberian logs Phytocoris nobi types of forests for inclusion in a series of (Anon., 1991), less so for New Zealand logs Pithanus maerl old-growth reserves. Naturally, field work (Anon., 1992) and is presently underway for Polymerus cast. will be required to even determine the pres- Chilean log importation. While this is not Scutelleridae: Vanduzeeina b. ence of such species on the proposed reserve. directly related to conservation, one has only Tingidae: This request is similar to others which we to look at Hawaii to see what impact non- Acalypta Mari have received to extrapolate our site-spe- indigenous species have had upon the local Acalypta saunc. cific information to other localities. biota of all types. One should examine, too, Coleoptera the forests of northeastern North America Conservation of Pacific Yew Carabidae: to see the damage that has occurred from Cychrus tuberc ( brevifolia) only a few diseases and insects, a forest Metrius contra I am completing a monograph on the changed forever. In spite of statements to Promecognath. arthropods associated with Taxus through- the contrary, the large-scale importation of Zacotus matth out the world, including the Pacific Yew, raw logs without absolute certainty of pest- Cicindelidae: Taxus brevifolia. Omus dejeani The great interest in Taxus free logs poses an enormous risk to the for- Curculionidae: species as a source of taxol, a cancer drug, ests of the west and to the native arthropods Lobosoma hor has vaulted this obscure understory tree into living in them. international prominence. Virtually noth- For other details si ing was known about arthropods on any THE FUTURE species of Taxus except on those species Many activities will influence the suc- 1992; Pimentel widely used as ornamentals, chiefly Taxus cessful conservation of arthropods in old- public land. baccata and T. cuspidata. Conservation growth forests, not the least of which will Effective arthi efforts for Taxus brevifolia naturally will be the achievement of a balance between growth forest wi benefit the few associated arthropods found forest harvest and conservation. This bal- of (1) the faun on it. The threat to the tree, and perhaps to ance is a highly politicized topic that became throughout the its arthropod fauna, is likely to be greatest an issue in the 1992 presidential campaign. ments, (4) ecolc from non-indigenous arthropod species A "forest summit" is planned for 1993 to the key taxa, (5 moving into natural stands from cultivars. try to resolve the conflicts. A major goal is and dynamics c The establishment of yew plantations for to agree upon a level of sustainable harvest. of different disc the production of foliage and stems for taxol Such action will create the template for any special habitats production may create populations of effort in forest arthropod conservation (7) appropriate arthropods able to move out into undis- (Southwood, 1977; New, 1984; Wilcove et monitoring pro turbed forested ecosystems as well. Thus, al., 1986; Warren and Key, 1991; Peterken, systematic know OLD-GROWTH FOREST ARTHROPOD CONSERVATION 585 tations should be TABLE 3. Examples of old-growth arthropods of conservation value.

Organism References Diplopoda Xestodermidae: ability of logs for Harpaphe haydeniana haydeniana (Wood) eased the interest n other countries. Insecta .port of raw logs Orthoptera Acrididae: Over the past two Boonacris alticola Rehn and Randall 1, 2, 3 tense interest in Gryllacrididae: Siberia, New Zea- Pristoceutophilus celatus (Scudder) 1, 2, 3 ous, 1991, 1992). Pristoceutophilus cercalis Caudell 1, 2, 3 quate quarantine Pristoceutophilus sargentae Gurney 1, 2, 3 log importations Tropidischia xanthostoma (Scudder) I, 2, 3 to bring in such Hemiptera: Heteroptera mprehensive reg- Enicocephalidae: iew that will pro- Boreostolus americanus Wygodzinsky and Stys 1, 2 inst the uninten- Lygaeidae: Plinthisus longisetosus Barber 1, 2 As and pathogens. Thylochromus nitidulus Barber 1, 2 his has been doc- Miridae: the SIberian logs Phytocoris nobilis Stonedahl 1, 2 New Zealand logs Pithanus maerkeli (Herrich-Schaeffer) 1, 2 ntly underway for Polymerus castilleja Schwartz 1, 2 Scutelleridae: While this is not 1, 2 ttion, one has only Vanduzeeina borealis californica Van Duzee Tingidae: what impact non- Acalypta lillianus Torre-Bueno 1, 2 lad upon the local Acalypta saundersi (Downes) 1, 2 uld examine, too, Coleoptera n North America Carabidae: as occurred from Cychrus tuberculatus Harris 1, 2 insects, a forest Metrius contract us Eschscholtz 1, 2 of statements to Promecognathus laevissimus DeJean 1, 2 ile importation of Zacotus matthewsi LeConte 1, 2 Cicindelidae: certainty of pest- Omus dejeani Reiche 1, 2 )us risk to the for- Curculionidae: native arthropods Lobosoma horridum Mannerheim 1, 2 For other details see references, viz., 1: Parsons et al.. 1991; 2: Lattin and Moldenke, 1992; 3: Lightfoot, 1986.

Ace the suc- 1992; Pimentel et al., 1992), especially on ate taxa in order to allow intelligent deci- i triropods in old-. public land. sions to be made (Wilson, 1985, 1989). ..ast of which will Effective arthropod conservation in old- balance between growth forest will require better knowledge ACKNOWLEDGMENTS rvation. This bal- of (1) the fauna, (2) species distribution My thanks to the many people who con- topic that became throughout the region, (3) habitat require- tributed to our knowledge of the arthropods dential campaign. ments, (4) ecological functions of at least of old-growth forests. What we know today finned for 1993 to the key taxa, (5) the community structure reflects their collected efforts. Thanks are s. A major goal is and dynamics of the fauna, (6) the impact due to the members of the H. J. Andrews istainable harvest. of different disturbance regimes upon the Research Group who educated me in the e template for any special habitats and the associated fauna, complexities of forest ecosystems. J. F. od conservation (7) appropriate and adequate sampling and Franklin, University of , has 1984; Wilcove et monitoring protocols, (8) and an adequate been a constant source of knowledge, inspi- y, 1991; Peterken, systematic knowledge base of the appropri- ration, and encouragement. D. A. Crossley, 586 JOHN D. LATTIN

Jr., University of , encouraged me Washington, Seattle. Coniferous Forest Biome lin (Tech. eds.). 198 in the early days when the tasks seemed Bull. 6. A story of fallen trees Franklin, J. F. and C. T. Dyrness. 1971. A checklist Pacif. Northwest For impossible. Thanks to Gary L. Parsons, of vascular plants on the H. J. Andrews Experi- Rep. PNW-GTR-22 Gerassimos Cassis, Andrew R. Moldenke, mental Forest, western Oregon. U.S. Dep. Agric.. Maser. C. and J. M. Trai Norman H. Anderson, Jeffrey C. Miller, Paul For. Serv., Pacif. Northwest For. Range Exp. Stn.. seen and unseen wor Hammond, D. Schowalter, Bon- PNW-138. Agric., For. Serv.. I Franklin, J. F. and C. T. Dyrness. 1973. Natural veg- Exp. Stn.. Gen. Ted nie Hall, Martha Brookes, and Frederick J. etation of Oregon and Washington. U.S. Dep. McIver, J. D., A. R. M Swanson for their efforts to see the anno- Agric., For. Serv., Pacif. Northwest For. Range 1990. Litter spiders tated checklist of the H. J. Andrews Exper- Exp. Stn., Gen. Tech. Rep., PNW-8. after clearcutting in imental Forest through to completion. Furniss, R. L. and V. M. Carolin. 1977. Western Northwest Environ. Andrew R. Moldenke deserves my special forest insects. U.S. Dep. Agric., For. Serv., Misc. Mispagel, M. E. and S. Publ. 1339. associated with van thanks for his extensive efforts to clarify the Goldsmith, F. B. 1991. Monitoring for conservation from foliar, ground. role of arthropods in forested ecosystems. and ecology. Chapman and Hall, London. ington, Seattle. Con Our many discussions over the years have Harmon, M. E. 1992. Long-term experiments on log Moldenke, A. R. 1990 provided me with inspiration and knowl- decomposition at the H. J. Andrews Experimental sand little legs. Wir Forest. U.S. Dep. Agric., For. Serv., Pacif. North- Moldenke. A. R. and E edge on many topics. My thanks to Anne west For. Range Exp. Stn., Gen. Tech. Rep.. PNW- braces of the H. J. Christie, Kerr Library, for help with critical GTR-280. western Cascade M( literature and to Deanna Watkins for help Harmon, M. E., W. K. Ferrell, and J. F. Franklin. batid mites (Acari in the preparation of the manuscript. The 1990. Effects on carbon storage of conversion of Agric., For. Serv., valuable comments from two anonymous old-growth forests to young forests. Science 247: Exp. Stn., Gen. Te( 699-702. Moldenke, A. R., B. L. reviewers are gratefully acknowledged. Harmon, M. E., J. F. Franklin, F. J. Swanson. P. Sol- E. Griswold. 1987 Support for this work from NSF grants lins, S. V. Gregory, J. D. Lattin, N. H. Anderson. of Douglas-fir and DEB-80-12122, BSR-85-14325, BSR-85- S. P. Cline, N. G. Aumen, J. R. Sedell, G. W. Northwest. U.S. Ii 16590, BSR-87-17434, and BSR-90-11663, Lienkaemper, K. Cromack, Jr., and K. W. Cum- Northwest For. Ra is gratefully acknowledged. mins. 1986. Ecology of coarse woody debris in PNW-GTR 207. temperate ecosystems. Adv. Ecol. 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