Northeastern Naturalist 23 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 Exploring the Microwilderness of Boston Harbor Islands National Recreation Area: Terrestrial Invertebrate All Taxa Biodiversity Inventory Jessica J. Rykken1,* and Brian D. Farrell1 Abstract - Between 2005 and 2011, we conducted a terrestrial invertebrate All Taxa Biodiversity Inventory (ATBI) in Boston Harbor Islands National Recreation Area, in order to document as many arthropod and gastropod species as possible in the park, and to understand how species were distributed across habitats and islands. Professional scientists, students, and citizen scientists collected ~160,000 invertebrates on 19 islands and peninsulas in the park, using a variety of trapping and collecting methods. More than 76,000 of these specimens were curated, identified, and databased, resulting in a total of 1732 distinct species and morphospecies. Of these, 232 species (13.4%) were species not native to North America. The introduced species included several new US and North American records, including 2 potential pests: Hishimonus sellatus (Mulberry Leafhopper) and the click beetle Athous haemorrhoidalis. Among native species, we documented several new state records, which expanded known ranges considerably in a few cases. Statistical estimates of absolute species richness for several representative taxa indicated that less-diverse groups (e.g., millipedes) were sampled almost completely by our methods, but additional sampling is needed to thoroughly inventory more-diverse taxa (e.g., ground beetles). The invertebrate ATBI lays the groundwork for future monitoring of focal groups such as pollinators. Introduction The All Taxa Biodiversity Inventory (ATBI) concept, first conceived by Janzen and Hallwachs (1994), has as its ultimate goal the identification and cataloguing of all species occurring within the boundaries of a park or other natural area in a relatively short period of time. Typically, ATBI efforts have been undertaken in known hotspots of biodiversity such as the Dominican Republic (Farrell 2005) or Great Smoky Mountains National Park (Nichols and Langdon 2007). In 2005, we initiated the terrestrial invertebrate phase of an ATBI in Boston Harbor Islands National Recreation Area (NRA). Prior to the terrestrial invertebrate ATBI, faunal inventories in Boston Harbor Islands had focused primarily on birds (Paton et al. 2005) and other vertebrate animals (Trocki et al. 2007), while the microwilderness—as renowned and local entomologist, E.O. Wilson, refers to the immense realm of invertebrates—had received disproportionately little attention. A notable exception was a comprehensive Macrolepidoptera survey conducted by Mello (2005). Boston Harbor Islands NRA is not regarded as a biodiversity hotspot. The park comprises 30 small islands and 4 peninsulas lying within 20 km of downtown Boston, MA. A long history of use and colonization by both Native and European 1Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA, 02138. *Corresponding author - jrykken@oeb.harvard.edu. Manuscript Editor: Christopher M. Hecksher Boston Harbor Islands National Recreation Area: Overview of Recent Research 2018 Northeastern Naturalist 25(Special Issue 9):23–44 Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 24 Vol. 25, Special Issue 9 Americans has altered island landscapes dramatically over the last several hundred years (Richburg and Patterson 2005). Overall species diversity in this urban island park was expected to be relatively low due to its temperate bioregion, disturbed landscapes, small size, and island isolation; however, this combination of traits also served to make the primary goal of the Boston Harbor Islands ATBI somewhat more attainable. The historical legacy of centuries of trade in the harbor has likely influenced the diversity of non-native species on the islands. Thus, an urban ATBI in a national recreation area presented a novel framework for exploring regional biodiversity. In addition to documenting as many terrestrial arthropod and gastropod species as possible in the park, we wanted to understand how species were distributed across habitats and islands/peninsulas. By conducting a terrestrial invertebrate inventory on an urban island archipelago, we hoped to explore themes about island colonization on a small spatial scale (Rykken and Farrell 2013), species resilience to human disturbance, and the relative dominance of non-native species (see Rykken and Farrell 2018 [this issue]). The data we collected during the invertebrate ATBI covered an extremely diverse array of taxa, and the results we present here provide a brief summary of our overall findings. Our intention is that our work will suggest topics or particular questions that invite further investigation and focused research in Boston Harbor Islands NRA. We also intended that biodiversity information could be used to inform management decisions and actions in the park. In addition to knowledge about individual species of interest (e.g., rare, pest, or newly introduced species), an ATBI can provide information about ecological roles and relationships, and species distributions and habitat associations. These kinds of data might help park managers prioritize conservation measures for particular communities or habitats in the park. Species data from parks can also contribute to larger regional datasets, providing new information for species-distribution maps and for monitoring range-shifts over time. The selection of particular taxa (e.g., pollinators) as indicators of ecosystem integrity can provide parks with valuable tools for measuring the long-term effects of local management actions or large-scale environmental threats such as climate change. Field-site Description Within Boston Harbor Islands NRA (42°18'31''N, 70°57' 51''W), the islands and peninsulas sampled for invertebrates varied in size from 1.1 ha to 104.5 ha. Most of the islands in the harbor were formed from glacial deposits of till, known as drumlins, and a few are bedrock (Rosen and FitzGerald 2004). The primary vegetation communities on most islands include forest, woodland, maritime shrub, old field, and beach strand, and non-native woody and herbaceous plant species dominate many of these communities (Elliman 2005). Salt-marshes and brackish marshes occur on several of the islands, but fresh water is extremely scarce. The mainland surrounding the harbor comprises several towns, varying from somewhat less-developed landscapes in Hingham and Weymouth in the south of the harbor, to heavily urbanized landscapes in Boston and Winthrop, including international port facilities for shipping and air Northeastern Naturalist 25 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 transport. Over the past several centuries, all of the more accessible and sizeable islands have hosted human structures and activities (Kales 2007). Currently, almost all of the islands in the park are open to human visitors, but they vary greatly in the intensity of anthropogenic traffic and impacts. Methods We sampled terrestrial invertebrates on 19 islands and peninsulas between 2005 and 2010. Nine islands (Bumpkin, Calf, Grape, Great Brewster, Ragged, Snake, Spectacle, Thompson, Worlds End) received an intensive structured sampling regime that targeted all arthropods and gastropods for at least 1 full growing season (May to October); Ragged Island was sampled only August–October during the pilot season in 2005. Eight islands and a peninsula were sampled intensively for bees in 2010 (Bumpkin, Grape, Great Brewster, Langlee, Peddocks, Spectacle, Thompson, Webb, Worlds End). Seven additional islands were visited sporadically for opportunistic collecting, bioblitzes, and/or were included in student projects (Deer, Georges, Long, Lovells, Middle Brewster, Outer Brewster, Rainsford). We used a variety of traps and methods to sample different habitats. On islands with structured sampling, we stratified the sampling by dominant habitat type: woodland, shrubland, meadow, beach, marsh, or pond edge. Passive-sampling methods included pitfall traps, Malaise traps, bee bowls (small pan traps), and light traps. We also sampled actively by hand-searching, and using aerial and sweep nets, beating sheets, and aspirators. A detailed description of our sampling design, collecting methods, sample processing, insect-curating procedures, QA/QC protocols, and taxonomic methods can be found in Rykken and Farrell (2013, 2015). All specimens were deposited in the entomology and invertebrate collections of the Museum of Comparative Zoology (MCZ), Harvard University, Cambridge, MA. Statistical analyses We selected 6 representative arthropod taxa to assess the completeness of our inventory. Collectively, these focal taxa represented a diversity of feeding groups and dispersal abilities (Table 1). Each focal taxon had been thoroughly sampled with appropriate trapping and collecting methods. Within each focal taxon, most or all of the specimens had been identified to species or morphospe cies by an expert. To estimate absolute species richness, we used sample-based abundance data to calculate the Chao 1 richness estimator with log-linear 95% confidence intervals (Chao 1984): SChao 1 = Sobs + F1 2 / 2F2 , where Sobs is the total number of species observed in all the samples (including all collecting methods) pooled, and Fi is the number of species that have exactly i individuals when all samples are pooled. Therefore, as the number of singletons and doubletons increases, the estimate increases. We conducted all statistical calculations in EstimateS version 8.2 (Colwell 2011) Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 26 Vol. 25, Special Issue 9 Results Between 2005 and 2010, more than 50 students, interns, and volunteers processed ~160,000 arthropods and gastropods collected on 19 islands. Of these, 76,539 specimens were identified by more than 40 taxonomists from North America and Europe. There are currently 1732 distinct species in the database (see Rykken and Farrell 2013 for complete species list), of which 1643 have valid species epithets; the remainder were recorded as morphospecies. High-resolution digital images of most identified species were also databased. The 1732 species were distributed among 7 taxonomic classes, 24 orders, and 201 families (Fig. 1; Appendix A); 232 species were known or suspected to be non-native to North America, representing 13.4% of the total. Among these were several new US and North American records (Table 2). Among native species, there were also several new state records confirmed by taxonomists (Table 2). Most of the species we collected were new records for the park, with the notable exception of the majority of Macrolepidoptera, which Mello thoroughly surveyed in 2001 and 2002 (Mello 2005). Taxa with relatively high proportions of non-native species included several of the less diverse, ground-dwelling, non-insect invertebrate groups: order Isopoda (sowbugs: 75% non-native), class Diplopoda (millipedes: 87% non-native), and class Gastropoda (snails and slugs: 36% non-native; Fig. 1). Among the more diverse insect orders (more than 10 species collected), Coleoptera (beetles) and Hemiptera (true bugs and relatives) had the highest proportion of non-native species (18% and 12%, respectively). We documented far more species of Coleoptera than of any other order, which is to be expected because Coleoptera are the most diverse order of insects in nature. However, the number of species documented for each order or class reflects only the specimens that were identified (and the availability of taxonomists to work on particular groups), and does not take into account all the specimens that were Table 1. Six focal taxa selected for species richness estimates in Boston Harbor Islands ATBI. Taxon Common name Feeding mode Dispersal mode Anthophila Bees Herbivore All species fly, strong fliers (pollen, nectar) Carabidae Ground beetles Predator Good runners, many can fly (some seed eaters) Curculionoidea (Anthribidae, Weevils Herbivore All species crawl, some Brentidae, Curculionidae) (various plant parts) also fly Diplopoda Millipedes Detritivore All species crawl, none fly Formicidae Ants Scavenger, predator, All species crawl, males herbivore (plants, and queens fly animals, fungi) Mycetophilidae, Keroplatidae Fungus gnats Fungivore (some spp. All species fly, weak fliers with predatory larvae) Northeastern Naturalist 27 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 collected but remained unidentified. We estimate that more than half of the specimens (~84,000) collected during the 6-y ATBI remain unidentified (Fig. 2). Most of the unidentified specimens are in the orders Hymenoptera (~14,000), Diptera (~61,000), and Araneae (~4000). Among the unidentified Hymenoptera, most are parasitic Apocrita. Only 3 families of dipterans (Syrphidae, Keroplatidae, and Mycetophilidae) were sent to specialists for identification; the remainder of the flies (i.e., the vast majority) remain unidentified. Species richness estimates for focal taxa Millipedes (comprising 6 families) were the least diverse group of the focal taxa, with 15 species; bees (comprising 5 families) were the most diverse, with 172 species. The observed species richness for millipedes equaled the estimated absolute species richness predicted by the Chao 1 richness estimator, and fungusgnat species richness fell within the 95% confidence interval of the estimate, while Figure 1. Number of native and non-native species (including morphospecies) identified across 4 classes (Gastropoda, Diplopoda, Collembola, Chilopoda) and 16 orders (within 3 classes: Arachnida, Insecta, and Malacostraca) during the Boston Harbor Islands ATBI. Coleoptera = beetles; Hymenoptera = wasps, ants, and bees; Hemiptera = true bugs and hoppers; Lepidoptera = moths and butterflies; Diptera = flies; Araneae = spiders; Orthoptera = crickets, grasshoppers, and katydids; Gastropoda = snails and slugs; Odonata = damselflies and dragonflies; Diplopoda = millipedes; Collembola = springtails; Isopoda = sowbugs and pillbugs; Dictyoptera = cockroaches and mantids; Amphipoda = scuds or beach fleas; Dermaptera = earwigs; Neuroptera = lacewings; Mecoptera = scorpionflies; Megaloptera = alderflies, dobsonflies, and fishflies; Chilopoda = centipedes; Opiliones = harvestmen. Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 28 Vol. 25, Special Issue 9 ant species richness fell just below the confidence interval (Fig. 3). The observed species richness for each of the other groups (weevils, ground beetles, and bees) comprised 76–82% of the estimated absolute richness and fell well below the 95% confidence interval (Fig. 3). Discussion Meeting inventory objectives The Boston Harbor Islands ATBI had 2 main scientific goals: (1) to document as many species as possible in the park, and (2) to describe and compare patterns of species distribution across habitats and islands. These 2 broad goals of biodiversity inventories have been categorized as “strict inventory” and “community characterization” by Longino and Colwell (1997), and while the former typically relies on active sampling by taxonomic experts to generate species lists (such as in bioblitzes), the latter uses structured, replicated methods to capture and compare variability over space and time. Strict inventory allows a much more streamlined collecting and processing effort, requiring just 1 or a few specimens to document Table 2. Species collected during the Boston Harbor Islands ATBI in 2005–2010 that represent new records or first published records (see footnotes) for Massachusetts (MA), the United States (US), and North America (NA). Order Family Species Record Origin Coleoptera Carabidae Amara aulica (Panzer) MA Europe Amara bifrons (Gyllenhal) MA Europe Apenes lucidulus (Dejean) MA N. America Harpalus rubripes (Duftschmid) MA Europe Laemostenus terricola terricola (Herbst) US Eurasia Chrysomelidae Epitrix pubescens (Koch)A NA Europe Longitarsus rubiginosus (Foudras)B US Europe Paria sexnotata (Say) MA N. America Elateridae Agriotes lineatus (L.) MA Europe Athous haemorrhoidalis (Fabricius) US Europe Athous cf. bicolor (Goeze)C NA? Europe Staphylinidae Lordithon obsoletus (Say) MA N. America Sepedophilus immaculatus (Stephens) NA Europe Hemiptera Cicadellidae Hishimonus sellatus (Uhler) NA Asia Hymenoptera Formicidae Camponotus caryae (Fitch) MA N. America Anergates atratulus (Schenk) MA Europe Myrmica scabrinodis Nylander NA Europe Pyramica metazytes Bolton MA N. America Halictidae Lasioglossum lionotum (Sandhouse) MA N. America AThis is the first published record of Epitrix pubescens in North America, although specimens have been collected from ON and QC (in 1975) and NH (in 1992) (Deczynski 2016). BThis is the first published record of Longitarsus rubiginosus in North America, although specimens were collected in NH in 1992 (D. Chandler, University of New Hampshire, Durham, NH, pers. comm.). CThis very likely represents a new record for North America, but confirmation of the species determination requires additional DNA analysis (H. Douglas, Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, ON, Canada, pers. comm.). Northeastern Naturalist 29 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 each species. Structured sampling, on the other hand, typically relies more on passive trapping techniques that can be deployed by non-specialists. In this process, large specimen-sample sizes that incorporate species redundancy and spatial/ temporal replication are critical for documenting and comparing community patterns among habitats or islands. In the Boston Harbor Islands ATBI, we attempted to combine both types of sampling methods, following the traditional and structured sampling model of the Great Smoky Mountains National Park ATBI (Nichols and Langdon 2007). However, luring taxonomists to collect in this small urban park proved challenging, and so the bulk of the sampling was done by non-specialists (i.e., students and volunteers), who used traps to document species and characterize communities. The trade-off for relying on structured trapping by inexperienced volunteers versus active sampling by experts is that more-cryptic species (e.g., specialist species associated with particular plants or rare microhabitats) are often missed or take longer to find. Among our focal taxa, observed species richness for the more diverse groups (weevils, ground beetles, bees) fell short (76–82%) of estimated absolute species richness. For less-diverse taxa, such as millipedes, ants, and fungus gnats, Figure 2. Number of identified specimens and estimated number of unidentified specimens collected in 4 classes (Gastropoda, Diplopoda, Collembola, Chilopoda) and 16 orders (within 3 classes: Arachnida, Insecta, and Malacostraca) during the Boston Harbor Islands ATBI. Taxa are ordered left to right as in Figure 1, in decreasing order of species richness. See Figure 1 for common names of taxa. Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 30 Vol. 25, Special Issue 9 structured sampling yielded the same number, or nearly as many species as the statistical estimates for absolute richness (note that ant collection was enhanced by being the focal taxon for a student project, which included systematic hand-collecting; Clark et al. 2011). Among non-focal groups, we believe our samples likely yielded a similar pattern in terms of the number of species predicted by statistical estimates. For example, we can be fairly certain that most if not all of the species of isopods (Isopoda), amphipods (Amphipoda), and earwigs (Dermaptera) have been documented on the islands we sampled, but more-diverse taxa such as rove beetles (Staphylinidae) or many of the diverse wasp and fly families were likely undersampled by our surveys. Additional types of traps or collecting may also have yielded a higher diversity of taxa. For example, we did not use pheromone or bait traps which target taxa such as bark beetles (Scolytinae), sap beetles (Nitidulidae), carrion beetles (Silphidae), mosquitoes (Culicidae), and ticks (Ixodida) far more efficiently than pitfall or Malaise traps. Similarly, more systematic active collecting such as timed branch-beating and/or sweep netting would likely have sampled a higher diversity of leaf feeders such as leaf beetles (Chrysomelidae) and leafhoppers (Cicadellidae). Among spiders, orb-weavers (e.g., Araneidae) are best collected by hand, while our pitfall traps targeted ground-dwelling species (e.g., Salticidae, Lycosidae). Beyond collecting, an important bias in the current biodiversity database resulted from the necessity of prioritizing the processing and curation of a subset of Figure 3. Observed (black column) and estimated (black plus white column) species richness for 6 focal invertebrate taxa in Boston Harbor Islands NRA. See Table 1 for scientific names. Estimates were calculated using the Chao 1 richness estimator; bars represent loglinear 95% confidence intervals. n = total number of collection samples used to calculate the estimates. Northeastern Naturalist 31 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 specimens from a collection of more than 160,000 based on the availability of taxonomists. The large number of samples collected also resulted in a sizeable backlog of specimens that are currently stored in vials of ethanol at the MCZ. Dipterans (flies), an extremely diverse order, represent the largest backlog, and hyper-diverse and abundant parasitic wasps in the suborder Apocrita are virtually absent in the database. Specialists for these taxonomically challenging groups are few, and often over-burdened by requests for their identification skills. Unfortunately, unidentified and/or uncurated specimen backlogs are the rule rather than the exception in comprehensive invertebrate inventories using structured sampling protocols (Parker and Bernard 2006), which points both to the need for longer-term funding for ATBIs (especially for processing, curation, and taxonomy), and to the need for more skilled taxonomists. This overall shortage of specialists for the most diverse groups, aptly described as the ‘taxonomic impediment’ (Taylor 1983), coupled with an ongoing decline in the field of taxonomy as specialists retire and fewer students enter the field (Hopkins and Freckleton 2002), will need to be addressed if ATBIs are truly expected to document all species. Noteworthy species Among 1732 identified species, there were many noteworthy finds, including 5 new introductions to North America and 3 species new to the US, as well as 11 confirmed new state records for native and non-native beetles (7 species), ants (3 species), and bees (1 species). All of the new records for ground beetles (Carabidae) are discussed in detail in Davidson et al. (2011), while newly introduced click beetles (Elateridae) are covered by Douglas (2011). Additional DNA barcoding is planned to confirm the identification of the European click beetle Athous cf. bicolor (Goeze) (H. Douglas, pers. comm.), which would represent the first documented arrival to North America of this species. Other species represented rare finds in Massachusetts. For example, Sphodros niger Hentz (Atypidae), a large mygalomorph purseweb spider, which we found on Calf and Grape Islands, has rarely been collected in Massachusetts (H. Levi [now deceased], Harvard University, Cambridge, MA, pers. comm.); there are existing records from Martha’s Vineyard, Cape Cod, and Walden Pond in Concord (Edwards and Edwards 1990). In Connecticut, the species is listed as “important” on the Connecticut Species of Greatest Conservation Need 2015 Wildlife Action Plan (www. ct.gov/deep/wildlife/pdf_files/nongame/ctwap/ctsgcn.pdf; accessed 11 January 2017). Another species also on this Connecticut list is the small, wetland-associated ground beetle Badister transversus Casey (Carabidae), of which we found a single specimen on Spectacle Island. Neither species is included on the Massachusetts List of Endangered, Threatened, and Special Concern Species. One of the more exciting non-native finds was a small, shore-dwelling ground beetle known previously from just a few specimens collected in Massachusetts in 1897, when it was (falsely) described as a new species, Bembidion puritanum, by Hayward. Erwin and Kavanaugh (1980) eventually synonymized the specimens with a European coastal species, B. nigropiceum Marsham. They speculated that Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 32 Vol. 25, Special Issue 9 the species had come to North America with the shipping trade in the 1800s, but had not persisted, thus explaining why no one had seen the species again in almost a century. Our rediscovery of B. nigropiceum on 3 islands in 2007 suggests that the beetle, which is small, flightless, and restricted in habitat, may have been present but overlooked for more than 100 years (Davidson and Rykken 2011). In some cases, it is still unclear whether a species is native or introduced. For example, we found the millipede Thalassisobates littoralis Silvestri (Nemasomatidae) on 6 islands, and in great abundance (>100 specimens) on Calf and Grape Islands. The species is native to coastal parts of western Europe down to the Mediterranean and is known in North America only from 1 site in Virginia and possibly 1 previous record from Massachusetts (Hoffman 1999). Thalassisobates littoralis is a littoral (beach) species that is prone to drifting, and there is some uncertainty about whether it arrived in North America with or without the aid of human transport (Enghoff 1987). Habitat associations and management considerations Rykken and Farrell (2013) presented an analysis of invertebrate-species distributions across islands in the park and showed that, while island size was positively correlated with species richness for some taxa, habitat type and diversity within islands were also influential for some taxa. One example is the relatively rare occurrence of fresh water, which has attracted a suite of hygrophilic invertebrate species to very small wetlands on Grape and Calf Islands, including 12 species of riparian ground beetles found nowhere else on the islands. On these 2 islands, pitfall traps in wetland habitats yielded from 15–57 more invertebrate species than in any other habitat type (e.g., meadow, shrub, beach, woodland). Fortunately, the freshwater marsh on Grape Island is managed for native plant diversity by active removal of invasive plants, and this practice will likely also benefit invertebrates. The small wet meadow on the western shore of Calf Island is obscure and not likely to be noted as a significant resource without evidence from inventories such as ours. A much more common habitat in this island park is the marine littoral zone, including sand and gravel beaches, dunes, salt-marsh edges, and the vegetative drift and wrack that accumulate in these areas. There are numerous invertebrate species closely associated with the littoral zone, and many of them specialize in 1 particular microhabitat. For instance, we found several of the species Majka and Ogden (2006) referred to as “beach-drift beetles”, including, the ant-loving beetle Brachygluta abdominalis (Aubé) (Staphylinidae), the clown beetle Hypocaccus fraternus (Say) (Histeridae), and the antlike flower beetle Sapintus pusillus (Laferté-Sénectère) (Anthicidae). Davidson and Rykken (2011) described in detail the microhabitat of the rediscovered ground beetle Bembidion nigropiceum as a 1-m–wide gravel tube “pushed up by the seawater at the high-tide line”. Majka and Ogden (2006) pointed out that the beach-drift environment is vulnerable to human disturbance, especially through “clean up” to make beaches appealing for recreation. The restricted habitats of B. nigropiceum and other species in the park suggest that managers should Northeastern Naturalist 33 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 consider beach habitat from a micro-scale perspective if active management actions, such as beach stabilization, are planned. Many herbivorous insects are closely associated with particular plant hosts for food or shelter or both. Although non-native plants are hosts for many insects on the islands, the native and critically imperiled maritime juniper woodland/shrubland community, found in small patches on World’s End, Langlee Island, and Ragged Island (Elliman 2005), is host to a suite of insects that specialize on Juniperus (juniper), including Callophrys gryneus (Hübner) (Juniper Hairstreak; Lycaenidae), Patalene olyzonaria (Walker) (Juniper Twig-geometer; Geometridae), and the leaf beetle Paria sexnotata (Say) (Chrysomelidae). The latter species is a new record for the state (E. Riley, Texas A & M University, College Station, TX, pers. comm.). A much more common and very abundant native plant on the islands is Rhus typhina L. (Staghorn Sumac), which provides important nesting habitat for some stem-nesting bees such as small carpenter bees in the genus Ceratina. Two species within this genus, C. calcarata Robertson and C. dupla Say, were among the most abundant and widespread bees we documented in the park. Many other plant hosts and microhabitats are important for terrestrial invertebrates, including dead and standing wood, fungi, and exposed soil or sandy bluffs (e.g., for soil-nesting bees). Managing for native invertebrate diversity on a landscape scale relies on many of the same strategies involved in promoting and maintaining native plant diversity. Programs to restore native plant communities, to promote diversity by maintaining a mosaic of habitats in all stages of succession, and to protect vulnerable communities from human disturbance will also benefit invertebrates. Timing and frequency of mowing, herbicide use, and other activemanagement strategies should also consider potential effects on the associated invertebrate fauna. Contributing to local and regional databases In a state and regional context, the more than 1500 newly documented species in the park (both native and non-native) contributed new occurrence records that can fill important knowledge gaps beyond park boundaries. In addition to the 8 new arrivals to the US or North America, 11 new species records for Massachusetts (confirmed by taxonomists) have extended known range limits for species such as the ant Pyramica metazytes Bolton that was previously known from only southeastern states, including Mississippi, Louisiana, Alabama, and Tennessee. Undoubtedly, we collected many more species representing new state records, though could not confirm them as such due to the overall scarcity of published checklists and even unpublished data for most Massachusetts invertebrates, with the exception of charismatic taxa such as butterflies, odonates, and tiger beetles. For instance, we identified 79 potential state records for beetles (including 15 introduced species) by searching through existing records in checklists for nearby states such as Maine (Majka et al. 2011) and Rhode Island (Sikes 2004, Sikes and Webster 2005), as well as for the entire northeastern North America region (Downie and Arnett 1996a, b). Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 34 Vol. 25, Special Issue 9 Park and state checklists provide valuable local data, but the ATBI database also contributed georeferenced specimen data and high-resolution digital images to regional and global databases including MCZBase (www.mczbase.mcz.harvard. edu), Encyclopedia of Life (www.eol.org), and Global Biodiversity Information Facility (www.gbif.org). These online databases provide publicly accessible portals to ecological, taxonomic, genetic, and geographic information for species found all over the world. As these databases gather additional biodiversity data from other parks and conservation lands, natural resource managers will be able to extract spatially explicit data sets that can be used to assess and monitor the effects of some of today’s most pressing large-scale environmental threats, such as climate change, within parks and across regions. Keeping track of introduced species One obvious utility of the ATBI in the Boston Harbor Islands has been to document newly arrived non-native species. Among the park’s newly discovered first arrivals to the US and North America, 2 species are known to be pests of agricultural crops in their native ranges: the leafhopper Hishimonus sellatus (Uhler) (Hemiptera: Cicadellidae) is a vector of mulberry dwarf phytoplasmas in Asia (Kawakita et al. 2000), and the wireworm Athous haemorrhoidalis (Fabricius) (Coleoptera: Elateridae) is a below-ground pest on crop plants (Douglas 2011). It is likely that these species also now occur on the mainland, and indeed, 2 photographic records of H. sellatus in eastern Massachusetts were submitted to the online arthropod identification website BugGuide (http://bugguide.net/node/view/611930; accessed 11 January 2017) in 2011 and 2012. Other non-native species collected in the park were recently introduced to the region, and park records will be important for documenting range expansions. Examples include Megachile sculpturalis Smith (Giant Resin Bee) from Asia, which has been observed to invade nests of the native Xylocopa virginica (L.) (Eastern Carpenter Bee; Laport and Minckley 2012), and the ambrosia beetle Ambrosiophilus atratus Eichhoff from Asia, which is currently being monitored by the USDA (Haack 2006). Invertebrates as indicator taxa for monitoring In a visionary workshop hosted by the National Park Service (NPS) in 1992, Ginsberg and others proposed that a national network of NPS sites be established to monitor invertebrate biodiversity, and set as a priority the development of protocols for selecting suites of species to serve as indicators of ecosystem health (Ginsberg 1993). Worldwide, invertebrate taxa have been used to monitor the integrity of various terrestrial and aquatic ecosystems because they possess traits that make them effective indicators, such as large population size, high diversity, short generation times, ease of sampling, and a range of sensitivities to environmental gradients such as moisture and temperature (Kremen 1992, McGeoch 1998). These characteristics also make them ideal candidates for monitoring effects of large-scale environmental threats such as global climate change, pollution, habitat fragmentation, and invasive species (Leal et al. 2012, Menéndez 2007). Northeastern Naturalist 35 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 Over the last decade more and more national parks have begun their own focused invertebrate inventory and monitoring programs to gather baseline data on taxa of concern such as pollinators (e.g., Rochefort 2016, Rykken 2015, Rykken et al. 2014). In Boston Harbor Islands, we extended the ATBI for an additional year by acquiring funding to set up a pilot bee-monitoring program (Rykken and Farrell 2015). This project not only added 23 new bee species to the ATBI database, but also established a structured sampling protocol and baseline database for future bee monitoring. There are many other possibilities for using invertebrates as indicators in park monitoring programs. For example, native plant restoration is a priority management action on many of the Boston Harbor Islands, and includes a vigorous, volunteer-based invasive-plant removal program. As native plant communities are gradually restored on the islands, it will be important to document if and how associated pollinator and/or other herbivorous insect communities shift in abundance, diversity, and species composition, as this will influence the successful long-term reestablishment of functioning ecosystems. The value of inventory By many measures, the Boston Harbor Islands terrestrial invertebrate ATBI was an ambitious and successful effort. Documenting diversity in a small urban island park provided novel information on island biogeography (Rykken and Farrell 2013), the prevalence of introduced species (see also Rykken and Farrell 2018 [this issue]), and, even more simply, on how diverse a small urban park can be. Diversity exceeded taxonomists’ expectations for many groups. For example, we more than doubled E.O. Wilson’s ant estimate for the park (25 species), by collecting 52 species. Our bee species count, at 172, represents just under half of the known bee taxa for Massachusetts (~388 species; M. Veit, Lawrence Academy, Groton, MA, pers. comm.). Comparisons to other coastal islands in Massachusetts also indicate relatively high species richness in Boston Harbor for some groups. For example, Purrington (1996) collected 102 carabid beetle species on Nantucket, compared to our 128 species. Of course, for most groups, we have no state checklists or local inventories with which to compare our numbers, and the ATBI itself provides a foundation of diversity data on which future inventories will build. We also hope that these data will inspire further inquiry into the ecology, natural history, population biology, and genetic diversity of invertebrates in Boston Harbor Islands, and that the National Park Service, in collaboration with scientists, will continue to incorporate invertebrate diversity into its educational programs and management considerations. Acknowledgments Funding was generously provided by the Stone Foundation, the Green Fund, and the National Park Service. The success of the ATBI relied on the skills and effort provided by many talented people, including: staff at the MCZ, Harvard University (Carmen Chavez, Whit Farnum, Amie Jones, Piotr Naskrecki, and Dave Wrobel); staff at Boston Harbor Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 36 Vol. 25, Special Issue 9 Islands NRA and Boston Harbor Islands Partnership (Marc Albert, Jennifer Bourque, Betsy Colby, Brian Conroy, Kelly Fellner, Kristen Hoffman, Beth Jackendoff, Bridget McDonald, Josh Parker, Andrew Pearson, Mary Raczko, Terri Teller, Dawn Tesorero, Gabe Wallman, David Weinstein); our veritable army of students, interns, and citizen scientists (Celina Abundis, Laila Alawa, Jeanne Andersen, Rosalind Becker, Emily Boehm, Dunbar Carpenter, Sung Won Cho, Maesen Churchill, Adam Clark, Betsy Colby, Susannah Corona, Erika D’Andrea, Alex Dolginow, Meredith Eustis, Shannon Fadden, Ashley Fillmore, Marcelle Goggins, Emily Hill, Anna Holden, Sean Hooley, Sean Kent, Daniel Kim, Josette Kimbrough, Lili Kocsis, Veronica Kratman, Ling Lin, Victoria Luu, Stephanie Madden, Amanda Newton, Emily Nguyen, Katie Nishimura, Kelley Nunn, Michael Peters, Alison Ravenscraft, Margaret Ross, Matan Shelomi, Georgia Shelton, Sebastian Velez, L. Ray Watkins, Jessica Worl, and Serena Zhao); and last, but not at all least, the taxonomists who contributed their expertise to provide us with species names (Robert Anderson, Mike Arduser, John Ascher, Adam Baldinger, Ross Bell, Ernest Bernard, Adam Brunke, Don Chandler, Adam Clark, Stefan Cover, Matt Dakin, Bob Davidson, Dan Dourson, Sam Droege, Robert Footit, Tom French, Jason Gibbs, Matt Gimmel, Gonzalo Giribet, Pat Gorring, Andy Hamilton, Richard Hoebeke, Joe Keiper, Sean Kent, Paul Lago, Serge Laplante, Jennifer Lenihan, Jim MacDougal, Stephanie Madden, Mark Mello, Joan Milam, Frank Model, Piotr Naskrecki, Allison Ravenscraft, Ed Riley, Wolfgang Rücker, Bjorn Rulik, Bruce Snyder, Bill Stubblefield, Katalin Szlavecz, Chris Thompson, Michael Veit, Ferenc Vilsics, and Dave Wagner). Literature Cited Chao, A. 1984. Non-parametric estimation of the number of classes in a population. Scandinavian Journal of Statistics 11:265–270. Clark, A.T., J. Rykken, and B.D. Farrell. 2011. The effects of biogeography on ant diversity and activity on the Boston Harbor Islands, Massachusetts, USA. PLoS ONE 6(11):e28045. Colwell, R.K. 2011. EstimateS: Statistical estimation of species richness and shared species from samples. Version 8.2. User's Guide and application published at http://purl.oclc. org/estimates. Accessed 15 February 2016. Davidson, R.L., and J.J. Rykken. 2011. Rediscovery of Bembidion (Lymnaeum) nigropiceum (Marsham) (= puritanum Hayward) (Coleoptera: Carabidae: Bembidiini) in Massachusetts, with remarks on biology and habitat. ZooKeys 147:48 7–496. Davidson, R.L., J. Rykken, and B. Farrell. 2011. Carabid beetle diversity and distribution in Boston Harbor Islands national park area (Coleoptera, Carabidae). ZooKeys 147:497–526. Deczynski, A.M. 2016. Morphological systematics of the nightshade flea beetles Epitrix Foudras and Acallepitrix Bechyné (Coleoptera: Chrysomelidae: Galerucinae: Alticini) in America north of Mexico. M.Sc. Thesis. Clemson University, Clemson, SC. 175 pp. Douglas, H. 2011. New records of European wireworm pests and other click beetles (Coleoptera: Elateridae) in Canada and USA. Journal of the Entomological Society of Ontario 142:11–17. Downie, N.M., and R.H. Arnett. 1996a. The Beetles of Northeastern North America, Volume I. The Sandhill Crane Press, Gainesville, FL. 880 pp. Downie, N.M., and R.H. Arnett.1996b. The Beetles of Northeastern North America, Volume II. The Sandhill Crane Press, Gainesville, FL. 840 pp. Northeastern Naturalist 37 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 Edwards, R.L., and E.H. Edwards. 1990. Observations on the natural history of a New England population of Sphodros niger (Araneae, Atypidae). Journal of Arachnology 18:29–34. Elliman, T. 2005. Vascular flora and plant communities of the Boston Harbor Islands. Northeastern Naturalist Special Issue 3:49–74. Enghoff, H. 1987. Thalassisobates littoralis (Silvestri): An amphiatlantic millipede (Diplopoda, Julida, Nemasomatidae). Entomologist’s Monthly Magazine 123:205–206. Erwin, T.L., and D.H. Kavanaugh. 1980. On the identity of Bembidion puritanum Hayward (Coleoptera: Carabidae: Bembidiini). The Coleopterists Bulletin 34:241–242. Farrell, B.D. 2005. From agronomics to international relations: Building an online encyclopedia of life in the Dominican Republic. Revista: Harvard Review of Latin America Fall 2004/Winter 2005:6–9. Ginsberg, H. 1993. Invertebrate monitoring in the National Park System. Technical Report NPS/NARURI/NRTR-93/02. National Park Service, Narragansett, RI. 17 pp. Haack, R.A. 2006. Exotic bark- and wood-boring Coleoptera in the United States: Recent establishments and interceptions. Canadian Journal of Forest Re sources 36:269–288. Hoffman, R.L. 1999. Checklist of Millipedes of North and Middle America. Special Publication Number 8. Virginia Museum of Natural History, Martinsville, VA. 584 pp. Hopkins, G.W., and R.P. Freckleton. 2002. Declines in the numbers of amateur and professional taxonomists: Implications for conservation. Animal Conservation 5:245–249. Janzen, D.H., and W. Hallwachs. 1994. All taxa biodiversity inventory (ATBI) of terrestrial systems: A generic protocol for preparing wildland biodiversity for non-damaging use. Report of a National Science Foundation Workshop, 16–18 April 1993, Philadelphia, PA. 132 pp. Kales, D. 2007. The Boston Harbor Islands: A History of an Urban Wilderness. The History Press, Charleston, SC. 158 pp. Kawakita, H., T. Saiki, W. Wei, W. Mitsuhashi, K. Watanabe, and M. Sato. 2000. Identification of mulberry dwarf phytoplasmas in the genital organs and eggs of the leafhopper Hishimonoides sellatiformis. Phytopathology 90:909–914. Kremen, C. 1992. Assessing the indicator properties of species assemblages for naturalareas monitoring. Ecological Applications 2:203–217. Laport, R.G., and R.L. Minckley. 2012. Occupation of active Xylocopa virginica nests by the recently invasive Megachile sculpturalis in upstate New York. Journal of the Kansas Entomological Society 85:384–386. Leal, I.R., B.K.C. Filgueiras, J.P. Gomes, L. Iannuzzi, and A.N. Andersen. 2012. Effects of habitat fragmentation on ant richness and functional composition in Brazilian Atlantic forest. Biodiversity and Conservation 21:1687–1701. Longino, J.T., and R.K. Colwell. 1997. Biodiversity assessment using structured inventory: Capturing the ant fauna of a tropical rain forest. Ecological Applications 7:1263–1277. Majka, C.G., and J. Ogden. 2006. Brachygluta abdominalis (Aubé) (Coleoptera: Staphylnidae) newly recorded in Canada, with notes on other beach-drift beetles. Proceedings of the Entomological Society of Washington 108:761–764. Majka, C.G., D.S. Chandler, and C.P. Donahue. 2011. Checklist of the Beetles of Maine, USA. Empty Mirrors Press, Halifax, NS, Canada. 328 pp. McGeoch, M.A.1998. The selection, testing, and application of terrestrial insects as bioindicators. Biological Reviews of the Cambridge Philosophical S ociety 73:181–201. Mello, M.J. 2005. Inventory of Macrolepidoptera and other insects in the Boston Harbor Islands national park area. Northeastern Naturalist Special Iss ue 3:99–144. Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 38 Vol. 25, Special Issue 9 Menéndez, R. 2007. How are insects responding to global warming? Tijdschrift voor Entomologie 150:355–365. Nichols, B.J., and K.R. Langdon. 2007. The Smokies all taxa biodiversity inventory: History and progress. Southeastern Naturalist Special Issue 1:27–3 4. Parker, C., and E. Bernard. 2006. The science approach to the Smokies ATBI. The George Wright Forum 23:26–36. Paton, P.W.C, R.J. Harris, and C.L. Trocki. 2005. Distribution and abundance of breeding birds in Boston Harbor. The Northeastern Naturalist 12:145–168. Purrington, F.F. 1996. Ground beetles of Nantucket Island, Massachusetts: 1995 (Coleoptera: Carabidae). Journal of the New York Entomological Society 104:95–103. Richburg, J.A., and W.A. Patterson III. 2005. Historical description of the vegetation of the Boston Harbor Islands: 1600–2000. Northeastern Naturalist S pecial Issue 3:13–30. Rochefort, R. 2016. Monitoring subalpine butterflies as climate changes. NPS Resource Brief. North Cascades National Park Service Complex and Mount Rainier National Park. Available online at https://www.nps.gov/articles/cascades-butterfly-project.htm. Accessed 11 April 2017. Rosen, P.S., and D. FitzGerald. 2004. Processes and evolution of Boston Harbor Islands: Peddocks and Lovells Islands. Pp. 197–210, In L. Hansen (Ed.). Guidebook for Field Trips in the Eastern Massachusetts Region. New England Intercollegiate Geological Conference, 96th Annual Meeting. 320 pp. Rykken J. 2015. Insect pollinators of Denali National Park and Preserve: A survey of bees (Hymenoptera: Anthophila) and flower flies (Diptera: Syrphidae). Natural Resource Report NPS/DENA/NRR—2015/952. National Park Service. Fort Collins, CO. 29 pp. Rykken, J.J., and B.D. Farrell. 2013. Boston Harbor Islands all taxa biodiversity inventory: Discovering the “microwilderness” of an urban island park. Natural Resource Technical Report NPS/BOHA/NRTR—2013/746. National Park Service. Fort Collins, CO. 167 pp. Rykken, J.J., and B.D. Farrell. 2015. Developing a pollinator-monitoring program for Boston Harbor Islands National Recreation Area: A pilot study. Natural Resource Report NPS/BOHA/NRR—2015/950. National Park Service, Fort Collins, CO. 31 pp. Rykken, J.J., and B.D. Farrell. 2018. Six-legged colonists: The establishment and distribution of non-native beetles in Boston Harbor Islands NRA. Northeastern Naturalist Special Issue 9:1–22. Rykken, J., A. Rodman, S. Droege, and R. Grundel. 2014. Pollinators in peril? A multipark approach to evaluating bee communities in habitats vulnerable to effects from climate change. Park Science 31:84–90. Sikes, D.S. 2004. The Beetle Fauna of Rhode Island: An Annotated Checklist. The Biota of Rhode Island Volume 3. The Rhode Island Natural History Survey, Kingston, RI. 296 pp. Sikes, D.S., and R.P. Webster. 2005. Bioinventory of Rhode Island Coleoptera: 45 new records. The Coleopterists Bulletin 59:311–327. Taylor, R.W. 1983. Descriptive taxonomy: Past, present, and future. Pp. 93–134, In E. Highley and R.W. Taylor (Ed.). Australian Systematic Entomology: A Bicentenary Perspective. CSIRO, Melbourne, Australia. 147 pp. Trocki, C.L., N.W. Talancy, and P.W.C. Paton. 2007. An inventory of amphibians, reptiles, nonvolant mammals, and select bird species on islands in Boston Harbor. Natural Resource Technical Report NPS/NER/NRTR—2007/094. National Park Service, Boston, MA. 43 pp. Northeastern Naturalist 39 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 Appendix A. Counts for species native to and non-native to North America across 201 families of arthropods and gastropods collected between 2005–2010 on 19 islands in Boston Harbor Islands NRA. Species richness Taxon Native Non-native Total PHYLUM ARTHROPODA Class Arachnida Order Araneae Agelenidae 1 1 Amaurobiidae 1 1 Araneidae 2 1 3 Atypidae 1 1 Corinnidae 5 5 Dysderidae 1 1 Gnaphosidae 3 3 Linyphiidae 1 1 Lycosidae 6 1 7 Miturgidae 1 1 Philodromidae 1 1 Salticidae 1 1 Segestriidae 1 1 Tetragnathidae 1 1 Thomisidae 4 1 5 Order Opiliones Phalangiidae 1 1 Class Chilopoda Order Lithobiomorpha Lithobiidae 1 1 Class Collembola Order Entomobryomorpha Entomobryidae 1 3 4 Isotomidae 3 3 Tomoceridae 1 1 Order Poduromorpha Neanuridae 2 2 Order Symphypleona Katiannidae 1 1 Sminthuridae 1 1 Class Diplopoda Order Julida Blaniulidae 4 4 Julidae 5 5 Nemasomatidae 1 1 Order Polydesmida Paradoxosomatidae 1 1 Polydesmidae 3 3 Order Polyxenida Polyxenidae 1 1 Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 40 Vol. 25, Special Issue 9 Species richness Taxon Native Non-native Total Class Insecta Order Coleoptera Aderidae 4 4 Anobiidae 4 4 Anthicidae 12 12 Anthribidae 3 3 Bostrichidae 1 1 Brachypteridae 1 1 Brentidae 5 1 6 Buprestidae 4 4 Byturidae 1 1 Cantharidae 4 1 5 Carabidae 110 18 128 Cerambycidae 31 31 Chrysomelidae 45 13 58 Cleridae 9 9 Coccinellidae 8 4 12 Corylophidae 1 1 Cryptophagidae 3 3 Cupedidae 1 1 Curculionidae 57 38 95 Dermestidae 1 1 2 Elateridae 46 3 49 Endomychidae 3 3 Erotylidae 2 2 Eucinetidae 1 1 Eucnemidae 5 1 6 Geotrupidae 2 2 Histeridae 2 2 Hydrophilidae 2 2 Laemophloeidae 4 4 Lampyridae 3 3 Languriidae 3 3 Latridiidae 6 2 8 Leiodidae 5 5 Lycidae 1 1 Melandryidae 5 1 6 Meloidae 2 2 Melyridae 5 5 Monotomidae 1 1 2 Mordellidae 22 22 Mycetophagidae 2 2 4 Nitidulidae 12 1 13 Order Coleoptera Oedemeridae 1 1 2 Passandridae 1 1 Northeastern Naturalist 41 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 Species richness Taxon Native Non-native Total Phalacridae 5 5 Ptilodactylidae 1 1 Pyrochroidae 1 1 Scarabaeidae 19 8 27 Scirtidae 6 6 Scraptiidae 3 3 Scydmaenidae 1 1 Silphidae 3 3 Silvanidae 1 1 Sphindidae 1 1 Staphylinidae 61 26 87 Synchroidae 1 1 Tenebrionidae 21 21 Tetratomidae 1 1 Trogidae 2 2 Trogossitidae 3 3 Order Dermaptera Carcinophoridae 1 1 Forficulidae 1 1 Order Dictyoptera Blattellidae 1 1 2 Mantidae 2 2 Order Diptera Asilidae 2 2 Calliphoridae 5 5 Dolichopodidae 1 1 Keroplatidae 6 2 8 Mycetophilidae 64 3 67 Stratiomyidae 1 1 Syrphidae 38 4 42 Tabanidae 2 2 Order Hemiptera Acanthosomatidae 1 1 Achilidae 1 1 Alydidae 2 2 Anthocoridae 2 2 Berytidae 2 2 Blissidae 1 1 Cercopidae 3 2 5 Cicadellidae 88 15 103 Order Hemiptera Cicadidae 1 1 Coreidae 1 1 Cydnidae 5 5 Flatidae 2 2 Fulgoridae 1 1 Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 42 Vol. 25, Special Issue 9 Species richness Taxon Native Non-native Total Geocoridae 1 1 Hebridae 1 1 Issidae 2 2 Lygaeidae 2 2 Membracidae 5 5 Mesoveliidae 1 1 Miridae 9 6 15 Nabidae 4 4 Pachygronthidae 2 2 Pentatomidae 16 16 Reduviidae 3 3 Rhopalidae 1 1 Rhyparochromidae 11 1 12 Saldidae 3 3 Thyreocoridae 2 2 Tingidae 1 1 Order Hymenoptera Andrenidae 27 1 28 Apidae 41 1 42 Bethylidae 8 8 Chrysididae 9 1 10 Colletidae 13 13 Crabronidae 46 3 49 Dryinidae 1 1 Formicidae 47 5 52 Gasteruptiidae 1 1 Halictidae 59 1 60 Ichneumonidae 3 3 Megachilidae 25 4 29 Mutillidae 5 5 Pompilidae 20 20 Sierolomorphidae 1 1 Siricidae 1 1 Sphecidae 9 9 Tiphiidae 2 2 Vespidae 20 2 22 Order Lepidoptera Acrolophidae 1 1 Cossidae 1 1 Crambidae 18 1 19 Drepanidae 1 1 Geometridae 21 1 22 Hesperiidae 8 1 9 Lasiocampidae 1 1 Limacodidae 2 2 Lycaenidae 5 5 Northeastern Naturalist 43 J.J. Rykken and B.D. Farrell 2018 Vol. 25, Special Issue 9 Species richness Taxon Native Non-native Total Mimallonidae 1 1 Noctuidae 96 5 101 Notodontidae 7 7 Nymphalidae 11 11 Papilionidae 2 2 Pieridae 2 1 3 Pyralidae 3 3 Sphingidae 4 4 Tortricidae 2 2 Yponomeutidae 2 2 Order Mecoptera Panorpidae 1 1 Order Megaloptera Corydalidae 1 1 Order Neuroptera Chrysopidae 2 2 Order Odonata Aeshnidae 1 1 Coenagrionidae 4 4 Corduliidae 2 2 Lestidae 1 1 Libellulidae 11 11 Order Orthoptera Acrididae 9 9 Gryllidae 11 11 Rhaphidophoridae 2 2 Tetrigidae 1 1 Tettigoniidae 5 2 7 Class Malacostraca Order Amphipoda Hyalidae 1 1 Talitridae 2 2 Order Isopoda Armadillidiidae 2 2 Oniscidae 1 1 Philosciidae 1 1 Porcellionidae 1 1 Scyphacidae 1 1 Trachelipodidae 1 1 Trichoniscidae 1 1 PHYLUM MOLLUSCA Class Gastropoda Order Stylommatophora Arionidae 3 3 Cochlicopidae 1 1 Northeastern Naturalist J.J. Rykken and B.D. Farrell 2018 44 Vol. 25, Special Issue 9 Species richness Taxon Native Non-native Total Discidae 1 1 2 Helicidae 2 2 Limacidae 2 2 Punctidae 1 1 Pupillidae 2 2 Strobilopsidae 1 1 Succineidae 1 1 Valloniidae 3 3 Vertiginidae 2 2 Zonitidae 6 2 8