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CProceedingsonservation of relevan the Hawaiiance of inse Entomologicalct monitoring S ociety (2013) 45:149–166 149

The Importance of Monitoring to Conservation Actions in Hawaii

Matthew J. Medeiros1, 2*, Jesse A. Eiben3, William P. Haines4, Raina L. Kaholoaa5, Cynthia B.A. King6, Paul D. Krushelnycky4, Karl N. Magnacca7, Daniel Rubinoff4, Forest Starr8, and Kim Starr8 1Urban School of San Francisco, San Francisco, CA 94117. 2University of California, Department of Integrative Biology, Berkeley, CA 94720. 3University of Hawaii at Hilo, College of Agriculture, Forestry and Natural Resource Management, Hilo, HI 96720. 4University of Hawaii at Manoa, Dept. of Plant and Environmental Protection Sciences, Honolulu, HI 96822. 5Haleakala National Park, Makawao, HI 96768. 6Hawaii Division of Forestry and Wildlife Native Invertebrate Program, 1151 Punchbowl Street, Room 325, Honolulu, Hawaii 96813. 7Oahu Army Natural Resources Program, PCSU, Schofield Barracks, HI 96857. 8Starr Environmental, Makawao, HI 96768. *Author to whom correspondence should be addressed: [email protected]

Abstract. Endemic insect species make up the overwhelming majority of Hawaii’s native fauna, and play many important ecological roles. Despite this, receive low levels of conservation funding, likely due to their small size, fluctuating popula- tion sizes, and lack of baseline data necessary to determine if they are threatened with extinction. To determine which insects are at risk, how insect populations fluctuate in natural areas, and which management actions are most beneficial to Hawaiian ecosystems, we propose that insects be monitored whenever possible. Insect monitoring should be broad, generating community-based metrics such as species richness, rather that focusing on individual species. Resultant data should be entered into a stable, central database. Rather than individual insect species being the explicit target of conservation, we emphasize that measures of insect diversity can provide an assessment of restoration efforts, and serve as a metric for prioritizing areas for conservation. We provide lists of additional recommenda- tions for land managers and research entomologists who wish to assist with insect conservation efforts.

Key words: database, endemic species, indicator species, invasive species, moni- toring, population trends

I. Hawaiian insect diversity and terrestrial biodiversity. In terms of sheer the importance of insect number of species, insects are the major conservation components of biodiversity in virtually Most biologists would agree that a any terrestrial ecosystem, and this is es- primary global goal of conservation is pecially true in Hawaii, where the current the protection of biodiversity, whether it native terrestrial vertebrate fauna consists is measured in terms of species (Myers et only of birds and a single species of bat. al. 2000), evolutionary lineages (Williams At last count, Hawaii had nearly 5500 et al. 1991), or ecological diversity (Soule described native insect species, nearly all et al. 2003). By any of these measures, in- of these endemic (Nishida 2002), and this sects account for the majority of Hawaiian number grows steadily as new species are 150 Medeiros et al. described. In fact, insects comprise 53% of tion rates as high as, or higher than other all described endemic life forms in Hawaii groups (McKinney 1999). Moreover, it is and 70% of all endemic , includ- possible that the continued decline and ex- ing both terrestrial and marine organisms tinction of plants and birds that are already (Figure 1; Eldredge 2000). Invertebrates favored with conservation resources (e.g., as a whole, including other and the endemic birds poouli, Melamprosops mollusks, account for 82% of all endemic phaeosoma; alala, Corvus hawaiiensis; life forms and 97% of endemic animals palila, Loxioides bailleui) may be due (Eldredge 2000). to the disintegration of ecosystems more Because insects are such an important broadly, and the decline of native insects, component of Hawaiian ecosystems in specifically. Thus, ignoring the plight of terms of species richness, it stands to native insects may contribute to the disap- reason that they fill many ecological pearance of those more ‘charismatic’ ele- niches and provide important ecological ments of biodiversity. Ultimately, focusing services. Although little is known about nearly all conservation resources on plants specific plant-pollinator relationships in and birds may not serve the long term Hawaii, insects are certainly the primary interests of those taxa, if the insects on pollinators of native plants. Based on floral which they directly or indirectly depend morphology, 67% of endemic flowering are left to disappear. plants are thought to be insect pollinated, compared to only 19% bird pollinated, II. Challenges to insect and 14% wind-pollinated (Sakai et al. conservation 2002). Insects are also important parts Despite insects being such a crucial of food webs, being the primary native component of Hawaiian biodiversity, ver- consumers of plants (especially in the tebrates and plants continue to receive the absence of herbivorous mammals), and lion’s share of conservation attention and important decomposers of organic matter. funding. To make matters worse for insect Insects are critical sources of protein for conservation efforts, Hawaii’s share of all native forest birds, especially when nation-wide conservation funding is vastly nesting and raising young (Mountain- underrepresented (Leonard 2008). Lack of spring 1987, Ralph and Fancy 1994), and funds for insect conservation is likely due the relationships between birds and their to a combination of factors that complicate insect prey can be highly specific (Banko such efforts, including practical problems et al. 2002). The Hawaiian hoary bat or faced by land managers, as well as nega- ōpeapea (Lasiurus cinereus semotus), an tive or indifferent public attitudes towards endangered species, feeds exclusively on insects. In general, the United States lags insects and other arthropods (Whitaker in its efforts to conserve invertebrates, and Tomich 1983, Jacobs 1999, Fullard with disproportionately poor representa- 2001). Of course, insects also serve as food tion of arthropods on the endangered spe- for each other, with extremely complex cies list (one per 16 vertebrates relationships among native herbivorous and plants). Hawaii, which lacks most of insects and their predators and parasitoids the vertebrates that garner the attention (Henneman and Memmott 2001). of continental conservationists but has Yet, this high diversity among insects a much higher percentage of endemic is not indicative of the security of their insects, might be expected to do better persistence, and insect species in Hawaii than the national average. Regrettably, have almost certainly suffered extinc- Hawaii is doing worse: a lower percentage Conservation relevance of insect monitoring 151

Figure 1. Insect species account for 70% of Hawaii’s total number of endemic species. of the Hawaiian arthropod fauna is listed or spend all or most of their lives under as endangered than at the national level rocks, high in the tree canopy, or buried (one arthropod for every 19 vertebrates in or leaf litter. This means that as- and plants), and current practice largely sessments of population status or extinc- ignores the conservation of insects. For tion of individual species require a highly example, the Kauai green sphinx (Tinos- focused search effort. Additionally, the toma smaragditis) has been collected less adult life stages of many insects are able than 20 times since its discovery. Despite to fly and are therefore highly dispersive, very little being known about its life his- making it extremely difficult to follow tory, no studies are currently being funded individuals over time. Tracking insects to learn more about it, even though this using GPS or radio technology is currently moth is the sole species of an endemic impossible for all but the largest insects, genus. Only a few Hawaiian insects are although marking with paints or tags is now federally listed, and these represent feasible for some species (see McGeoch et a small fraction of the species at risk and al. 2011 for an introduction to the broader an insignificant proportion of Hawaii’s literature regarding insect monitoring). overall insect diversity. Compared to vertebrates and many Why does this lack of emphasis on in- plants, insects usually have relatively short sect conservation exist in the first place? lifespans, rapid generation times and often Perhaps the most obvious difficulty re- high reproductive output, meaning that garding insect conservation is that most populations can fluctuate greatly, both insects are very small and cryptic, often temporally and spatially. Some native noticed only after careful observation of insects exhibit “boom or bust” population an ecosystem. Many insects are nocturnal, dynamics, persisting at unnoticeable levels 152 Medeiros et al. most of the time, but then experiencing of its documented history—which includes occasional population explosions when the vast majority of Hawaiian insects, par- they are common, or even considered ticularly in highly-diverse radiations—it pests. For most species, the underlying is especially difficult to ascertain when causes of population fluctuations are not species are declining or under threat, since understood, likely being dependent on they may not be detected for years or even interactions between climate, resources, decades at a time. and mortality factors such as predators Given the challenges and limitations and pathogens, and not always following outlined above, how should researchers, predictable seasonal patterns. Because of land managers, administrators and others these short-term fluctuations, it can be dif- interested parties proceed to make insects ficult to detect long-term trends in insect a more integral part of conservation in populations, even when they can be ac- Hawaii? We offer our perspectives on curately sampled. To account for seasonal how best to approach this question in the variation, it is usually necessary to sample following sections. multiple times per year, sometimes over several years. This can quickly become a III. How and why should we daunting task when dealing with multiple monitor insects? species and multiple locations. Thus, As mentioned above, knowledge of how these methods are essentially abandoned to conserve insects often suffers from in practice, despite the fact that such data a dearth of information. This applies to are necessary to understand conservation understanding both the status of species or needs and large scale population patterns. populations and, to a somewhat lesser ex- Assessing the long-term viability of tent, how to mitigate threats to them. Some insect species can be difficult because of threats to insect species and communities this lack of historical baseline informa- are fairly well characterized (such as tion on distributions and abundances. For invasive social , see below), most native Hawaiian birds and plants, and the solutions to these problems are scientists have at least some records of theoretically straightforward, if often dif- where species formerly occurred, and ficult to implement. Other forces degrade how common they once were. In contrast, Hawaiian ecosystems more broadly and historical information regarding insect often interact with one another, and these distributions is far from complete, and likely exert impacts on insects that are for most species, is spotty or completely complex, are challenging to measure, and missing. Some insects that are currently as a result are currently poorly supported rare or localized may have always been with empirical data. This also means that rare, while for others, rarity may be a the best ways to alleviate these impacts symptom of decline due to novel threats. for insects may not always be clear. We Many insects that were among the most therefore suggest that a critical need at common in the 1890s, and some even more this juncture is an increased investment recently, have become extinct or nearly in insect monitoring, which will serve so. Three examples include the carabid several inter-related goals. beetle Blackburnia tantalus (Liebherr and First, increased monitoring will begin Polhemus 1997), the facilis to allow more accurate assessments of (Daly and Magnacca 2003), and the wasp population or species trends for at least Euodynerus nigripennis (Williams 1927). a subset of the native fauna, which are For any species that has been rare for most currently almost entirely lacking. Second, Conservation relevance of insect monitoring 153 monitoring insect communities will pro- a single group will consistently indicate vide much additional information to land trends for all or possibly even most other managers and state officials about overall groups. Similarly, one group may be es- patterns and trends in diversity within pecially sensitive to one type of stressor, protected natural areas. We reiterate that and thus be a good “indicator” for that insects and their relatives comprise the impact, but another group may be a better majority of the biodiversity that is the indicator for another impact. For example, object of this protection. A third benefit Krushelnycky and Gillespie (2010) found is that the monitoring of insect communi- that native and beetles were ties, when conducted in conjunction with among the most sensitive taxonomic management actions, can begin to clarify groups to invasive ants in Hawaii, that whether commonly used management other groups were much more variable strategies actually do provide significant in their responses, and that trophic group benefit to insects (as is often assumed or and population density were generally at least hoped), and ideally whether some more effective for predicting sensitivity are more effective than others. For a more to ants than was taxonomic identity. In thorough discussion of the advantages comparison, preliminary assessments of using arthropod diversity to inform suggest that Hawaiian Orthoptera and conservation practices, see the arguments larval Lepidoptera may be particularly of Kremen et al. (1993) and Oliver and sensitive to predation by rodents, but also Beattie (1996). that the vulnerability of taxa may vary Of course, the effectiveness of moni- between sites and be context-dependent toring efforts is dependent on the con- (P. Krushelnycky, unpublished data). Non- tinuation of the sort of general collecting native plant invasions may produce yet a and taxonomic research that has been a different pattern for insect communities. mainstay of entomological work. This Therefore, it will often be difficult to know research provides critical information that which indicators to select until more com- gives conservation assessments meaning prehensive sampling demonstrates which and validity, and at the same time can groups respond to a particular stressor provide an alternate means to evaluate and which do not. Practically speaking, species status. Finally, an effective way this means that broad sampling is the best to aggregate this information is needed: current approach, until such datasets can a centralized invertebrate database is be used to determine whether a smaller critical to synthesizing abundance and subset of taxonomic or functional groups distribution information generated from can be used as indicators for the health of all types of collecting and monitoring ef- the larger insect community (Basset et al. forts (McGeoch et al. 2011). We elaborate 2004). on these points below. When assessing the impacts of threats A. Using a broad approach to insect or management actions, community- monitoring. The use of specific indicator wide metrics, such as endemic species groups presents an appealing way to deal richness, diversity or proportion of the with the complexity of monitoring insect community that is native (by species or communities (McGeoch 1998, Niemi and individuals) should be evaluated, even McDonald 2004, Gerlach et al. 2013). if trends within specific taxonomic or However, different insect groups often functional groups are also investigated. respond differently to different factors, Community-wide metrics are important to so it is not likely that population trends in use because measures that span multiple 154 Medeiros et al. insect groups will be much less sensitive to but also to make a species-specific rec- the idiosyncratic fluctuations of particular ommendation to conserve it. Some of the species or populations. Changes in these well-known Aeolian-dependent species gross-level, synthetic metrics over time such as the wekiu bug ( wekiuicola) between different inventory or sampling (Eiben and Rubinoff 2010) and Haleakala efforts may be insightful even when dif- grasshopper moth (Thyrocopa apatela) ferent methods were used, although we (Medeiros 2009) are specific examples; strongly suggest being as consistent as other well-known species with a high level possible in sampling methodology if a of cultural or ecological importance may known goal is to make such comparisons. benefit from species-specific monitoring For example, an inventory of arthropods as well, such as yellow-faced in the at Haleakala National Park (Krushelnycky genus Hylaeus. In each of these cases a et al. 2007) conducted roughly 25 years significant investment into the species’ after an earlier one (Beardsley 1980) life history is or was necessary. In ad- allowed for some limited comparisons dition, research on specific taxonomic regarding overall patterns of adventive groups has provided clear evidence of the species establishment, but differences in wider impacts of native habitat conversion: the goals and methods between the two Leblanc et al. (2013) found that a suite of surveys hampered detailed assessments Hawaiian Drosophilidae was strongly tied of faunal changes over the intervening to native forest stands. period. Assessments of native arthropod In sum, we suggest that land managers diversity require that insects be identified partner with entomologists to conduct to low taxonomic levels, necessitating both general arthropod monitoring in locations expertise and time investment. However, and situations of interest, especially before because the insect fauna of Hawaii is and after conducting particular manage- disharmonic, with many groups absent, ment actions. Even when insects are not species-level identification is considered a specific target of management actions, more feasible than continental tropical they can serve as a valuable measure of ecosystems. Additionally, many groups of the effectiveness of habitat protection. insects can be classified as native or non- By tracking increases or decreases in the native after identification to genus, family, proportion of native insect diversity or or even order, and it may be possible to abundance, we can use insects as a kind use morphospecies to quantify diversity of “conservation currency.” Initial moni- within some groups. toring will establish species lists and/or That said, while we generally advo- community composition at a given local- cate that researchers and land managers ity, with repeated observations providing attempt to sample insects as broadly as important information about how specific possible, specialized monitoring focusing management actions affect insect conser- on single species or groups of interest may vation and recovery. Whenever possible, sometimes be needed or valuable (see Mc- it is important to simultaneously monitor Geoch et al. 2011 for a list of examples of at a comparable site where management species of interest from South Africa). In actions are not implemented, to tease apart some cases, when land managers request the effect of climate and other factors conservation recommendations from en- unrelated to management. tomologists, enough may be known about Our goal here is not to provide a list a specific insect to confirm not only that of specific sites and methods for this the insect is threatened with extinction, monitoring, because the decision to un- Conservation relevance of insect monitoring 155 dertake insect monitoring will depend tifications are still incomplete due to heavily on available funding, manpower, the unavailability of specialists in these and other logistical and situation-specific groups, or to an inadequate present state considerations. Similarly, appropriate of taxonomic knowledge concerning methods can also be highly site- and goal- the groups. specific. However, several examples that This passage highlights the importance illustrate our aim include efforts by the of current taxonomic work, as well as Oahu Army Natural Resources Program identification of specimens by experts. to monitor insect community response to In addition, the compilation of infor- intensive rodent trapping in the Waianae mation generated in taxonomic studies Mountains, an effort by Pacific Rim Con- and other types of “general collecting” servation and the Hawaii Department of can in some cases yield insight into spe- Land and Natural Resources (DLNR) to cies trends and conservation status. For monitor insect response to the construc- example, Vorsino et al. (2013) found a tion of a predator-proof fence at Kaena substantial distributional shift and range Point Natural Area Reserve, and an effort reduction over time in Omiodes continu- by DLNR, the U.S. Geological Survey and atalis, by comparing data from histori- the Maui Forest Bird Recovery Program cal collections with those from modern to survey insect diversity and abundance surveys. The consistent entry of this type to ascertain food resources for endangered of collecting information, as well as in- forest birds on East Maui. Given sufficient formation from broader inventories and progress and momentum in this direction, monitoring efforts, into a central statewide a working group could be established database will provide a vast amount of to provide more specific guidance and information that would be difficult to perhaps make recommendations for a compare and synthesize otherwise. For network of monitoring sites. instance, Howarth and Preston (2007) B. Establishing a statewide inverte- recently documented insect species found brate database. The continued descrip- on lands surrounding Kahului airport on tion of native Hawaiian insect diversity Maui. While this information is extremely is critical to its conservation, especially useful for monitoring long-term changes considering its high level of endemism. All in insect communities, it is currently not conservation assessments and decisions easy to conduct a quick search of these are ultimately based on our understand- data. To aid scientists, land managers, ing of the native biota. As a result of this and environmental planners gathering constraint, previous attempts at compil- information on the changing distribu- ing complete lists of species for a given tions of taxonomic groups, a new online time and place within Hawaii have been State Invertebrate Database (SID) is being made with varying degrees of success. developed. The database is being gener- For example, Beardsley (1980) published ated by the State of Hawaii Department a catalog of all insect species, native and of Land and Natural Resources, Division non-native, found in the crater district of of Forestry and Wildlife, and will be a Haleakala National Park. There, he wrote specimen-level database that includes (p. 1), geographical (GIS) locality data whenever The Catalog is still relatively incom- available. An ultimate goal of the database plete. Specimens belonging to certain is to catalogue all museum specimens taxonomic groups have not yet been from collections made in Hawaii, and identified. In other groups, many iden- provide a central repository into which 156 Medeiros et al. future specimen and collection data can sitoid, predator, scarcity of host plant, or be integrated. The SID will allow users complex interactions between all of these. to generate distribution maps for given Second, even when the primary threat to species, species lists for given localities, an insect species is known, if that threat and answer broad questions which to is highly specific and difficult to alleviate, date have been difficult to address. For doing so may not be the wisest use of re- example, researchers will be able to ana- sources, given the large number of species lyze changes in habitat use by species over that would be ignored in the process. The time, land managers will be able to better general action of keeping ecosystems as prioritize locations for management and intact as possible maximizes management conservation efforts, and groups planning efficiency by benefiting an entire suite for new protected areas will be able to of species (Asquith 1995; Gagné 1982; determine whether rare species have been Howarth and Gagné 2012). documented from a particular location of When a species-specific approach is concern. warranted or required (e.g. under the Endangered Species Act), it is best to IV: Recommendations for target species whose protection will land managers benefit many other species or even entire Generally, species tend to have high ecosystems. Drosophila flies are a good extinction rates in fragmented habitats example: They mainly breed in subdomi- (Hanski 1999, Rybicki and Hanski 2013). nant trees and require moderately dense Therefore, as with many other organisms, shade, so protecting their hosts requires the best strategy is often to protect large maintaining a native canopy and eliminat- tracts of land, rather than many small, ing factors that hinder plant reproduction, widely spaced patches of high-value habi- such as pigs and rats. Management actions tat (Samways 2007). That said, minimum that benefit the flies therefore also benefit a habitat sizes for insects are often smaller number of other species, including plants. than for vertebrates, so efforts to protect How should ecosystems in Hawaii even small areas may have a significant be broadly protected? As stated above, payoff in terms of insect conservation. there is little robust evidence that com- Some rare and elusive insects can be mon management practices are effective conserved effectively in small ranges that for conserving insects in Hawaii, though offer some natural protection, perhaps many of us assume that they are. Until partly due to remoteness and/or inacces- more information becomes available, we sibility, though some invasive species still suggest following recommendations that pose problems in such areas (New 2009). are based on our understanding of impor- With the caveat that there may some- tant threats. For example, Beardsley (1980) times be exceptions, we agree with listed four management actions that he Samways (2007) that the most effective considered critical to preserving endemic conservation strategy is a generalized ap- insect biodiversity, particularly at high proach, protecting habitats from universal elevations. These four actions are no less threats, rather than attempting to protect relevant today than they were over thirty insects on a case-by-case, species-by-spe- years ago when first suggested. We highly cies basis. This is largely for two reasons: recommend that management agencies First, the most important limiting factor direct available resources toward these for any given insect species is not often actions, when relevant to the ecosystem known with certainty, whether it be a para- in question. To paraphrase and update Conservation relevance of insect monitoring 157 Beardsley’s (1980) recommendations, impacts of ants are complicated and not these four actions are: always easy to document, it is nonetheless 1. Ungulate control. Beardsley wrote clear that ants pose a serious threat to that feral ungulates are a threat to insects many endemic insects (Cole et al. 1992, and should be excluded, since they con- Krushelnycky and Gillespie 2008, 2010). tribute to the decline of plant communities Numerous ant species have invaded many (though alien weed control within fenced habitats in Hawaii, and some continue to areas is also important; see Cabin et al. spread (e.g., Hartley et al. 2010). Eradica- 2000). Host plant specialists as well as tion or control of ants using baits and toxi- the consumers that feed upon them are cants would benefit a wide range of species therefore harmed by ungulate rooting and (Krushelnycky et al. 2005), although the browsing. Ungulate damage in Hawaii non-target effects of control methods must has been well documented by a number be considered, especially if treatments are of papers; the literature has been synthe- conducted area-wide on a regular basis. sized in two position papers by the Hawaii Unfortunately, ant eradication is usually Conservation Alliance (2005; 2007). We very difficult, and success depends on add that rodent control is important as various factors including the identity of well, since rodents are direct predators of the target species and most importantly arthropods as well as major seed predators the size of the infestation. There are now of plants they depend on. many examples of relatively small scale Ungulate exclusion and control tech- eradications globally (Hoffmann et al. niques have been mastered in Hawaii and 2011), including the eradication of Phei- have become standard practice for most dole megacephala from an Oahu offshore natural areas. There are many examples islet (Plentovich et al. 2011) and the likely of successful ungulate control and eradica- eradication of Wasmannia auropunctata tion (Cole et al. 2012; Tunison et al. 1995). on Maui (Vanderwoude et al. 2010). How- If the island is large, eradication will ever, we know of only a single instance likely only be local and fences must be where more than one invasive ant species maintained forever. Smaller islands, such was simultaneously eradicated from a as Kahoolawe, can have all the ungulates managed area (Hoffmann and O’Connor removed, requiring no additional costs for 2004), and in Hawaii multiple problematic ungulate control. ant species often co-occur, which can Rodent control is much harder than present greater management challenges. ungulate control and is usually done over For example, the eradication of bigheaded a limited area. There are many examples ants (P. megacephala) from Mokuauia of successful rodent eradication on small Islet was followed by a dramatic increase islands and atolls (Howald et al. 2007). in the number of yellow crazy ants, Ano- However, control of rodents on large plolepis gracilipes (Plentovich et al. 2011). islands will likely be restricted to small 3. Wasp control. Invasive yellowjack- high value locations, through the use ets (Vespula pensylvanica) have played of persistent trapping or installation of a major role in diminishing native insect predator-proof fences. populations through direct predation 2. Ant control. Entomologists since (Gambino 1992, Wilson et al. 2009), Perkins (1913) have noted the devastation though as Howarth and Gagné (2012) that ants have caused to the Hawaiian point out, quantifying the extent of wasp insect biota. Hawaii is believed to have impacts is difficult. Regardless, yellow- no native ant species, and although the jackets are major pests of both humans and 158 Medeiros et al. other animals and their nests should be trends as native species respond to the destroyed when possible. Although wasp invasives. control is resource intensive and must be To prevent invasive species from enter- done on a continuous basis, techniques ing native habitats, transportation of ma- for successful wasp control are known, terials into these areas should be closely and local control can also benefit humans monitored. For example, insects such as that frequent those areas. Yellowjackets ants can colonize construction materials have been documented to displace native while they are stored in baseyards, and en- pollinators (Wilson and Holway 2010), tire colonies can then be transported into and management of yellowjackets has natural areas. Construction equipment been shown to increase seed set of native and materials that have previously been plants (Hanna et al. 2013). in an area with invasive species should be 4. Ask visitors to stay on trails. This is power-washed and closely inspected for the most habitat-specific recommendation seeds, dirt, and insects before being al- in Beardsley’s (1980) report, as the alpine lowed into conservation areas. Fence con- cinder desert is such a small area, and a struction is an important management tool place where many arthropods take shelter used to protect native habitats in Hawaii under rocks. However, trampling any area from ungulate damage, and predator-proof in Hawaii still dominated by native plants fences may become a more specific tool will have detrimental impacts to the habi- for insect conservation (Watts et al. 2008). tat, and this is worth reminding people. However, fence lines also frequently serve as corridors for the spread of non-native Howarth and Gagné (2012) also provide species. Equipment sanitation and post- an updated list of general management construction monitoring are integral to actions relevant to insect conservation in reducing the introduction and spread of Hawaii, though some of these actions are invasive species into conservation areas, geared more toward government agencies and these kinds of actions should be trig- rather than individual land managers or gered when construction is proposed and researchers. Their suggestions include written into contracts before construction calls to control invasive species, improve begins. Plants and associated planting quarantine measures for shipments to Ha- media used within natural areas should be waii, and improve the testing and review closely inspected before they are brought for potential biocontrol introductions. We into the field. recommend these actions as well. One of We recommend planting native plants the biggest threats to native Hawaiian spe- wherever practical. Many native insects cies is the introduction of invasive alien can only utilize native plants, and even if species, often insects themselves (see common native plant species are estab- McGeoch et al. 2011). When invasive spe- lished far away from natural areas, the cies are introduced into already shrinking potential resource opportunities increase native habitats, they may displace native for native insects (see Roets and Pryke plant species, directly prey upon native 2013). For example, the native tephritid fly insects, or otherwise degrade existing (Trupanea artemisiae) is only known from habitats. Resource managers must be the native hinahina (Artemisia australis aware of any new invasive species, and and A. mauiensis). When A. mauiensis regular monitoring of insect populations was planted at a number of sites many (see above) can help to detect these new miles from the known range of this fly, introductions and to track population the fly was able to find the plants and Conservation relevance of insect monitoring 159 successfully breed (Starr and Starr 2012). non-native plants in some circumstances This type of scenario is also believed to can still harbor a number of endemic in- have occurred on the once mostly barren sects. We are not suggesting that invasive island of Kahoolawe: As more host plants plants ever be included in restoration were planted on the island, native insect efforts, only pointing out that a knee-jerk species associated with these plants were reaction of rapidly removing all non-native discovered (Starr and Starr, personal ob- plants is not always the best approach. servations). In sum, our recommendations for land On the other hand, non-native plant con- managers are as follows: trol is not always immediately desirable. 1. Incorporate monitoring of insect com- Though many native Hawaiian insects are munities (including funding these ef- very host specific and can only utilize na- forts) into management plans, especially tive plants, other endemic insects are able when these plans include important new to utilize non-native plants. For example, management actions the native tephritid flyTrupanea crassipes 2. Ungulate and rat control has only been reared from the non-native 3. Ant control Spanish needles (Bidens pilosa). Many 4. Wasp control native Hylaeus bees heavily utilize the 5. Asking visitors to stay on trails non-native tree heliotrope (Tournefortia 6. Control and monitoring of other inva- argentea) (Magnacca and King 2013). sive species (often plants and insects) Assuming this non-native tree does not 7. Frequent cleaning and sanitation of display invasive tendencies, removing all construction and fence-building it from locations where there are native equipment bees would not be in the bees’ best interest 8. Close inspection of plants and soil that unless an equally attractive plant resource are to be brought into the field was immediately available. The native 9. Planting and restoring native vegetation Hawaiian grass leafroller moth (Omiodes wherever practical (with the caveat that continuatalis) utilizes many non-native stands of non-native plants should not grasses as a larval host, including the be destroyed without some consider- non-native kikuyugrass (Cenchrus clan- ation of their possible role in insect destinus). Perhaps the most conspicuous conservation) example is the endangered Blackburn’s 10. Encourage general collecting of in- sphinx moth (Manduca blackburni) that sects by research entomologists (see mostly uses the non-native tree tobacco below) (Nicotiana glauca), a common roadside weed, as its larval host plant. Some gen- We recognize that many of these rec- eralist native insects such as predators ommendations are practices that land and detritivores may rely on the structural managers are already doing, or attempting complexity provided by stands of mature to do, and managers may feel frustrated trees, regardless of whether they are na- with the current lack of novel methods of tive or non-native, so removal of large insect conservation. However, long-term non-native trees may not be advisable monitoring of insect communities is glar- until outplanted native trees have become ingly absent from most management plans. established. Managers should take into Although such monitoring can be chal- account potential deleterious effects on lenging, and requires significant financial native insects when weighing costs and investment, we feel strongly that one of the benefits of non-native plant control, as most important ways in which insects can 160 Medeiros et al. inform conservation is via the data they the research in the context of ongoing provide. This requires a shift in perspec- management interests. While time and tive on the part of both land managers resource constraints don’t always make and entomologists. In addition to asking, it practical for entomologists to gather “What can we do to conserve Hawaiian data relevant for conservation, there are insects?” we might ask, “What can insects certainly instances when useful data can tell us about how well we are conserving be generated relatively easily while con- Hawaiian biodiversity?” Rather than being ducting a larger research program. For the explicit target of conservation, insect example, many endemic Hawaiian birds diversity can provide a measure of success rely heavily on caterpillars as a source of restoration efforts, or serve as a metric of protein, and it is likely that the decline for prioritizing areas for conservation. Of in endemic caterpillars is a contributing course, our hope is that more data on how factor in the decline of these birds (see conservation practices affect insect com- Asquith 1995 for an overview of this munities will eventually inform us as to issue). Land managers might ask a field which are most effective for conserving lepidopterist doing a taxonomic revision particular insect groups, enabling us to to collect data on the abundance of cater- revisit the question, “What can we do to pillars observed during their fieldwork. conserve Hawaiian insects?” Depending on the biology of the target species, and the frequency at which sites V. Recommendations for are visited, the lepidopterist may be able research entomologists to generate a useful ecological dataset We suggest that communication be- on caterpillar diversity and abundance. tween resource managers and research Such information may not be the primary entomologists is a necessary prerequisite objective of the entomologist, but this type for effective and long-term insect conser- of communication between managers and vation in Hawaii. Information that can entomologists may lead to publications help managers prioritize activities or for- that would not otherwise have been pos- mulate management plans include insect sible. We do stress, however, that small inventory and distribution data (such as datasets or those that sample for only that to be compiled in the SID, and that a short period of time will be highly already available in various publications), subject to error due to sometimes large specific threats to native insect species natural fluctuations in population sizes of in these areas if known, and sometimes, insects, and such data must therefore be what can be done to alleviate these threats. interpreted with caution. Dialogue between researchers and manag- We recognize that land managers may ers can lead to insect conservation actions sometimes be frustrated when looking that benefit multiple species and resources. for relevant information in the academic Here, we outline several ways research literature, in particular when ecological entomologists can help land managers information such as distributions, host plan for conservation actions. plants, and rarity are buried in species We recommend that research ento- descriptions and the legends of phyloge- mologists, when practical to do so, “check netic trees. Research entomologists are in” with land managers before starting therefore encouraged to highlight relevant research projects, particularly those in- information for insect conservation, such volving new collections. This provides as maps, lists of species by localities, and an opportunity for managers to discuss host plant records, and ideally report it Conservation relevance of insect monitoring 161 directly to land managers. In addition, cades ago and very few collected recently. data-free statements in papers declaring, Clearly, this makes it difficult for experts for example, that a given insect group in any given group to track the changing “can be used as an indicator for overall distribution of endemic insects using ecosystem health,” are of limited utility. natural history museums. Publishing such statements without indi- To remedy these problems, we make cating how and why a particular group are two recommendations: First, although it biological indicators, and without making may not be realistic in the grant-driven specific management recommendations, is modern world to expect graduate students nearly meaningless to land managers who and researchers to have time to become are attempting to take real conservation the kinds of general entomologists more actions on a daily basis. When applicable, commonly seen in decades past, it is pos- researchers should report any information sible for students and researchers alike to to managers if they believe that an aspect participate in regular “bioblitzes.” These of current management practices may be are carefully organized and intense col- harming native insects, or if changes in lecting expeditions that focus on gathering management practices could significantly as many specimens as possible in a short benefit native insects. time, and identifying them as quickly as One major trend in contemporary possible. With a variety of specialists at- entomology is the decline of the “gen- tending a bioblitz together, there is excel- eral entomologist.” Starting in graduate lent opportunity to learn things outside of school, entomologists—like practitioners one’s own area of expertise. We encourage of other fields—tend to become more and the organization of additional bioblitzes more specialized as they progress toward and other detailed biological inventories a degree. For non-applied entomologists, (see Howarth and Preston 2007) in the keeping up with the exponential increase future. We recommend that land manage- of relevant scientific papers coupled with ment agencies suggest areas they would the joint pressures to publish and earn like to learn more about as possible sites grants in fields as diverse as phylogenet- for future bioblitzes, since such activities ics to biomechanics leads most students can provide at least a qualitative assess- to focus narrowly on one group. This ment of native arthropod diversity. ultimately leads to researchers who are Second, we recommend that when experts in one area but would be hard- practical to do so, researchers request on pressed to evaluate conservation concerns, research permit applications that they be much less identify other types of arthro- allowed to collect generally while in the pods, outside of their narrow study system. field, and that they collect, curate, and A related problem is that entomologists deposit as many specimens into museums typically only collect their target group as time permits (with the exception of of organisms during fieldwork. Reasons threatened and endangered, or otherwise include lack of ability to identify non- known rare species). This would help to target organisms, lack of time to curate alleviate the huge gaps in data currently them, and also reluctance on the part of suffered by existing natural history collec- land-management agencies to approve the tions and allow experts for given groups to collection of non-target specimens. The use recently collected museum material to result is fewer and fewer specimens in quickly learn about the status of species research collections, many of which have even when they don’t have the time or re- an abundance of specimens collected de- sources to conduct field work themselves. 162 Medeiros et al. Field researchers may not always have the laborative collecting efforts time or resources to curate and identify 4. When practical, ask for permission to specimens that are not directly related collect generally, and quickly deposit to an ongoing research project, but when specimens outside your area of exper- they are willing to undertake the effort of tise into natural history museums. properly preparing specimens, we believe it is in the best interests of land managers Lastly, we remind research entomolo- to allow it. Having specimens in natural gists who have the opportunity to do so history museums that can later be identi- that sharing fun, interesting, and impor- fied by experts would be preferable to the tant insights into insect natural history current dearth of recent specimens. with interpretive staff and even visitors Of course, this practice depends on can help give staff and visitors alike a cooperation from land managers and broader understanding and appreciation others who review and approve collect- of the insect world, ultimately leading to ing permits: We recommend that when increased support for protection of insect considering collecting permit applica- populations. Posters given to interpretive tions, resource management agencies centers, classroom visits, and even im- should reassess the current standard of promptu conversations while in the field, only allowing small numbers of “target” are all examples of ways we can positively specimens to be collected by a particular impact the future of insect conservation. researcher. Rather, whenever an entomolo- gist goes on a collecting trip, permitting Notes agencies could encourage them to col- This paper was conceived in large part lect, curate, and deposit specimens from by discussions that took place during a a wide variety of groups. We have listed symposium that took place at the 2012 this recommendation in the above section Hawaii Conservation Conference entitled (“Recommendations for land managers”). “They Are the 99%: Challenges and Opportunistic collecting by entomologists Successes of Conserving Invertebrates, is very unlikely to negatively impact in- Hawaii’s Smallest Majority.” Other than sect populations, given several aspects of the corresponding author (MM), authors insect biology (Morris 1987). are listed alphabetically. In sum, our recommendations for ento- mologist researchers are as follows: Acknowledgments 1. Check in with land managers before Collectively, we thank Frank Howarth, projects, when possible, to determine Betsy Gagné, Wayne Gagné, Rosemary whether it is feasible to generate datas- Gillespie, Sam Gon, Ken Kaneshiro, Bill ets that are of conservation utility Mull, Steve Montgomery, Jim Liebherr, 2. Refrain from making statements in Dan Polhemus, George Roderick, Fred publications that suggest conservation Stone, and Lorena “Tap” Wada, among utility of taxa without stating how this others, for being inspirational advocates is possible; rather, present land manag- for Hawaiian insect conservation. Robert ers with easy-to-interpret distribution Dudley provided logistical support for maps, hostplant records, and checklists MM. Michael Samways provided valuable for species you study, as well as any comments that improved the manuscript. practical recommendations or sugges- Funding was provided in part by the Na- tions you may have tional Science Foundation (NSF) award 3. Participate in bioblitzes and other col- #DEB-0918341, USDA Cooperative Conservation relevance of insect monitoring 163 State Research, Education and Extension Hawaiian forest. Biotropica 44: 463–471. (CSREES) projects HAW00942-H and Daly, H.V., and K.N. Magnacca. 2003. 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