[Note: at the January 2005 meeting of the Steering Committee of the Pennsylvania Biological Survey, the Invertebrate Technical Committee was reorganized into four separate Technical Committees: Terrestrial Arthropods (excluding Arachnids), Aquatic Arthropods, Mollusks, and Arachnids. Individual web pages for the four Technical Committees will be produced as appropriate. The information below is compiled for all Pennsylvania invertebrates.]
The Invertebrate Technical committee (ITC) was organized under the original formation of the Survey in 1979, and produced "Chapter 2 – Invertebrates" for the 1985 Species of Special Concern in Pennsylvania publication. This committee is composed of a broad representation of Pennsylvania’s scientific, conservation and natural resource agencies including: The Carnegie Museum of Natural History, The Academy of Natural Sciences of Philadelphia and the Pennsylvania State University, Pennsylvania Fish and Boat Commission, Pennsylvania Department of Agriculture, Pennsylvania Department of Environmental Resources – Bureau of Water Quality and the U.S.Fish and Wildlife Service, The Nature Conservancy and Western Pennsylvania Conservancy. The task of this committee is tremendous. Not only it is responsible for approximately 10-12,000 + taxa, for which there is a dearth of information, but a state government trustee has not been identified for the terrestrial invertebrate fauna. Pennsylvania’s Wild Resource Conservation Fund has already supported a few invertebrate research projects. Subcommittees formed to date: Invertebrate Trusteeship Subcommittee and Mollusk Subcommittee.
photograph courtesy of Merlin Benner
11,702 species recorded in
282 are Species of Special Concern
37 are considered Extirpated
38 are known only from historical records
INVERTEBRATES: REVIEW OF STATUS IN PENNSYLVANIA
John E. Rawlins
Carnegie Museum of Natural History
Charles W. Bier
Western Pennsylvania Conservancy
Study of invertebrate animals, especially insects, in Pennsylvania has a long history dating from before Linnaeus (1750s) and includes some of the earliest publications on the North American fauna. During the 1800s, most native species of invertebrates in Pennsylvania were named, but basic information on the biology of only a few species was published. The fauna of adjacent states (especially New York and New Jersey) was well studied, and by the early 1900s the northeastern United States had become the most thoroughly studied region for invertebrates in the New World. By the 1950s, a few of the dominant groups of invertebrates had been monographed in publications specifically addressing the fauna of Pennsylvania (e.g., Ortmann  for freshwater mussels; Tietz  and Forbes  for Lepidoptera). Many other Pennsylvania invertebrates were included in revisionary treatments of specific families or genera but such information has not been summarized or reviewed in a meaningful or useful manner for management or conservation purposes.
Despite a long history of study, no invertebrate group has been inventoried in Pennsylvania with the same rigor as the vertebrate groups. Although some aquatic taxa have credible baseline inventories, many invertebrate species are still documented only by original literature references. Some species have only recently been described (Carle 1993) and many others are still not described and without published information of any kind. Even groups with great public appeal, such as butterflies, have their current distribution and biology in Pennsylvania more poorly understood than in any adjacent state (e.g., Iftner et al. 1992, Allen 1997). Unlike vertebrates, aquatic organisms, or vascular plants, no regulatory agency in state government is formally charged with stewardship for terrestrial invertebrates, a group which contains most of the species of organisms now living in the state.
Invertebrates are present in every conceivable biotic habitat, and in most ecosystems they constitute the groups with greatest species richness. Invertebrates are ecologically involved with virtually every biotic process occurring in natural communities, from pollination, herbivory, and predation to soil formation, disease transmission, nutrient cycling, and decomposition to name only a few. Most habitat types found in adjacent states are present in Pennsylvania. This broad habitat diversity, the relatively large size of the state, and its central location make it probable that the total number of invertebrate species in Pennsylvania is not exceeded by any contiguous state. An overview of invertebrate taxa occurring in Pennsylvania is given in Appendix 1.
The great diversity of invertebrates, especially insects, in natural systems makes their abundance or absence a powerful indicator of more than just the composition of biological communities, but also provides information on structural aspects of those systems (especially trophic relationships) and on the viability of processes influencing their integrity and stability over time (Kim 1993, New 1993). Invertebrates as a group are the dominant element of biodiversity in natural systems, second only to plants in biomass and unsurpassed in terms of the nature and number of their ecological associations and interactions (Wilson 1987).
STATUS OF PENNSYLVANIA INVERTEBRATES
The total number of species of organisms in Pennsylvania is here estimated to be 21,884, of which 11,702 (53.2%) are invertebrates (including Protista other than algae), 5,143 (23.5%) are plants (including algae), 3,619 (16.5%) are fungi and lichens, and 710 (3.2%) are vertebrates (Fig. 1). Insects constitute 46% of total biodiversity in the state, and at least 80% of those are terrestrial.
Fig. 1. Species diversity of the major groups of organism in the Pennsylvania biota.
This figure is based on an estimated 21,884 species of organisms in Pennsylvania.
Current information regarding the conservation status of only a few invertebrate species are known, and those are limited almost entirely to aquatic groups of insects and mollusks. For several aquatic groups baseline surveys are either nearly complete or actively under way. Freshwater mussels (Mollusca: Unionidae) are perhaps the best inventoried group, with much new information from several sources confirming the current condition of what was once an exceptionally diverse fauna (Ortmann 1919). Modern surveys have been conducted in all drainage basins by the Academy of Natural Sciences in Philadelphia (ANSP) and the Western Pennsylvania Conservancy (WPC) in collaboration with the Carnegie Museum of Natural History (CMNH) and in conjunction with the Pennsylvania Fish and Boat Commission and the US Fish and Wildlife Service (e.g., Bier 1994, Bogan 1993). Field work by WPC staff, A. Bogan, T. Proch, and others has increased awareness of rare or Endangered species and has augmented historical information available from institutional collections at CMNH and ANSP. A summary publication on the Pennsylvania fauna is in preparation by Bogan and Proch.
The only serious baseline inventories of insects to date for the entire state concentrate on aquatic lineages. Dragonflies and damselflies (Odonata) are relatively well known (170 species), beginning with early work by Beatty and Beatty (1971a, 1971b), and continued by C. Shiffer (1985). Recent work on Presque Isle (Masteller 1993) and the Allegheny National Forest (Bier and Rawlins 1994, Bier et al. 1997) has added greatly to knowledge of the fauna in northwestern counties. Although new species have been described (Carle 1980, 1993) and species thought to have been extirpated have been rediscovered (Bier and Rawlins 1994), both spatial and temporal coverage for Odonates is still very weak and much remains to be done.
Baseline inventories have been completed for other orders of aquatic insects, supported by the Pennsylvania Wild Resource Conservation Fund and Pennsylvania Fish and Boat Commission. These include 312 species of Trichoptera (caddisflies; Masteller and Flint 1992a, 1992b), and 136 species of Plecoptera (stoneflies; work by Surdick and Kim 1976, Earle 1994, Grubbs 1996, Masteller 1997) and a thorough treatment of Simuliidae (black flies: Adler and Kim 1986). Recent surveys are important contributions, but they are not geographically complete and lack historical context with literature and collection-based occurrence data. New species have been discovered and described for both Trichoptera (Sykora and Weaver 1976, 1978; Sykora and Harris 1994) and Plecoptera (Baumann and Grubbs 1996), and work continues on both orders by S. Harris, J. Sykora, J. Earle, S. Grubbs, and others.
No terrestrial lineage of invertebrates in Pennsylvania has been adequately inventoried, including conspicuous lineages such as butterflies, tiger beetles, large moths, land snails, and spiders. Robber flies (Diptera: Asilidae) were reviewed for Pennsylvania in the form of a list (Shelley 1979), but more recent information is not available. Knowledge for some large groups is especially weak, and novices soon learn that even species-level identification is a major undertaking. Butterflies (Lepidoptera: Papilionoidea) and skippers (Lepidoptera: Hesperioidea) have never been thoroughly surveyed across the state, although substantial information is available in the literature, collections, and private field notes of many observers. J. Prescott of Erie spent the later years of his life accumulating distribution information on the Pennsylvania butterflies (Prescott 1984). Other serious enthusiasts, notably D. Wright and R. Yahner, have contributed greatly to our growing knowledge of butterfly biology and status in Pennsylvania (Yahner 1996, 1998). The Nature Conservancy (TNC) has been monitoring and protecting the critically imperiled regal fritillary butterfly (Speyeria idalia), and both TNC and WPC have investigated populations of other butterflies of special concern. No thorough survey with adequate geographical breadth and with necessary historical context has been undertaken, and no in-depth publication on the Pennsylvania fauna is available. The same is true for virtually every other terrestrial group. Land snails (Mollusca: Gastropoda) remain unstudied except for county records recorded in Hubricht (1985). Spiders (Arachnida: Araneae) are virtually unknown outside collections and systematic revisions with only limited fieldwork by J. Buchkovich.
The nocturnal Macrolepidoptera ("larger moths") have been studied extensively in recent years by workers at CMNH, WPC, and TNC (Rawlins et al. 1997, Rawlins et al. 1998a), but no general publication is available despite some excellent models from the Ohio Biological Survey (e.g., Rings et al. 1992 on the noctuid moth fauna). Not a single family in the largest order of living invertebrates, the beetles (Coleoptera), has been thoroughly reviewed for Pennsylvania in recent decades. Coleopterists at CMNH have been studying several large families of ecological importance in Pennsylvania (e.g., ground beetles [Carabidae], carrion beetles [Silphidae], long horned beetles [Cerambycidae], and others), but no formal survey or inventory has been initiated. As for smaller orders of insects and minor lineages of arachnids (daddy longlegs, ticks, and mites) and myriapods (centipedes and millipedes), the level of knowledge is appallingly low with only one recent study examining a few of the frequently encountered organisms at Presque Isle (Masteller et al. 1993).
The Pennsylvania Natural Diversity Inventory (PNDI) currently recognizes 282 species of invertebrates of real or potential special concern for conservation, only 2.4% of the estimated number of invertebrate species (Appendix 2). PNDI recognition of these taxa is unofficial, and only 2 freshwater mussels are formally listed as Endangered for Pennsylvania. Of species on the PNDI list, 13.1% are thought to be extirpated from the state and an additional 13.5% are known from historical records only (Fig. 2). More than 5% of potential taxa of special concern have undetermined status (no investigation of actual status has been attempted) and nearly 9% have unknown status (investigation of status was attempted but no conclusion was ascertained). Comparison of the above figures for invertebrates with those for vascular plants or vertebrates will reveal (1) how little is known about Pennsylvania invertebrates and (2) how poorly protected invertebrates are with respect to informed awareness of their conservation status. No Pennsylvania invertebrate is thought to be extinct globally, but so little is known about factors necessary to interpret occurrence data that few conclusions can be made with confidence. These factors include information on species ranges to determine those genuinely extralimital in Pennsylvania; data on habitat requirements and restrictions, especially those involving unusual or limiting habitats such as barrens, wetlands, viable watercourses, and caves; and information on obligatory ecological associations (foodplants, pollinators, hosts for parasitoids, and fungal associates to name just a few).
Fig. 2. Conservation status of Pennsylvania invertebrates of special concern
as registered in the Pennsylvania Natural Diversity inventory (PNDI).
This figure is based on 282 species of special concern as of July, 1998.
The Pennsylvania Natural Diversity Inventory is maintained by WPC, TNC, and the Pennsylvania Bureau of Forestry, and has contributed greatly to status recognition for Pennsylvania organisms. For invertebrates PNDI has assessed status of some species, has conducted limited monitoring for a few species, and has generated special concern lists (Appendix 2), albeit unavoidably incomplete and biased in coverage in favor of taxa for which information is available. Originally PNDI suffered from limited interaction with invertebrate experts in museums and universities and by not being part of a more concerted effort, but this has improved through collaborative efforts, especially with CMNH.
Pennsylvania is behind other states in the northeastern United States in terms of recognizing and classifying species of special concern. For every group except fish, Pennsylvania lists fewer species of concern than the average of other state listings from Maryland northward (French and Pence 1996). And of all groups, invertebrates have the poorest coverage in Pennsylvania, largely due to the absence of government stewardship for terrestrial species.
An increasing number of non-native invertebrates have been recorded in Pennsylvania, formally registered in the recently established North American Non-Indigenous Arthropod Database (NANIAD) developed at The Pennsylvania State University and soon to appear on the World Wide Web through the United States Department of Agriculture, Animal and Plant Health Inspection Service, Plant Protection and Quarantine (USDA/APHIS/PPQ). In August 1998, NANIAD listed 2,419 species of non-indigenous arthropods as having been reported from the United States, of which 152 species were cited specifically for Pennsylvania, and a few others listed only from the general region including Pennsylvania (J. Grehan, pers. comm.). Many European insects have long been known to have populations in Pennsylvania, but others are just being documented (i.e., Idaea dimidiata, Geometridae, was just noted in Rawlins et al., 1998a). With more than a third of the vascular plants in Pennsylvania being alien, it is not surprising that many exotic insects in the state are herbivores or graminivores feeding on an introduced plant, e.g., Calophasia lunula (Noctuidae) feeding on Linaria biennis (Scrophulariaceae), and Harpalus (Ophonus) puncticeps (Carabidae) on seeds of Daucus carota (Apiaceae).
Some additional exotic species are expected in Pennsylvania as they are known to be established elsewhere in North America (e.g., European Noctua pronuba [Noctuidae] has gradually expanded its range southward from Canada [Passoa and Hollingsworth 1996], and was just recently established across Pennsylvania [1998 records from Berks and Greene Counties, J. Rawlins, pers. obs.]). The sudden appearance of others may be due to rapid expansion or may be a misconception based on the absence of authoritative monitoring. For example, the European Trechus quadristriatus (Carabidae) was just recorded from Pennsylvania (Rawlins et al. 1998a) and was only recently first recorded from Michigan and Ontario. Several exotic species have become serious pests (e.g., the gypsy moth, Lymantria dispar [Lepidoptera: Lymantriidae], and the zebra mussel, Dreissena polymorpha [Bivalvia: Mytilacea; French 1990]), but others are considered beneficial (e.g., honey bees) and several were intentionally introduced for biocontrol of weeds and pest insects. As baseline inventories increase in rigor and taxonomic breadth, it is expected that viable populations of many additional alien species of invertebrates will be discovered in Pennsylvania.
THREATS TO INVERTEBRATE BIODIVERSITY IN PENNSYLVANIA
The most significant threats to the ecological and evolutionary viability of invertebrate populations in Pennsylvania are the same as those influencing vascular plants and vertebrates. Of greatest concern in recent years are (1) man-induced disturbances such as result from agricultural practices (tillage, weed control, decreased floristic diversity, pesticides, and fertilizers); (2) forestry methods (timber harvest, pesticides, changes in both taxonomic composition and structural diversity of second growth forests); (3) land use changes (urbanization, suburban sprawl, road construction); (4) habitat conversion (wetlands draining, dam construction, and aquatic degradation from channelization, dredging, and pollution); (5) introduction of exotic predators and/or competitors; (6) decrease in the size and spatial continuity of habitats (fragmentation); (7) changes in background environmental conditions influencing the availability of limiting microhabitats (acid rain, soil degradation, aerial pollution, temperature change); (8) reduction in viable population size (loss of genetic diversity and susceptibility to extirpation); and (9) undesirable management effects including decreased plant diversity and changes in vegetation due to overpopulation of white-tailed deer (Odocoileus virginianus).
Perhaps the greatest threat to invertebrate biodiversity in Pennsylvania is continuing ignorance of the fauna, its biology, and ecological interactions. This unfamiliarity is due to a number of special challenges confronting modern study of invertebrates.
1. The tremendous number of species and taxonomic complexity of invertebrates renders their identification difficult and often requires the attention of a specialist. Species recognition and delimitation is often based on microscopic attributes or characters discernible only after special techniques of dissection or preparation have been employed. The lack of authoritatively curated collections, and the absence of modern systematic treatments for many taxa greatly complicates the ease and accuracy of identification. In addition, the large number of taxa and exceptionally small number of specialists translates into great difficulty in obtaining authoritative identifications or verifying tentative determinations (Fig. 3). Finally, few groups are understood well enough systematically to make it possible to identify them without retention of voucher specimens. In fact, most invertebrate identifications are suspect and not authoritative unless specimen-based documentation is employed, meaning each data record is backed by an actual specimen for which the identification may be verified in the future as knowledge of the group improves. This is true not only for microscopic taxa, but also necessary for most other invertebrates with the exception of a few large and distinctive species.
2. Specialists competent to provide authoritative identifications and information on many invertebrate groups are few in number, often not residing within the state, and usually overloaded with identification requests for their speciality. Systematic expertise includes more than just identification skills, and specialists are often the best source of information on biology, development, ecology, and conservation for the same taxa. The number of experienced systematists is declining as the number of students training for careers in systematics decreases, and the availability of systematic expertise will remain one of the more limiting factors for invertebrate study in the foreseeable future.
3. Stewardship mandated by Pennsylvania law and assigned to a government agency is lacking for all terrestrial invertebrates, and conservation and study of those groups is frequently just ignored. Aquatic groups have received better attention from the agency responsible for their oversight, the Fish and Boat Commission, but non-game diversity still suffers from management attention emphasizing game species.
4. Public perceptions, concerns, and attitudes about species are not equal across the biota. "Invertebrates" comprise an artificial group of dissimilar organisms, and public awareness of them is often confused or imprecise. In addition, obscure but ecologically significant invertebrate taxa are not recognized by the public, and therefore not valued by them. More significant is the fact that limited funds to study and conserve biodiversity are not distributed equally among species, and invertebrates invariably receive a disproportionately small allocation for research and management.
5. Invertebrates are perceived as not having ecological relevance. This is due to lack of awareness of their ecological interactions, and the continuing practice of inventory and monitoring for target species or lineages without context in the associated flora and fauna. Modern biodiversity assessment is never exclusively limited to a single species, and the critical importance of invertebrates will become evident as monitoring and management become more ecosystem-based and of necessity involve simultaneous assessment of many related organisms across a broad range of habitats.
6. The economic utility of developing our knowledge of invertebrates is not suitably appreciated by stakeholders (agriculturalists, foresters, land-owners, conservationists, scientists, and others). This is mostly due to lack of communication about invertebrate species and failure to demonstrate that knowledge of invertebrate status is good business, whether it be for crop protection or as indicators of ecosystem health.
7. Lack of a comprehensive and well coordinated system statewide for gathering and disseminating information on invertebrates prevents prompt attention to their needs, whether it be conservation of special concern taxa or management of pests. A computerized system capable of rapidly processing and distributing information on a large number of species is needed, together with establishment of uniform standards and protocols for inventory, monitoring, and research on invertebrate taxa by as many collaborators as possible.
Fig. 3. Limitations of systematic expertise on organisms in Pennsylvania,
expressed as a ratio of the total number of species in a group to the
number of resident systematic specialists. The figure is based on the
listing of specialists by groups as given in Kim and Mulfinger (1996).
CURRENT PROJECTS AND INITIATIVES INVOLVING INVENTORY AND MONITORING OF PENNSYLVANIA INVERTEBRATES
Several groups of aquatic insects are being actively inventoried at this time, including Ephemeroptera by G. Hoover (The Pennsylvania State University). Almost half of the Pennsylvania counties have been sampled and more than 230 species documented to date. A comprehensive survey of crane flies, the largest family of true flies (Diptera: Tipulidae), has been the object of baseline work by C. Young (CMNH) and J. Gelhaus (ANSP) that combines all available historical information with new collections from throughout the state. More than 200 species have been documented in this abundant group of decomposers.
Baseline studies for aquatic groups other than insects are just starting, including important groups like crayfish (Crustacea: Decapoda) by T. Nuttall (Lock Haven University). TNC has been monitoring the Refton cave planarian in Chester County, but other non-arthropod aquatic taxa are almost completely unknown. The full diversity of aquatic invertebrates, including aquatic beetles (Coleoptera), several large families of true flies (Diptera), aquatic bugs (Heteroptera), copepods, ostracods, cladocerans, annelids, and rotifers remain almost entirely undocumented for Pennsylvania.
Several habitat-based studies have recently been conducted in Pennsylvania that have greatly increased the quantity and quality of information available on invertebrate resources. A unique aspect of these studies is that they document terrestrial invertebrate diversity coincident with plant diversity. This greatly expands the utility of information on all taxa, and incorporates structural and process aspects of biodiversity in addition to the usual composition component (presence or absence of species). These include study of selected insect groups on barrens habitats (e.g., Wheeler 1991), and a continuing broad inventory by CMNH, TNC, and WPC on 6 major barrens habitats across the state that includes 10 orders of insects in 73 families and the vascular flora, with temporal coverage over the full season. TNC sampled Lepidoptera during a study of serpentine barrens corridors in the Nottingham/Chrome/Goat Hill area of Chester County, and CMNH is working with the Hawk Mountain Sanctuary Association in Berks County on a two-year survey of nocturnal Macrolepidoptera and other insect lineages.
Recently completed insect surveys on the Allegheny National Forest by CMNH and WPC have recorded much new information on hundreds of species, documented by tens of thousands of specimens. These include studies in northern and Allegheny hardwoods habitats influenced by outbreaks of the elm spanworm (Ennomos subsignarius, Geometridae; Rawlins, et al. 1997), oak forests influenced by gypsy moth defoliation (Lymantria dispar, Lymantriidae; Rawlins, et al. 1998a), and wetlands potentially affected by the forest tent caterpillar (Malacosoma disstria, Lasiocampidae; Rawlins et al. 1998b). Entomologists at The Pennsylvania State University are undertaking survey work to provide biological context for restoration of natural habitats at the Gettysburg National Battlefield. Even the single day "BioBlitz" events popularizing biodiversity in large city parks in Pittsburgh (CMNH) and Philadelphia (ANSP) have provided much new information on the composition and status of urban invertebrates.
REPOSITORIES, DATABASES, AND GAPS IN INFORMATION
The major repositories for specimen-based information on Pennsylvania invertebrates are concentrated in the collections of CMNH (insects and mollusks), ANSP (strength in mollusks, aquatic insects, and some freshwater non-arthropod taxa), and The Pennsylvania State University (arthropods, especially insects). The number of Pennsylvania invertebrates in these museums is not certain, but must exceed 4 million specimens, a number far greater than the total of all other organismal documentation for the state. The Smithsonian Institution (National Museum of Natural History) in Washington, D.C. and the American Museum of Natural History in New York also have significant holdings of Pennsylvania invertebrates, and the Museum of Biological Diversity at Ohio State University in Columbus, has important holdings of freshwater mussels from Pennsylvania. The vast majority of invertebrate specimens from Pennsylvania in these repositories (more than 95%) are not databased, and in many cases their current level of curation is such that authoritative verification of identification is needed. Information on some species of special concern has been databased, especially for freshwater mussels and some butterflies by the PNDI. Curatorial standards and collection conditions are improving in major Pennsylvania repositories, funded by support for improvement of research facilities from the National Science Foundation (CMNH and ANSP). Future priority should be given to supporting database development based on specimens already curated in these collections and to render secure the maintenance of the primary databases these institutions maintain.
An unprecedented amount of coordinated interaction among institutional databases will be necessary to ensure accuracy, to avoid redundancy in documentation, to rapidly update primary databases, and to maximize the speed with which such information is communicated to secondary databases and general users. For optimal oversight and development of data resources on Pennsylvania invertebrates, a centralized mechanism for data management is needed to coordinate gathering and dissemination of specimen-based information. Collaborators should seek to fill gaps in information for the state as a whole, with special priority for increasing spatial representation for all invertebrate taxa; increasing taxonomic coverage with emphasis on groups of conservation concern and economic importance; and augmenting phenological information to provide accurate information on voltinism, overwintering stages, and activity periods. Additional information needs include data on immature stages (especially Holometabola) and ecological requirements including associates, soils, and meteorological conditions. There is great need for specimen-based documentation of associations with clearly labelled and authoritatively determined voucher specimens, including non-invertebrate associates (host plants, parasite hosts, prey, pollinators, and so on).
ACTIONS TO IMPROVE INVENTORY AND MONITORING OF PENNSYLVANIA INVERTEBRATES
Responsible stewardship for invertebrate resources in Pennsylvania requires action in each of following areas:
A. Educate the public, legislature, and government agencies in order to develop a base of understanding and support for invertebrate issues, including recognition of the special challenges itemized above.
B. Assign responsibility for stewardship of invertebrate groups, especially resolve state trusteeship for terrestrial taxa.
C. Develop a strategic and comprehensive plan for baseline inventory that interfaces realistically with a plan for sustained, long-term monitoring in order to provide adequate geographical, temporal, and taxonomic coverage of Pennsylvania invertebrates. Essential to such a plan is the creation of a centralized system for gathering and disseminating information on invertebrates and other organisms statewide. Present systems are inadequate for total biodiversity management and at best only address taxa of special concern or economic interest (e.g., PNDI).
D. Provide a stable and continuing source of funding to remedy the financial uncertainties associated with the present limited support through regulatory agencies and funding sources such as the Pennsylvania Wild Resources Conservation Fund. State and Federal agencies should continue and expand their support for study of aquatic groups, but attention must also be given to terrestrial taxa. Strategies for linking private and public funds should be developed to avoid redundant efforts and solidify commitment of universities, museums, conservancies, and other organizations concerned about invertebrates.
E. Address the serious problem of limited systematic expertise on invertebrates by establishing programs that foster the education and employment of such specialists in Pennsylvania.
Establishing a meaningful system for inventory and monitoring of invertebrates in Pennsylvania will be a substantial task and will not be accomplished if the study of various lineages remains disconnected and isolated as in the past. Unlike more widely dispersed studies of vertebrates and plants, such an effort for invertebrates will be accomplished most effectively by establishing a central information system to coordinate and support inventory work by specialists and their supporting institutions. As baseline inventories are developed, efficient strategies for long-term monitoring can be developed that are minimally redundant and cost-effective. A collaboration for stewardship of invertebrate biodiversity in Pennsylvania requires state support, and will not survive without explicit sanction from an appropriate government agency or allocation. The effort to establish such a system in Pennsylvania will be worth it, as the survival and health of the largest portion of Pennsylvania biodiversity is at stake.
We appreciate the assistance and information provided by invertebrate specialists working on the Pennsylvania biota, including R. Androw, B. Barton, A. Bogan, R. Davidson, G. Hoover, T. Nuttall, D. Schweitzer, C. Shiffer, S. Thompson, and C. Young. In particular we benefited from access to information and resources at the Western Pennsylvania Conservancy and The Carnegie Museum of Natural History. K.C. Kim and J. R. Grehan generously provided timely information from NANIAD on exotic species. Above all, financial support from the Pennsylvania Wild Resource Conservation Fund and encouragement from its director, Frank Felbaum, is greatly appreciated.
Adler, P.H., and K.C. Kim. 1986. The black flies of Pennsylvania (Simuliidae, Diptera): bionomics, taxonomy, and distribution. Bulletin 856, The Pennsylvania State University College of Agriculture, Agricultural Experiment Station, University Park, Pennsylvania, 88pp.
Allen, Thomas J. 1997. The butterflies of West Virginia and their caterpillars. Univ. of Pittsburgh Press, Pittsburgh, Pa. 50 color plates. 388pp.
Baumann, R.W., and S.A. Grubbs. 1996. Two new species of Soyedina (Plecoptera: Nemouridae) from the Appalachian Mountains. Ent. News 107:220-224.
Beatty, G.H., and A.F. Beatty. 1971a. The Odonata of Pennsylvania: list and cross- references to literature containing data on occurrence in the state. Proc. Pa. Acad. Sci. 45:111-120.
Beatty, G.H., and A.F. Beatty. 1971b. The distribution of Pennsylvania Odonata. Proc. Pa. Acad. Sci. 45:147-167.
Bier, C.W. 1994. A survey of the mussels (Unionidae) in portions of the French Creek Basin of northwestern Pennsylvania. W. Pa. Cons. Report to the US Fish and Wildl. Serv. 97pp.
Bier, C.W., and J.E. Rawlins. 1994. A survey of the dragonflies and damselflies of the Clarion River and its tributaries near the Allegheny National Forest, Pa. Final report to Allegheny Nat. For., US For. Ser., US Dept. of Agr. 80pp.
Bier, C.W., J.E. Rawlins, R.L. Davidson, and D.P. Koenig. 1997. A survey of Odonata (Insecta) and Unionidae (Mollusca) associated with streams in the Allegheny National Forest, Pa. Report to Allegheny Nat. For., US For. Serv., US Dept. Agr. 133pp.
Bogan, A.E. 1993. Freshwater bivalves (Mollusca: Unionidae) of the Monongahela River Basin and direct tributaries to the Ohio River in Pennsylvania. Report to the US Fish and Wildl. Serv. 38pp.
Campbell, J.M. 1893. The cladoceran species of inshore habitats of Lake Erie at Presque Isle. J. Pa. Acad. Sci. 67:115-119.
Carle, F.L. 1980. A new Lanthus (Odonata: Gomphidae) from eastern North America with adult and nymphal keys to American Octogomphines. Annals of the Ent. Soc. Amer. 73:172-179.
Carle, F.L. 1993. Sympetrum janeae spec. nov. from eastern North America, with a key to Nearctic Sympetrum (Anisoptera: Libellulidae). Odonatologica, 22:1-16.
Earle, J. 1994. New records of stoneflies (Plecoptera) from Pennsylvania. Ent. News, 105:80-84.
Felbaum, F., B. Mitchell, K. McKenna, J. Hassinger, A. Shiels, J. Hart, and D. Brauning. 1995. Endangered and Threatened Species of Pennsylvania. Wild Res. Cons. Fund, Harrisburg, Pa. 80pp.
Forbes, W.T.M. 1954. Lepidoptera of New York and neighboring states. Part III. Noctuidae. Memoir 329, Cornell Univ. Agr. Exp. Station, Ithaca, N.Y. 433pp.
French, J.R.P., III. 1990. The exotic zebra mussel - a new threat to endangered freshwater mussels. End. Spec. Tech. Bull. 15:3-4.
French, T.W., and D.M. Pence. 1996. Endangered, Threatened and special concern animal species in the northeastern states: A list of species recognized by state and Federal laws. Compilation by Northeast Nongame Tech. Com., NE Wildl. Admin. Assoc. and US Fish and Wildl. Serv. 29pp.
Grubbs, S.A. 1996. Stoneflies (Plecoptera) of the Powdermill Nature Reserve, southwestern Pennsylvania. Ent. News 107:255-260.
Hubricht, L. 1985. The distributions of the native land mollusks of the eastern United States. Fieldiana (Zool.), New Series 24:1-191.
Iftner, David C., John A. Shuey, and John V. Calhoun. 1992. Butterflies and skippers of Ohio. Bull. Oh. Biol. Surv. (New Series) 9:1-212.
Kim, K.C. 1993. Biodiversity, conservation and inventory: why insects matter. Biodiv. and Conser. 2:191-214.
Kim, K. and D.J. Mulfinger. 1996. Biodiversity inventory specialists directory [Draft]. The Pa. State Univ. Compiled for Pa. Wild Res. Cons. Fund. 84pp.
Kim, K.C., H.D. Pratt, and C.J. Strojanovich. 1986. The sucking lice of North America: An illustrated manual for identification. The Pennsylvania State University Press, University Park, Pa.
Masteller, E.C. 1993. Odonata (dragonflies and damselflies) of Presque Isle State Park and Lake Erie, Erie County, Pa. J. Pa. Acad. Sci. 67:137-138.
Masteller, E.C. 1997. Plecoptera biodiversity of Pennsylvania. Rpt. to Pa. Wild Res. Cons. Fund. Second Ed. Volumes 1 and 2 [Vol. 1:1-17, 3 tables, maps; Vol. 2, data compilation]. The Pa. State Univ. at Erie, The Behrend College.
Masteller, E.C., and O.S. Flint. 1992a. Trichoptera biodiversity of Pennsylvania. Report to Pa. Wild Res. Cons. Fund. Second Ed. Volumes 1 and 2 [unpaginated]. The Pa. State Univ. at Erie, The Behrend College.
Masteller, E.C., and O.S. Flint. 1992b. The Trichoptera (caddisflies) of Pennsylvania: an annotated checklist. J. Pa. Acad. Sci. 66:68-78.
Masteller, E.C., G. Kedzierski, and S. Spichiger. 1993. Terrestrial arthropods of Presque Isle State Park, Pennsylvania, including Pseudoscorpiones, Opiliones, Isopoda, Diplopoda, Chilopoda, and selected insects. J. Pa. Acad. Sci. 67:127-131.
New, T.R. 1993. Angels on a pin: Dimensions of the crisis in invertebrate conservation. Amer. Zool. 33:623-630.
Opler, P.A. (ed.). 1985. Invertebrates. Pages 81-165 in H.H. Genoways and F.J. Brenner, eds. Species of Special Concern in Pennsylvania. Carnegie Mus. Nat. Hist. Special Publication No. 11, vi + 430pp.
Ortmann, A.E. 1909. The destruction of the freshwater fauna in western Pennsylvania. Proc. Amer. Philos. Soc. 48:90-110.
Ortmann, A.E. 1919. Monograph of the naiades of Pennsylvania. Part 3. Systematic account of the genera and species. Memoirs of the Carnegie Mus. 8:xiv + 384pp.
Passoa, S., and C.S. Hollingsworth. 1996. Distribution, identification and rate of spread of Noctua pronuba (Lepidoptera: Noctuidae) in the northeastern United States. Ent. News 107:151-160.
Prescott, J.M. 1984. The butterflies of Presque Isle State Park, Erie County, Pennsylvania. Melsheimer Ent. Ser. 34:19-23
Purrington, F.F. and R.L .Davidson. 2000. New southerly distribution records for the boreal carrion beetle, Nicrophorus vespilloides (Coleoptera: Silphidae). Entomological News 111(5): 355-358
Rawlins, J.E., C.W. Bier, S.A. Thompson, R.L. Davidson, C.W. Young, and R.A. Androw. 1998a. Study of nocturnal Macrolepidoptera, Sarcophagid flies, and Carabid beetles in oak habitats historically exposed to pesticide applications targeted on the gypsy moth on the Allegheny National Forest, Pennsylvania. Final rep. to US For. Ser. Allegheny Nat. For. 357pp.
Rawlins, J.E., C.W. Bier, S.A. Thompson, R.L. Davidson, C.W. Young, and R.A. Androw. 1998b (in prep.). Survey of non-target insects in wetlands associated with outbreaks of the forest tent caterpillar, Allegheny National Forest, Pennsylvania. Final Rep. US For. Ser., Allegheny Nat. For. [In preparation]
Rawlins, J.E., C.W. Bier, C.W. Young, R.L. Davidson, S.A. Thompson, and R.A. Androw. 1997. Study of nocturnal Macrolepidoptera in forest ecosystems associated with population outbreaks of the elm spanworm on the Allegheny National Forest, Pennsylvania. Final rep. to US For. Ser., Allegheny Nat. For. 414pp.
Rings, R.W., E.H. Metzler, F.J. Arnold, and D.H. Harris. 1992. The owlet moths of Ohio. order Lepidoptera, family Noctuidae. Oh. Biol. Sur. Bull., New Series, 9:vi + 219, 16 pl.
Shelly, Todd E. 1979. A list of known robber flies of Pennsylvania (Diptera: Asilidae). Ent. News 90:95-99.
Shiffer, C.N. 1985. Species accounts on Odonata. Chapter 2. Invertebrates. Opler, P.A. (ed.). Pages 79-165 in H. H. Genoways and F.J. Brenner, eds. Species of Special Concern in Pennsylvania. Carnegie Mus. Nat. Hist., Special Publ. No. 11, vi + 430pp.
Surdick, R.F., and K.C. Kim. 1976. Stoneflies (Plecoptera) of Pennsylvania: a synopsis. The Pa. State Univ., College of Agr., Agr. Exp. Station, Bull. 808:1-73.
Sykora, J.L., and S.C. Harris. 1994. Five new species of Hydroptila from eastern United States (Insecta: Trichoptera: Hydroptilidae). Annals Carnegie Mus. 63:67-75.
Sykora, J.L., and J.S. Weaver, III. 1976. A new species of Rhyacophila (Trichoptera, Rhyacophilidae) from western Pennsylvania. Annals of Carnegie Mus. 46:29-32.
Sykora, J.L., and J.S. Weaver, III. 1978. Three new species of Trichoptera from western Pennsylvania. Annals of Carnegie Mus, 47:1-12.
Thorne, S.G., K.C. Kim, B.J. McGuinness, and K.C. Steiner. 1995. A heritage for the 21st century: conserving Pennsylvania's native biological diversity. Pa. Fish and Boat Comm., Harrisburg, Pa. 60pp.
Tietz, H.M. 1952. The Lepidoptera of Pennsylvania. A Manual. The Pa. State College, School of Agr., State College, Pa. xii + 194pp.
Wheeler, A.G. 1991. Plant bugs of Quercus ilicifolia: myriads of mirids (Heteroptera) in pitch pine-scrub oak barrens. J. N. Y. Ent. Soc. 99:405-440.
Wilson, E.O. 1987. The little things that run the world: the importance and conservation of invertebrates. Cons. Biol. 1:344-346.
Yahner, R.H. 1996. Biodiversity conservation of butterflies and skippers in agricultural landscapes of Pennsylvania. Final rep. to Wild Res. Cons. Fund, Harrisburg, Pa. 70pp.
Yahner, R.H. 1998. Butterfly and skipper use of nectar sources in forested and agricultural landscapes of Pennsylvania. J. Pa. Acad. Sci. 3:104-108.
Overview of invertebrates known to occur in Pennsylvania.
Common Name Taxonomic Group
flagellates Superclass Mastigophora 11-100 Freshwater or parasitic
opalinates Superclass Opalinata 2-10 Gut commensals invertebrates
amebae, radiolaria, Superclass Sarcodina 11-100 Freshwater or parasitic
sporozoans Subphylum Sporozoa 2-10 Parasitic
microsporidians Subphylum Cnidospora 2-10 Parasitic
ciliates Subphylum Ciliophora 101-1000 Freshwater or parasitic
sponges Family Spongillidae 2-10 Freshwater
hydras, freshwater Class Hydrozoa 2-10 Freshwater
flatworms Class Turbellaria 11-100 Freshwater or terrestrial
flukes Class Trematoda 2-10 Parasitic
tapeworms Class Cestoda 2-10 Parasitic
nemerteans Phylum Nemertina 1 Freshwater
rotifers Phylum Rotifera 11-100 Freshwater or terrestrial
gastrotrichs Phylum Gastrotricha 2-10 Freshwater
nematodes Phylum Nematoda 101-1000 Freshwater, terrestrial, parasitic
horsehair worms Phylum Nematomorpha 2-10 Freshwater, soil, larvae parasitic
spiny-headed worms Phylum Acanthocephala 2-10 Parasitic
operculate snails Order Prosobranchia 11-100 Freshwater or terrestrial
non-operculate snails, Order Pulmonata 11-100 Freshwater or terrestrial
freshwater clams Order Schizodonta >65 Freshwater
fingernail clams Order Heterodonta 11-100 Freshwater
polychaetes Class Polychaeta 2-10 Freshwater
earthworms Class Oligochaeta 11-100 Freshwater or terrestrial
leeches Class Hirudinea 11-100 Freshwater or terrestrial
Water bears, tardigrades 2-10 Freshwater or terrestrial
scorpions Order Scorpiones 1 Terrestrial
pseudoscorpions Order Pseudoscorpiones 2-10 Terrestrial
spiders Order Araneae 101-1000 Terrestrial
daddy longlegs, Order Opiliones >10 Terrestrial
ticks and mites Order Acarina 101-1000 Freshwater, terrestrial, parasitic
fairy shrimp, Subclass Branchiopoda 11-100 Freshwater
water fleas, and kin
ostracods Subclass Ostracoda 11-100 Freshwater
copepods Subclass Copepoda 11-100 Freshwater or terrestrial
branchiurans Subclass Branchiura 2-10 Freshwater or parasitic
mysidaceans Order Mysidacea 1? Freshwater
wood lice, pill bugs, Order Isopoda 2-10 Freshwater or terrestrial
amphipods Order Amphipoda 2-10 Freshwater or terrestrial
crayfish, crabs, shrimp Order Decapoda 11-100 Freshwater or terrestrial
proturans Order Protura 2-10 Terrestrial
diplurans or entotrophs Order Entotrophi 2-10 Terrestrial
springtails Order Collembola 11-100 Freshwater or terrestrial
bristletails Order Archaeognatha 2-10 Terrestrial
silverfish and firebrats Order Thysanura 2-10 Terrestrial
Mayflies Order Ephemeroptera >230 Freshwater
dragonflies and Order Odonata >170 Freshwater
walkingsticks Order Phasmatodea 2-10 Terrestrial
grasshoppers, katydids, Order Orthoptera 11-100 Terrestrial
roaches and mantids Order Dictyoptera 2-10 Terrestrial
termites Order Isoptera 2-10 Terrestrial
stoneflies Order Plecoptera >136 Freshwater
earwigs Order Dermaptera 2-10 Terrestrial
booklice and barklice Order Psocoptera 11-100 Terrestrial
zorapterans or angel Order Zoraptera 1 Terrestrial
chewing lice Order Mallophaga 11-100 Terrestrial
biting lice Order Anoplura 2-10 Terrestrial
thrips Order Thysanoptera 11-100 Terrestrial
hoppers, cicadas, scales,
Order Hhomoptera 101-1000 Terrestrial
true bugs Order Heteroptera 101-1000 Freshwater or terrestrial
Dobsonflies and Order Megaloptera 2-10 Freshwater
ant-lions, owlflies, Order Neuroptera 11-100 Freshwater or terrestrial
beetles Order Coleoptera >1000 Freshwater or terrestrial
twisted-wing insects Order Strepsiptera 2-10 Terrestrial
scorpionflies Order Mecoptera 2-10 Terrestrial
caddisflies Order Trichoptera >312 Freshwater
moths and butterflies Order Lepidoptera >1000 Freshwater or terrestrial
true flies Order Diptera >1000 Freshwater or terrestrial
fleas Order Siphonaptera 2-10 Terrestrial
sawflies, wasps, bees, Order Hymenoptera >1000 Terrestrial
millipedes Class Diplopoda 11-100 Terrestrial
pauropods Class Pauropoda 2-10 Terrestrial
symphylans Class Symphyla 2-10 Terrestrial
centipedes Class Chilopoda 11-100 Terrestrial
tongue worms Phylum Pentastomida 2-10 Parasitic
moss animals Phylum Bryozoa 2-10 Freshwater
entoprocts Phylum Entoprocta 1? Freshwater
Invertebrate species listed in the
Pennsylvania Natural Diversity Inventory (PNDI), July 1998.
See explanation of state and global ranks at bottom of table.
Common Name Taxonomic Group
State Rank Global Rank
a freshwater sponge Spongilla lacustris S1 G?
Refton cave planarian Sphalloplana pricei S1 G1
eastern pearlshell Margaritifera S1 G4
dwarf wedgemussel Alasmidonta heterodon SX G1
triangle floater Alasmidonta undulata S3S4 G4
brook floater Alasmidonta varicosa S2 G3
three-ridge Amblema plicata S2 G5
alewife floater Anodonta implicata SH G5
cylindrical papershell Anodontoides ferussacianus S2S4 G5
purple wartyback Cyclonaias tuberculata SX G5
fanshell Cyprogenia stegaria SX G1
butterfly mussel Ellipsaria lineolata SX G4
elephant ear Elliptio crassidens SX G5
northern lance Elliptio fisheriana SH G4
Atlantic spike Elliptio producta S2 G4Q
northern riffleshell Epioblasma torulosa rangiana S1 G2T2
snuffbox Epioblasma triquetra S1S2 G3
Wabash pigtoe Fusconaia flava S2 G5
long-solid Fusconaia subrotunda S1S2 G3
cracking pearlymussel Hemistena lata SX G1
pink mucket Lampsilis abrupta SX G2
yellow lampmussel Lampsilis cariosa S2S3 G3G4
pocketbook Lampsilis ovata S2S3 G5
eastern lampmussel Lampsilis radiata S2 G5
white heelsplitter Lasmigona complanata S1 G5
creek heelsplitter Lasmigona compressa S2S3 G5
green floater Lasmigona subviridis S1 G3
fragile papershell Leptodea fragilis S1 G5
tidewater mucket Leptodea ochracea SX G4
eastern pondmussel Ligumia nasuta S1 G4G5
black sandshell Ligumia recta S3S4 G5
threehorn wartyback Obliquaria reflexa SX G5
hickorynut Obovaria olivaria SX G4
ring pink Obovaria retusa SX G1
round hickorynut Obovaria subrotunda S1 G4
orange-foot pimpleback Plethobasus cooperianus SX G1
sheepnose mussel Plethobasus cyphyus S1 G2G3
clubshell Pleurobema clava S1 G2
round pigtoe Pleurobema coccineum S2S3 G4
ohio pigtoe Pleurobema cordatum SX G3
rough pigtoe Pleurobema plenum SX G1
pyramid pigtoe Pleurobema pyramidatum SX G2
pink heelsplitter Potamilus alatus S1 G5
eastern floater Pyganodon cataracta S2S3 G5
rabbitsfoot Quadrula cylindrica S1 G3
monkeyface Quadrula metanevra SX G4
pimpleback Quadrula pustulosa SX G5
mapleleaf Quadrula quadrula S1 G5
salamander mussel Simpsonaias ambigua S1 G3
lilliput Toxolasma parvum S1 G5
pistolgrip mussel Tritogonia verrucosa S1 G4
fawnsfoot Truncilla donaciformis S1 G5
deertoe Truncilla truncata SX G5
paper pondshell Utterbackia imbecillis S1S3 G5
rayed bean mussel Villosa fabalis S1 G1G2
rainbow mussel Villosa iris S1 G5
a sac-spider Oxysoma cubana S? G?
an orb-weaver spider Singa eugenie S? G?
a sand spider Arctosa littoralis S? G?
Pennsylvania cave scud Crangonyx dearolfi S1 G1G2
Allegheny cave scud Stygobromus allegheniensis S2S3 G4
Biggers' cave scud Stygobromus biggersi S1 G1G2
Franz's cave scud Stygobromus franzi S1 G2
Shenandoah valley cave Stygobromus gracilipes S1 G2
Pizzini's cave scud Stygobromus pizzinii S1 G2
stellmack's cave scud Stygobromus stellmacki S1 G1
Potomac groundwater Stygobromus tenuis potomacus S1 G4T3Q
an ostracod Dactylocythere suteri SU GU
a crayfish Cambarus carolinus S3? G4
a crayfish Cambarus dubius S? G5
a crayfish Cambarus monongalensis S3? G5
a crayfish Cambarus thomae S? G5
n. clearwater crayfish Orconectes propinquus S3S4 G5
a crayfish Orconectes virilis S? G5
white river crayfish Procambarus acutus SU G5
Mississippi grass shrimp Palaemonetes kadiakensis SU G4
Franz's cave isopod Caecidotea franzi S1 G1
an isopod Caecidotea kenki S1 G3
Price's cave isopod Caecidotea pricei S2S3 G3
six-banded longhorn Dryobius sexnotatus SH G?
a tiger beetle Cicindela ancocisconensis S1 G3
a tiger beetle Cicindela formosa generosa S1 G5T5
beach-dune tiger beetle Cicindela hirticollis S2S3 G5
little white tiger beetle Cicindela lepida SH G4
a tiger beetle Cicindela limbalis S3 G5
cobblestone tiger beetle Cicindela marginipennis SX G2G3
a tiger beetle Cicindela patruela S2S3 G3
a tiger beetle Cicindela scutellaris rugifrons SH G5
a tiger beetle Cicindela splendida SH G5
a tiger beetle Cicindela unipunctata SH G4
Virginia big-headed tiger Megacephala virginica SH G5
American burying beetle Nicrophorus americanus SH G1
a burying beetle Nicrophorus marginata SX G?
black lordithon rove Lordithon niger SX G1
an ant Lasius minutis S? G?
Packard's lichen moth Cisthene packardii S1S3 G5
lead colored lichen moth Cisthene plumbea S1 G5
lichen moth Crambidia cephalica S1S2 G4
pure lichen moth Crambidia pura SU G4
phyllira tiger moth Grammia phyllira SH G4
joyful holomelina moth Holomelina laeta SU G5
chestnut case-bearer Coleophora leucochrysellaSX G?
an inchworm moth Apodrepanulatrix liberaria S3 G4
southern pine looper Caripeta aretaria S1 G4
a wave moth Cyclophora nanaria S1S2 G5
November moth Epirrita autumnata henshawi SU G5T5
broad-lined erastria moth Erastria coloraria S1 G4
blueberry gray Glena cognataria S1 G4G5
esther moth Hypagyrtis esther S2S3 G5
a wave moth Idaea violacearia S1 G4
Barrens itame Itame sp 1 (cf. Itame inextricata) S1 G3
twilight moth Lycia rachelae S1 G4
Barrens metarranthis Metarranthis apiciaria SH GU
promiscuous angle Semiothisa promiscuata S1 G4
shiny gray carpet moth Stamnodes gibbicostata SU G4
arogos skipper Atrytone arogos SX G3G4
dusted skipper Atrytonopsis hianna S3 G4G5
golden-banded skipper Autochton cellus SH G4
Arctic skipper Carterocephalus palaemonmanda S2 G5T5
columbine duskywing Erynnis lucilius S1S2 G4
mottled duskywing Erynnis martialis S1S2 G4
persius duskywing Erynnis persius S1S2 G4T2
black dash Euphyes conspicuus S3 G4
sedge skipper Euphyes dion S1 G4
dotted skipper Hesperia attalus slossonae SX G4T3
Leonard's skipper Hesperia leonardus S3S4 G4
cobweb skipper Hesperia metea S2S3 G4G5
salt-marsh skipper Panoquina panoquin SH G5
mulberry wing Poanes massasoit S2 G4
broad-winged skipper Poanes viator SU G5T4
broad-winged skipper Poanes viator zizaniae S1 G5T5
southern grizzled skipper Pyrgus wyandot S1 G2
scalloped sootywing Staphylus hayhurstii S1 G5
eastern cloudywing Thorybes confusis SH G4
dot-lined white moth Artace cribraria S1 G5
southern tolype moth Tolype minta S1 G5
tolype moth Tolype notialis S1 G?
red-banded hairstreak Calycopis cecrops S2S3 G5
Appalachian blue Celastrina neglectamajor S3S4 G4
sooty azure Celastrina nigra S1S2 G4
northern hairstreak Fixsenia favonius ontario S1S3 G4T4
silvery blue Glaucopsyche lygdamus S2 G5T4
Henry's elfin Incisalia henrici S2S3 G5
frosted elfin Incisalia irus S2 G3G4
hoary elfin Incisalia polia S1 G5
karner blue butterfly Lycaeides melissa samuelis SX G5T2
bog copper Lycaena epixanthe S2 G4G5
bronze copper Lycaena hyllus S2 G5
olive hairstreak Mitoura grynea S3 G5
hickory hairstreak Satyrium caryaevorum SU G4?
black-waved flannel Lagoa crispata S1 G5
Melsheimer's sack bearer Cicinnus melsheimeri S1 G5
Barrens dagger moth Acronicta albarufa SX G3G4
a dagger moth Acronicta lanceolaria SU G4
southern variable dart Anomogyna elimata SU G5
a borer moth Apamea burgessi SH G4
a borer moth Apamea cristata SU G4
a sallow moth Apharetra purpurea S2 G4
a cutworm moth Aplectoides condita S1 G4
a mallow moth Bagisara gulnare SU G4
straight lined mallow Bagisara rectifascia SU G4
boreal fan moth Brachionycha borealis SH G4
marbled underwing moth Catocala marmorata SX G4
miranda underwing moth Catocala miranda SU G4
precious underwing moth Catocala pretiosa SX G4T2
pine woods underwing Catocala sp. 1 S1 G5
a bird-dropping moth Cerma cora S? G3G4
a sallow moth Chaetaglaea cerata S1 G3G4
Barrens chaetaglaea Chaetaglaea tremula SH G5
marvel moth Chytonix sensilis S1 G4
pink star moth Derrima stellata SH G4
festive midget moth Elaphria festivoides complex SU G5
George's midget moth Elaphria georgei S? G4
pointed sallow Epiglaea apiata S2S3 G5
violet dart moth Euxoa violaris SH G4
a noctuid moth Fagitana littera SH G4
sundew cutworm moth Hemipachnobia monochromatea S2S3 G4
hop vine borer moth Hydraecia immanis SU G4
a borer moth Hydraecia stramentosa SU G4
a noctuid moth Lemmeria digitalis S2 G4G5
American brindle moth Lithomoia germana S3S4 G5T5
Thaxter's pinion moth Lithophane thaxteri SH G4
an owlet moth Macrochilo hypocritalis S? G4
Doll's merolonche Merolonche dolli S1 G3
footpath sallow moth Metaxaglaea semitaria S2 G5
northern brocade moth Oligia hausta S1 G4
a borer moth Papaipema aerata SH GH
columbine borer Papaipema leucostigma SU G4
a borer moth Papaipema marginidens SU G4
flypoison borer moth Papaipema sp 1 S2 G2
an oak moth Phoberia orthosioides S3 G4
a rustic moth Platyperigea meralis S1 G4
a cutworm moth Properigea sp 1 S1 G2G3
pink sallow Psectraglaea carnosa S1 G3
anointed sallow moth Pyreferra ceromatica SX GU
aureolaria seed borer Rhodoecia aurantiago SH G4
a cutworm moth Richia acclivis S1S2 G4G5
a cutworm moth Richia grotei S1 G4
a quaker moth Ulolonche modesta S1 G5
broad sallow moth Xylotype capax S3 G4
a zale moth Zale curema S1 G4
a zale moth Zale metata S? G5
oblique zale moth Zale obliqua S1 G5
pine barrens zale Zale sp 1 S1 G3Q?
a zale moth Zale submediana S2 G4
pine barrens Zanclognatha martha S1S2 G4
a prominent moth Datana ranaeceps SH G4
gorgone checkerspot Chlosyne gorgone SH G4
Harris' checkerspot Chlosyne harrisii S3 G4
silvery checkerspot Chlosyne nycteis S3? G5
tawny crescent Phyciodes batesii SH G3G4
pasco crescent Phyciodes selenis S3S4 G5
faunus anglewing Polygonia faunus S3S4 G5
gray comma Polygonia progne SU G5
Diana Speyeria diana SX G3
regal fritillary Speyeria idalia S1 G3
giant swallowtail Papilio cresphontes S2 G5
pink-edged sulphur Colias interior SH G5
olympia marble Euchloe olympia S1 G4
checkered white Pontia protodice SH G4
northern metalmark Calephelis borealis S1 G3G4
spiny oakworm moth Anisota stigma S? G5
regal moth Citheronia regalis SU G4
pine devil Citheronia sepulcralis SH G4
Barrens buckmoth Hemileuca maia S1S2 G4
midwestern fen Hemileuca sp 3 S1 G3G4
American chestnut Synanthedon castaneae SH G3G5
graceful clearwing Hemaris gracilis SH G4
Franck's sphinx moth Sphinx franckii SH G4
yponomeutid moth Swammerdamia castaneae SX GHQ?
earwig scorpionfly Merope tuber SU G3G5
spotted blue darner Aeshna clepsydra S2S3 G4
spring blue darner Aeshna mutata S1 G3G4
long-legged green Anax longipes S1S2 G5
ocellated darner Boyeria grafiana S3 G5
southern bog darner Gomphaeschna antilope SH G4
blue-nosed darner Nasiaeschna pentacantha S2 G5
black-banded bandwing Calopteryx aequabilis S2 G5
superb jewelwing Calopteryx amata S2S3 G3G4
Appalachian jewelwing Calopteryx angustipennis SU G4
sparkling jewelwing Calopteryx dimidiata SH G5
titian ruby-spot Hetaerina titia S2 G5
two-spotted dancer Argia bipunctulata SU G4
eastern dancer Argia tibialis SH G5
resolute damsel Coenagrion resolutum S1 G5
boreal bluet Enallagma boreale S2 G5
lateral bluet Enallagma laterale S1 G3
elegant skimmer Dorocordulia lepida S2 G5
stripe-winged baskettail Epitheca costalis SH G4
Uhler's sunfly Helocordulia uhleri S3 G5
Allegheny river skimmer Macromia alleghaniensis SH G4
ski-tailed emerald Somatochlora elongata S2 G5
forcipate bog skimmer Somatochlora forcipata S2 G5
Michigan bog skimmer Somatochlora incurvata S1 G3
lined bog skimmer Somatochlora linearis S1 G5
Williamson's bog Somatochlora williamsoni S1 G5
forked clubtail dragonfly Arigomphus furcifer S2 G5
abbreviated clubtail Gomphus abbreviatus S2 G3G4
moustached clubtail Gomphus adelphus S? G4
harpoon clubtail Gomphus descriptus S1S2 G4
brotherly clubtail Gomphus fraternus S2S3 G5
lined clubtail Gomphus lineatifrons SX G4
rapids clubtail Gomphus quadricolor S1S2 G3G4
Roger's clubtail Gomphus rogersi S1 G4
wide-tailed clubtail Gomphus ventricosus SX G3
green-faced clubtail Gomphus viridifrons S1 G3
Zorro clubtail Lanthus parvulus S3S4 G3G4
irregular snaketail Ophiogomphus anomalus S1 G3
Edmund's snaketail Ophiogomphus edmundo SX G1
midget snaketail Ophiogomphus howei S1 G3
twin-horned snaketail Ophiogomphus mainensis S3 G4
obscure clubtail Progomphus obscurus S2 G5
river clubtail dragonfly Stylurus amnicola SX G3G4
marked clubtail Stylurus notatus SX G3G4
oblique clubtail Stylurus plagiatus SX G5
zebra clubtail Stylurus scudderi S1 G3G4
Canadian white-faced Leucorrhinia proxima S2 G5
dwarf skimmer Nannothemis bella SH G4
saffron-bordered Sympetrum costiferum S1? G5
Thorey's grayback Tachopteryx thoreyi S2S3 G4
Helma's cheumatopsyche Cheumatopsyche helma S1 G1G3
Vannote's cheuma- Cheumatopsyche vannotei SH GH
Explanation of Natural Heritage Program Element Ranking
Global and State Element Ranks are listed and defined below. The ranking system is utilized by Natural Heritage Programs, such as the Pennsylvania Natural Diversity Inventory, as a way of comparing the relative conservation status of plants, animals, and natural communities, especially if other information on status does not exist. This system was developed by The Nature Conservancy in conjunction with the Natural Heritage Program Network.
STATE RANK ("S-RANK")
SX Apparently extirpated from the state
SH Known from historical records only
SU Unrankable; status investigated but uncertain
S? Unranked; status unknown and not investigated
S1 Critically imperiled (endangered) in state because of extreme rarity (5 or fewer occurrences or very few remaining individuals or acres) or because of some factor(s) making it especially vulnerable to extirpation from the state
S2 Imperiled (threatened) in state because of rarity (6 to 20 occurrences or few remaining individuals or acres) or because of factor(s) making it vulnerable to extirpation in the state
S3 Rare or uncommon in state (on the order of 21 to 100 occurrences)
GLOBAL RANK ("G-RANK")
GX Believed to be extinct throughout range
GH Known from historical records only
GU Unrankable; status investigated but uncertain
G? Unranked; status unknown and not investigated
G1 Critically imperiled (endangered) globally (5 or fewer occurrences)
G2 Imperiled (threatened) globally (6 to 20 occurrences)
G3 Either very rare and local throughout its range or found locally in a restricted range (21 to 100 occurrences)
G4 Widespread, abundant, and apparently secure globally
G5 Demonstrably widespread, abundant, and secure globally
Q Taxonomic status is questionable
NOTE: "T-RANK" is the conservation status rank for taxa such as subspecies: G4T2, indicating that the subspecies is rarer than the full species.