Progress toward a list of saproxylic beetles (Coleoptera) in the southeastern USA

Clayton R. Traylor; Michael D. Ulyshen; J. Winston Cornish; Gabriel Tigreros; Joseph V. McHugh

doi:10.3897/zookeys.1232.143989

Research Article

Progress toward a list of saproxylic beetles (Coleoptera) in the southeastern USA

Clayton R. Traylor^‡§, Michael D. Ulyshen^|, J. Winston Cornish^§, Gabriel Tigreros^¶§, Joseph V. McHugh^§

‡ Temple University, Philadelphia, United States of America

§ University of Georgia, Athens, United States of America

| USDA Forest Service, Southern Research Station, Athens, United States of America

¶ The Jones Center at Ichauway, Newton, United States of America

Corresponding author: Clayton R. Traylor ( clayton.r.traylor@gmail.com )

Academic editor: Christopher Majka

This is an open access article distributed under the terms of the CC0 Public Domain Dedication.

Citation: Traylor CR, Ulyshen MD, Cornish JW, Tigreros G, McHugh JV (2025) Progress toward a list of saproxylic beetles (Coleoptera) in the southeastern USA. ZooKeys 1232: 1-95. https://doi.org/10.3897/zookeys.1232.143989

ZooBank: urn:lsid:zoobank.org:pub:E7F3DF85-80E1-41FB-8DB4-25E9460FCC9F

Abstract

Deadwood-dependent (saproxylic) insects represent a large proportion of forest biodiversity, are major contributors to ecosystem processes, and are conservation priorities due to their sensitivity to changing forest conditions. Despite relevance across much of the world, research on saproxylic biodiversity has been concentrated in Europe where interest was first generated. A major impediment for this field elsewhere is a lack of resources to determine which species are saproxylic. Here, we attempt to facilitate research on saproxylic beetles (Coleoptera) in the southeastern USA by compiling information from 18 published studies and theses in the region. A list of 1,393 taxa (species or genera) from 74 families is provided with deadwood associations. This includes 891 taxa from 71 families that were reared or emerged from deadwood, and 831 taxa from 61 families that were collected from bulk trapping methods and considered to be saproxylic, or were previously included in a list of regional deadwood taxa. Additionally, for 293 taxa from non-economically important families that were considered to be saproxylic in a recently published study, known saproxylic habits, microhabitat associations, and conservation notes are listed. Sixty-eight of these species represent new state records in Georgia, USA. Although a checklist of saproxylic species is needed for the southeastern USA, it is precluded by a dearth of knowledge about the natural history and distribution of species in the region. Increasing our understanding of these species’ habitat requirements is essential for understanding biodiversity responses to changing forest conditions and assessing conservation needs.

Key words:

Coastal Plain, forest entomology, forest management, oak-pine forest, Piedmont, southern Appalachian Mountains, southern pine forest

Introduction

Saproxylic insects, those dependent on deadwood and associated resources for all or part of their life-cycle, are important contributors to ecosystem processes and account for a large portion of biodiversity within forests (Stokland et al. 2012). Many of these species aid in wood decomposition and nutrient cycling within forests (Ulyshen 2016), although assemblages are composed of species occupying a variety of functional roles, including phloeophages, xylophages, mycophages, and predators (Ulyshen and Šobotník 2018). Outside of deadwood, many saproxylic insects additionally perform, or are involved in, other ecosystem processes, such as pollination (Ulyshen et al. 2023). Estimates from northern and central Europe suggest that saproxylic species comprise approximately 20–25% of all invertebrates within forests (Siitonen 2001; Graf et al. 2022), although estimates are higher within some taxonomic groups (e.g., 42–65% for Coleoptera; Köhler 2000; Martikainen et al. 2000; Stenbacka et al. 2010; Gibb et al. 2013). Furthermore, many species facultatively use deadwood, increasing the proportion of forest invertebrate species utilizing this resource to 50–70% (Graf et al. 2022). Outside of Europe, the proportion of saproxylic species within forests has only been estimated for Coleoptera in the Atlantic Canada region: 63–79% (Majka 2009). Although saproxylic species occur in most insect orders (Ulyshen 2018), beetles (Coleoptera) are the most studied due to their taxonomic and ecological diversity (Gimmel and Ferro 2018).

Since the seminal work of Speight (1989) documenting the decline of saproxylic insects from intensive forest management and land-use change across Europe, great attention has been devoted to assessing the conservation status of saproxylic insects in the region (Seibold et al. 2015a; Cálix et al. 2018; Hagge et al. 2021) and documenting their basic habitat requirements and ecology (Siitonen and Saaristo 2000; Harvey et al. 2011; Horák et al. 2011; Lindman et al. 2023). Saproxylic communities are sensitive to the amounts and diversity of deadwood resources, and the quality and age of forest habitat more generally (Stenbacka et al. 2010; Gibb et al. 2013; Janssen et al. 2016, 2017; Seibold et al. 2016, 2017). Thus, much effort has been spent developing forest management practices that benefit these species, including deadwood retention or imitation of natural tree mortality dynamics (Thorn et al. 2014; Gossner et al. 2016; Heikkala et al. 2016; Roth et al. 2019; Ekström et al. 2021). Moreover, minimum thresholds of deadwood volume, under which certain species decline, have been determined for certain forest types (Müller and Bütler 2010; Lachat et al. 2012; Gossner et al. 2013). Overall, these efforts have provided evidence-based recommendations for saproxylic insect conservation in Europe.

Although saproxylic insects throughout the world are facing threats of deforestation and intensive forest management, research has been concentrated in Europe where awareness was first generated. But interest is rising in other regions where forests are the dominant vegetation (Davies et al. 2008; Seibold et al. 2015b). For example, researchers in Canada have devoted effort to this field (reviewed in Langor et al. 2008) and have produced conservation-priority lists of saproxylic species (Majka 2007). Similarly, there has been an increase in attention in the southeastern USA to assess host usage and habitat quality (e.g., Campbell et al. 2008a, b; Ulyshen and Hanula 2009a, 2010; Klepzig et al. 2012; Ferro et al. 2012a, b; Traylor et al. 2022a, 2023a, 2024). A global representation of saproxylic research is needed, as deadwood dynamics—to which saproxylic fauna are adapted—vary regionally from climatic and biotic differences (e.g., temperature and termites; Zanne et al. 2022), and the ecological relationships that guide conservation recommendations may change from these biogeographical differences. For example, the relationship between saproxylic insect biodiversity and deadwood volume may change in different climates (Lassauce et al. 2011; Lachat et al. 2012; Seibold and Thorn 2018), as ectothermic larvae can develop within smaller amounts of deadwood in warmer climates (Müller et al. 2015). Thus, it is important for management recommendations to be developed separately for each region based on the sensitivities and ecological roles of the endemic fauna within the context of local climatic conditions.

A major impediment to studying saproxylic insect biodiversity is simply recognizing which species are saproxylic. Decisions about inclusion or exclusion of species from analyses of saproxylic communities may influence the results, as well as the management practices drawn from them. This problem is especially relevant when biodiversity is assessed from trap samples, which collect a wider range of species and functional guilds than sampling directly from substrates (Micó et al. 2020). Currently, regional checklists of saproxylic species have been produced for Europe (Köhler 2000; Schmidl and Bußler 2004) and portions of Canada (e.g., Majka and Pollock 2006; Majka 2007; references cited in Langor et al. 2008). Additionally, resources are available for a few better-known and economically important insect groups in the USA, such as bark beetles (Curculionidae: Scolytinae; Wood 1982), woodboring beetles (Cerambycidae; Lingafelter 2007), and jewel beetles (Buprestidae; Nelson et al. 2008). Aside from these examples, few resources are available to aid in this task in North America as regional checklists with annotated saproxylic associations simply do not exist. Thus, studies of saproxylic biodiversity using trap samples are burdened with finding scattered primary literature documenting the natural history of each species (if any exist). Otherwise, decisions about saproxylic status must be presumed based on current classification and patterns of known natural history for congeneric or confamilial species.

Here, we provide two resources to aid research on saproxylic beetles in the southeastern USA. First, from published studies or theses in the southeastern region, we provide a list of taxa (species or genera) that either 1) emerged or were hand-collected directly from deadwood, 2) were considered to be saproxylic based on a classification process, or 3) were previously included in a regional list of common deadwood taxa. Secondly, we provide a summary of the known deadwood associations of 293 taxa from non-economically important families collected in a survey of saproxylic beetles in Georgia, USA (Traylor et al. 2023a). These resources should facilitate studies of saproxylic biodiversity in the southeastern USA and elsewhere in North America and form the foundation for a regional saproxylic checklist.

Methods

Saproxylic definition

The definition of “saproxylic” has evolved over time to fit a growing understanding of deadwood resources and the species that depend on them (Alexander 2008; Graf et al. 2022). Today, a saproxylic species is considered “any species that depends, during some part of its life cycle, upon wounded or decaying woody material from living, weakened or dead trees” (Stokland et al. 2012: 6). In addition to species dependent on wood, this definition broadly includes species dependent on non-wood resources found in or on living, weakened, or dead trees, such as the inner bark (phloem), sap runs, or slime fluxes in tree wounds. While it may seem bizarre for saproxylic species to use resources in living trees, it is widely recognized that important microhabitats can occur and accumulate over a tree’s life, such as dead branches in the canopy or interior rot forming tree hollows (Micó 2018; Seibold et al. 2018). Additionally, this definition includes species that depend indirectly on saproxylic resources. For example, species dependent on wood-decaying fungi are considered saproxylic, as well as species representing higher trophic levels (e.g., predators, parasitoids, and parasites) that are dependent on other saproxylic species as hosts or prey (Stokland et al. 2012). Moreover, species that do not consume wood or other saproxylic organisms may still be saproxylic, such as bees that nest in deadwood or tree cavities (Ulyshen 2018).

An important distinction exists between species that are obligately dependent on deadwood and associated resources, versus those that use these habitats facultatively. In general, saproxylic species are those that are obligately dependent, and thus would disappear if those resources were removed from forests (Stokland et al. 2012). However, distinguishing facultative or obligative use of deadwood (i.e., non-saproxylic vs saproxylic) can be difficult and largely depends on incomplete information known about a species’ biology.

List of saproxylic Coleoptera compiled from previous studies

From 18 published studies and theses in the southeastern USA (Table 1), we compiled a list of coleopteran taxa (species and genera) that either 1) emerged or were hand-collected directly from deadwood, 2) were considered saproxylic following some explicit determination process, or 3) were included in a previous regional list of common deadwood insects. Fifteen of these studies provided species lists of Coleoptera that either emerged or were hand-collected directly from deadwood in experimental methods (Table 1). Two studies provided lists of species that were considered to be saproxylic after a determination process. Klepzig et al. (2012) used pitfall trapping to collect insects and considered beetle species as saproxylic, non-saproxylic ground beetles, or other based on familiarity with the taxa and published natural history. Traylor et al. (2023a) collected beetles with flight intercept traps and litter-sifting and classified the collected species as saproxylic or not (see below). Finally, one study created a list of wood-feeding beetles that are commonly collected in the southern USA (Hanula 1996). Many other, often older, publications provide accounts of deadwood associations based on field observations (e.g., Savely 1939; Howden and Vogt 1951), but those are not included here. Other studies assumed saproxylic status based on the known natural history of closely related taxa, but did not go through any explicit determination process (e.g., Campbell et al. 2008a, b); therefore, they were not included.

Table 1.

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Experimental studies providing species lists of Coleoptera with deadwood associations. Coleoptera were emergent from deadwood (method = E), hand-collected from deadwood (H), or classified (C) as saproxylic in the southeastern United States. The state(s) and ecoregion(s) sampled from each study are provided. Level III Ecoregions follow https://www.epa.gov/eco-research/ecoregions-north-america. The list of Coleoptera can be found in Suppl. material 1.

Study	State	Type III Ecoregion(s)	Method
Ferro and Gimmel 2014	LA	8.3.6 (Mississippi Valley Loess Plains)	E
Ferro and Nguyen 2016	LA	8.3.5 (Southeastern Plains); 8.3.6 (Mississippi Valley Loess Plains); 8.3.7 (South Central Plains); 8.5.2 (Mississippi Alluvial Plain); 8.5.3 (Southern Coastal Plain); 9.5.1 (Western Gulf Coastal Plain)	E
Ferro et al. 2009	LA	8.3.6 (Mississippi Valley Loess Plains)	E
Ferro et al. 2012a	TN	8.4.4 (Blue Ridge)	E
Ferro et al. 2012b	TN	8.4.4 (Blue Ridge)	E
Garrick et al. 2019	AL, NC, TN	8.4.4 (Blue Ridge); 8.4.9 (Southwestern Appalachians)	H
Gil 2008	LA	8.3.6 (Mississippi Valley Loess Plains)	H
Hanula 1996	southern region		C
Klepzig et al. 2012	LA, MS, NC, TX	8.3.5 (Southeastern Plains); 8.3.7 (South Central Plains); 8.5.1 (Middle Atlantic Coastal Plain)	C
Tigreros 2024	GA	8.3.5 (Southeastern Plains)	E
Traylor et al. 2023a	GA	8.3.4 (Piedmont)	C
Ulyshen 2014	MS	8.3.5 (Southeastern Plains)	E
Ulyshen and Hanula 2009a	SC	8.3.5 (Southeastern Plains)	E
Ulyshen and Hanula 2009b	SC	8.3.5 (Southeastern Plains)	E
Ulyshen and Hanula 2010	SC	8.3.5 (Southeastern Plains)	E
Ulyshen et al. 2010	GA	8.3.4 (Piedmont)	E
Ulyshen et al. 2020	MS	8.3.5 (Southeastern Plains)	E
Ulyshen et al. 2024	SC	8.3.5 (Southeastern Plains)	E

Only species or generic determinations (including subgeneric distinctions, when possible) were included in our lists; morphospecies at the tribe, subfamily, or family rank were excluded. We checked the validity of all taxonomic names using the Integrated Taxonomic Information System (www.ITIS.gov), or with the most recent taxonomic revision of the group or other published literature.

Annotated list of saproxylic Coleoptera

We summarized saproxylic habits of 293 coleopteran taxa considered to be saproxylic from Traylor et al. (2023a). Briefly, from March–September 2020 in Clarke County, Georgia, USA, beetles were collected from forests using two methods: 1) a flight intercept trap (two clear, intersecting panels attached over a white bucket) suspended 5 m aboveground and baited with ethanol, and 2) leaf litter sifting and extraction via a Berlese funnel (see Traylor et al. 2022b, 2023a for details). For each species, we provide summary statistics of collection data. Full data of species’ abundance per sampling location, species’ abundance per sampling period for each collection method, and forest details are publicly available at a repository (Traylor et al. 2023b).

The saproxylic determination process was as follows. Species were identified and considered as saproxylic or not based on known natural history information indicating that they either develop in, or are primarily found within, deadwood or other saproxylic habitats. We used Gimmel and Ferro (2018) to exclude some families and subfamilies from saproxylic consideration. Species and morphospecies with no known natural history information were also considered saproxylic if all their congeners with known natural history could be considered saproxylic. See Traylor et al. (2023a) for more information.

Here, we assembled known natural history information for each saproxylic species (or genus for morphospecies) based on published accounts, rearing/emergence records, or verifiable images with documented natural history information from Bugguide (www.bugguide.net). Additionally, we provide specific information on microhabitat usage (if known) and conservation notes (if applicable). Unless stated otherwise, we also provide general distributions for each species from occurrences recorded in the Global Biodiversity Information Facility (GBIF; www.gbif.org) and BugGuide (www.bugguide.net). Non-mutually exclusive distribution categories are defined as follows: the southeastern United States (Virginia south to Florida, west to Arkansas and eastern Texas), the eastern United States (Minnesota south to Louisiana, and eastward), eastern North America (the eastern United States and Canada, Ontario and eastward), central United States (the “plain states”, i.e., Wisconsin south to Texas, and west to Montana and New Mexico), central North America (central United States and Canada, Manitoba west to Alberta). The assembled information is not intended to be a full review of each species’ natural history. Instead, the information is meant to provide a summary of deadwood associations and saproxylic habits exhibited by each species that can be used to aid researchers in classifying species as saproxylic or not and to give some background on required habitats or wood types. Natural history information unrelated to deadwood was not included (e.g., that adult Mordellidae visit flowers). Because natural history and host record reviews already exist for economically important groups in this region, such as Buprestidae (Nelson et al. 2008), Cerambycidae (Lingafelter 2007), and Curculionidae: Scolytinae (Wood 1982), we do not include these taxa below unless they are new state records in Georgia, USA.

Results

In total, 1,393 beetle taxa (species or genera) from 74 families had deadwood associations recorded in the 18 studies or theses (Suppl. material 1). This includes 891 taxa from 71 families that were reared or emerged from deadwood, and 831 taxa from 61 families that were collected from bulk trapping methods and considered to be saproxylic or were listed as common associates of deadwood. We urge readers to be cautious in the use of this list, as some species that were reared or emerged from deadwood are likely to be facultatively saproxylic. Additionally, some taxa considered to be saproxylic could be reclassified as facultative users of deadwood if more detailed knowledge of their natural history became available and supported the change.

Below, we provide the deadwood habitat associations of 293 saproxylic beetle taxa (281 species and 12 genera for which only morphospecies were assigned) collected from 40 sites during a survey conducted in the Piedmont region of Georgia, USA (Traylor et al. 2023a). Natural history data relevant to saproxylic habits are summarized for each species, and bionomic information is included when possible. All taxa were considered obligately saproxylic, but as before, additional information learned about the listed taxa in the future may require that some be considered facultative. Sixty-eight species are new state records in Georgia, USA and have been marked as such (* indicates that museum specimens exist from Georgia, but they are not formally published: An Annotated List of the Coleoptera of Georgia, v. 1.0; https://site.caes.uga.edu/ugca/turnbow-and-smith/). Abundance for each species per site with geographic coordinates, per sampling method, and per collection date can be found in a data repository (Traylor et al. 2023b).

Aderidae

Cnopus impressus (LeConte, 1875)

Collection information.

Eastern North America.

Saproxylic habits.

Bores into dead wood, such as persimmon (Diospyros L. (Ebenaceae)), and adults also occur on dead trees, including white oak (Quercus alba L.) stems (Valentine 1998).

Bostrichidae

Lichenophanes bicornis (Weber, 1801)

Collection information.

Bothrideridae

Bothrideres cryptus Stephan, 1989

Collection information.

USA: Georgia (new state record*): Clarke Co.: two individuals from two sites. Caught in flight trap from 21 April–2 June 2020.

Distribution.

Eastern North America (Stephan 1989).

Saproxylic habits.

Occur under the dry bark of dead oaks, where it parasitizes cerambycid larvae of the genus Parelaphidion Skiles, 1985 (Stephan 1989).

Bothrideres geminatus (Say, 1826)

Conservation.

Significantly higher abundance in primary (= old-growth) than secondary (= second-growth) forests in the southern Appalachian Mountains (Ferro et al. 2012a).

Cerylon unicolor (Ziegler, 1845)

Collection information.

USA: Georgia: Clarke Co.: 13 individuals from 10 sites. Caught in flight trap from 25 March–9 September 2020.

Distribution.

North America.

Saproxylic habits.

Adults occur under the bark of various dead hardwood and conifer trees and on fungi, and larvae develop under the bark of hardwoods (Lawrence and Stephan 1975); emerged from loblolly pine, water oak (Quercus nigra L.), and especially sweetgum, and associated with portions of dead trees close to the ground (Ulyshen and Hanula 2009a).

Ciidae

Cleridae

Chariessa pilosa (Forster, 1781)

Collection information.

Conservation.

Occurrence probability increases with the amount of landscape forest cover in the Piedmont (Traylor et al. 2024).

Cymatodera bicolor (Say, 1825)

Collection information.

USA: Georgia: Clarke Co.: a single individual from one site. Caught in flight trap from 9–21 April 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Prey upon bark and woodboring beetles and occurs under bark of various hardwoods, including hickory, and “cedar” (presumably Juniperus L. (Cupressaceae)) (Hopkins 1893; Knull 1951; Dorshorst and Young 2008).

Acalles clavatus (Say, 1831)

Collection information.

Conservation.

Significantly higher abundance in secondary (= second-growth) than primary (= old-growth) forests in the southern Appalachian Mountains (Ferro et al. 2012a).

Apteromechus pumilus (Boheman, 1837)

Collection information.

USA: Georgia: Clarke Co.: five individuals from four sites. Caught in flight trap from 25 March–16 June 2020.

Distribution.

Southeastern United States (Ciegler 2010).

Saproxylic habits.

unknown; larvae of Apteromechus Faust, 1896 mine the bark of recently dead trees (Kissinger 1963).

Cophes fallax (LeConte, 1876)

Collection information.

Saproxylic habits.

Larvae develop, and adults occur gregariously in decaying wood of many hardwoods and larch, including dead portions and exposed wood in living trees (Blackman and Stage 1918, 1924; Anderson 1952); adults also occur in solid, dry wood and old stumps (Beutenmuller 1893; Hoffmann 1942); associated with moderately decayed wood (Ferro et al. 2012a).

Conservation.

Significantly higher abundance in secondary (= second-growth) than primary (= old-growth) forests in the southern Appalachian Mountains (Ferro et al. 2012a); rare in the Maritime Provinces of Canada, possibly due to the history of intensive forest management in the region altering forest composition and structure (Majka 2007).

Tomolips quercicola (Boheman, 1845)

Collection information.

USA: Georgia: Clarke Co.: eight individuals from eight sites. Caught in flight trap from 21 April–30 June 2020.

Distribution.

Eastern North America south to Central America.

Saproxylic habits.

Emerged from loblolly pine, water oak, and sweetgum in bottomland forests (Ulyshen and Hanula 2009a); adults occur in rotten deadwood, tree hollows, and under bark of various trees, including beech and magnolia (Magnolia L. (Magnoliaceae)) (Blatchley 1928; Kissinger 1955; Ciegler 2010).

Xyleborus pfeilii (Ratzeburg, 1837)

Saproxylic habits.

Larvae are predators of woodboring insects in dead pine logs and stumps (Frost 1916; Savely 1939); occupies logs throughout decomposition (Savely 1939; Ulyshen and Hanula 2010).

Conservation.

Occurrence probability increases in old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2024).

Dipropus soleatus (Say, 1839)

Collection information.

Conservation.

Occurrence probability increases in old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2024).

Parallelostethus attenuatus (Say, 1825)

Eucinetidae

Eucinetus strigosus LeConte, 1875

Collection information.

USA: Georgia: Clarke Co.: a single individual from one site. Sifted from leaf litter from 27–28 July 2020.

Distribution.

Eastern United States.

Saproxylic habits.

unknown; species of Eucinetus Germar, 1818 breed in slime molds and basidiomycete fungi growing on dead wood, and often occur under bark or in decaying wood (Wheeler and Hoebeke 1984).

Larvae develop in the heartwood of decayed hardwoods (e.g., maple; Peterson 1960), and have emerged from dry, dead wood of various hardwoods (Knull 1946; Muona 2000); associated with moderately decayed logs (Ferro et al. 2012a).

Conservation.

Significantly higher abundance in secondary (= second-growth) than primary (= old-growth) forests in the southern Appalachian Mountains (Ferro et al. 2012a); significantly associated with old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2023a).

Melasis pectinicornis Melsheimer, 1844

Collection information.

Conservation.

Rare in the Maritime Provinces of Canada, possibly due to the history of intensive forest management in the region altering forest composition and structure (Majka 2007).

Nematodes atropos (Say, 1836)

Collection information.

USA: Georgia: Clarke Co.: 36 individuals from 19 sites. Caught in flight trap from 19 May–27 July 2020.

Distribution.

Eastern North America and Mexico.

Saproxylic habits.

larvae, pupae, and adults have been found together within the base of a dead black oak (Van Horn 1909); emerged from water oak logs (Ulyshen and Hanula 2009a), hardwood twigs (Ferro and Nguyen 2016), and American beech, elm, and maple (Dury 1904).

Nematodes penetrans (LeConte, 1852)

Collection information.

Hybosoridae

Ceratocanthus aeneus (MacLeay, 1819)

Collection information.

USA: Georgia: Clarke Co.: two individuals from two sites. Caught in flight trap from 27 July–26 August 2020.

Distribution.

Southeastern United States.

Saproxylic habits.

Larvae develop within the moist, rotting contents of hollow trees (Choate 1987); adults also occur under bark (Hoffman 2006).

Germarostes aphodioides (Illiger, 1800)

Laemophloeus fasciatus Melsheimer, 1846

Collection information.

USA: Georgia: Clarke Co.: three individuals from three sites. Caught in flight trap from 9 March–26 August 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Adults occur under bark (Thomas 1993).

Genus Lathropus Erichson, 1846

Collection information.

USA: Georgia: Clarke Co.: 18 individuals from 13 sites. Caught in flight trap from 9 March–9 September 2020.

Saproxylic habits.

Species of Lathropus occur under bark and in association with bark beetles (Thomas 1993); however, some evidence suggests the association is incidental and that Lathropus feed on fungi (Uliana 2003; Thomas 2010).

Phloeolaemus chamaeropis (Schwarz, 1878)

Collection information.

USA: Georgia: Clarke Co.: two individuals from two sites. Caught in flight trap from 16 June–14 July 2020.

Distribution.

Southeastern United States (Thomas 1993).

Saproxylic habits.

Adults occur under the bark of hardwood logs (sometimes freshly dead with fermenting phloem), especially oak, and apparently feed on fungi (Thomas 1993); emerged from water oak and sweetgum logs (Ulyshen and Hanula 2009a) and occur on freshly dead southern red oak (Gil 2008).

Placonotus zimmermanni (LeConte, 1854)

USA: Georgia: Clarke Co.: 289 individuals from 33 sites. Caught in flight trap from 9 March–9 September 2020.

Distribution.

Eastern United States.

Saproxylic habits.

Larvae develop in, and adults occur on Hypoxylon growing on oaks, including live oak (Quercus virginiana Mill.) and southern red oak, apparently feeding on conidia (Lawrence 1977; Gil 2008).

Conservation.

Occurrence probability increases in old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2024).

Enicmus tenuicornis LeConte, 1878

Collection information.

USA: Georgia (new state record): Clarke Co.: six individuals from four sites. Caught in flight trap from 6 May–30 June 2020.

Distribution.

North America.

Saproxylic habits.

Feeds on slime molds in the genus Stemonitis Gleditsch (Stemonitaceae) (Lawrence and Newton 1980); emerged from spruce (Picea A.Dietr), fir (Abies Mill.), and pine (all in Pinaceae) in the Pacific northwest (Hatch 1961), and mainly associated with coniferous forests in Atlantic Canada (Majka et al. 2009).

Leiodidae

Genus Agathidium Panzer, 1797

Collection information.

USA: Georgia: Clarke Co.: 69 individuals from 21 sites, not identified to species-level. Caught in flight trap and sifted from leaf litter from 9 March–9 September 2020.

Saproxylic habits.

Species of Agathidium are specialist consumers of slime molds (Lawrence and Newton 1980) and commonly occur under bark, in wood, or on fungi (Downie and Arnett 1996).

Conservation.

Significantly higher abundance in primary (= old-growth) than secondary (= second-growth) forests in the southern Appalachian Mountains (Ferro et al. 2012a).

Genus Anisotoma Panzer, 1797

Collection information.

USA: Georgia: Clarke Co.: a single individual from one site, not identified to species-level. Caught in flight trap from 19 May–2 June 2020.

Saproxylic habits.

Species of Anisotoma are specialist consumers of slime molds (Lawrence and Newton 1980) and occur in decaying wood (Wheeler 1979).

Lucanidae

Platycerus quercus (Weber, 1801)

Dircaea liturata LeConte, 1866

Collection information.

Conservation.

In the Piedmont, significantly associated with forests within sparsely forested landscapes (<50% forest; Traylor et al. 2023a) and occurrence probability decreases with the amount of landscape forest cover (Traylor et al. 2024).

Micromalthidae

Micromalthus debilis LeConte, 1878

Collection information.

USA: Georgia (new state record): Clarke Co.: a single individual from one site. Caught in flight trap from 14–27 July 2020.

Distribution.

Eastern North America, but spread to other regions (Philips and Young 2000).

Saproxylic habits.

Develops within red-rotted hardwoods, especially oak and chestnut (Philips and Young 2000); displays a unique and complicated life cycle with separate phases of paedogenesis and parthenogenesis (see Philips and Young (2000) for overview and Barber (1913a, b) for original description).

Monotomidae

Genus Bactridium LeConte, 1861

Collection information.

USA: Georgia: Clarke Co.: 11 individuals from nine sites, unable to identify to species-level. Caught in flight trap from 9 March–11 August 2020.

Saproxylic habits.

Species of Bactridium occur under bark of hardwoods and feed on ascomycete fungi (McElrath and McHugh 2018).

Rhizophagus sayi Schaeffer, 1913

Collection information.

USA: Georgia: Clarke Co.: three individuals from two sites. Caught in flight trap from 9–25 March 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Occur under the bark of deciduous trees where it is a predator and likely fungivore (Bousquet 1990); found in association with bark beetles (McElrath and McHugh 2018).

Conservation.

Significantly associated with young forests (regrown since 1938 and pine dominated) in highly forested landscapes (> 50% forest) in the Piedmont (Traylor et al. 2023a).

Emerged from fresh swamp white oak logs (Quercus bicolor Willd.) (Powell et al. 2016, G.S. Powell in litt.), and from moderately decayed loblolly pine logs (Ulyshen et al. 2020; Traylor et al. 2023c).

Conservation.

Significantly associated with forests in highly forested landscapes (> 50% forest) in the Piedmont (Traylor et al. 2023a).

Mordellistena masoni Liljeblad, 1918

Collection information.

USA: Georgia (new state record*): Clarke Co.: 98 individuals from 27 sites. Caught in flight trap from 2 June–11 August 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Emerged from freshly cut loblolly pine logs (MU pers. obs.; Traylor et al. 2023c).

Conservation.

Occurrence probability increases in old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2024).

Nosodendridae

Nosodendron unicolor Say, 1824

Collection information.

USA: Georgia: Clarke Co.: 15 individuals from 12 sites. Caught in flight trap from 9 March–26 August 2020.

Distribution.

Eastern United States.

Saproxylic habits.

Larvae develop within slime fluxes on hardwood trees, and adults occur in this habitat as well (Hayes and Chu 1946); the beetles feed on fermenting sap enriched with microorganisms and any predation is likely facultative (Ivie 2002a).

Conservation.

Occurrence probability increases in old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2024).

Passandridae

Catogenus rufus (Fabricius, 1798)

Collection information.

USA: Georgia: Clarke Co.: seven individuals from six sites. Caught in flight trap from 25 March–26 August 2020.

Distribution.

Eastern North America, south to Central America.

Saproxylic habits.

Larvae parasitize the pupae of woodboring beetles, and adults can occur under loose bark of dead and dying hardwood and conifer trees (Dimmock 1882; Fiske 1905).

Saproxylic habits.

Emerged from dry sections of black oak and bear oak (Quercus ilicifolia Wangenh.), with the bear oak record apparently coming from outer bark of a living tree (Ford 1973).

Conservation.

Significantly associated with old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2023a).

Petalium debile Fall, 1905

Collection information.

Collection information.

USA: Georgia: Clarke Co.: a single individual from one site. Caught in flight trap from 19 May–2 June 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Larvae develop under loose bark on the upper side of decaying logs (Young 2002b).

Conservation.

Significantly higher abundance in primary (= old-growth) than secondary (= second-growth) forests in the southern Appalachian Mountains (Ferro et al. 2012a).

Neopyrochroa femoralis (LeConte, 1855)

Collection information.

USA: Georgia: Clarke Co.: two individuals from two sites. Caught in flight trap from 25 March–9 April 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Larvae develop under bark and in decayed wood of standing dead trees (Young 2002b).

Conservation.

Silvanidae

Cathartosilvanus imbellis (LeConte, 1854)

Collection information.

USA: Georgia: Clarke Co.: five individuals from three sites. Caught in flight trap from 19 May–11 August 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Occur almost exclusively under bark, especially of oaks (Thomas 1993); emerged from loblolly pine, water oak, southern red oak, ash, and sweetgum (Gil 2008; Ulyshen and Hanula 2009a, 2010; Ulyshen et al. 2012); may rapidly colonize freshly burned logs (Ulyshen et al. 2010).

Nausibius major Zimmermann, 1869

Collection information.

USA: Georgia (new state record): Clarke Co.: three individuals from two sites. Caught in flight trap from 2 June–14 July 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Emerged from red oak (Quercus rubra L.) (Majka et al. 2008) and eastern white pine (Fall 1931); adults occur in rotting hardwood logs (Dunford and Young 2004; Garrick et al. 2019).

Hymenorus pilosus (Melsheimer, 1846)

Collection information.

Conservation.

Apparently lost after 1925 from the fauna of Plummers Island, Maryland, possibly due to changing forest conditions (Steiner 2008).

Mycetochara fraterna (Say, 1824)

Collection information.

USA: Georgia (new state record*): Clarke Co.: four individuals from four sites. Caught in flight trap from 21 April–2 June 2020.

Distribution.

Eastern United States, transcontinental in Canada.

Saproxylic habits.

Adults occur under bark of aspen (Campbell 1978); species of Mycetochara Berthold, 1827 have been reared from the decaying interior of a living maple tree (McClarin 2008b).

Conservation.

Apparently lost after 1925 from the fauna of Plummers Island, Maryland, possibly due to changing forest conditions (Steiner 2008).

Genus Neomida Latreille, 1829

Collection information.

USA: Georgia: Clarke Co.: a single individual from one site, unable to be identified to species-level due to damage. Caught in flight trap from 9–25 March 2020.

Saproxylic habits.

All life stages occur in various fungi where the larvae develop, and adults also occur under bark of pine and oak (Ciegler 2014).

Platydema ellipticum (Fabricius, 1798)

Collection information.

USA: Georgia: Clarke Co.: a single individual from one site. Caught in flight trap from 25 March–9 April 2020.

Distribution.

Eastern United States.

Saproxylic habits.

Larvae develop within the shelf fungus F. Gilva (Weiss 1919).

Conservation.

Occurrence apparently stable from 1900–present on Plummers Island, Maryland, despite losses of other species possibly due to changing forest conditions (Steiner 2008).

Collection information.

USA: Georgia: Clarke Co.: a single individual from one site. Caught in flight trap from 30 June–14 July 2020.

Distribution.

North America.

Saproxylic habits.

Adults occur under bark, in decaying wood or in hollows of a variety of trees, as well as in or on polypore fungi (Pollock 2012; Ciegler 2014).

Eustrophus tomentosus Say, 1827

Collection information.

USA: Georgia: Clarke Co.: 105 individuals from 23 sites. Caught in flight trap from 25 March–9 September 2020.

Distribution.

North America.

Saproxylic habits.

Adults occur under bark, in decaying wood or in hollows of a variety of trees, in addition to in or on polypore fungi (Pollock 2012; Ciegler 2014).

Conservation.

Rare in the Maritime Provinces of Canada, possibly due to the history of intensive forest management in the region altering forest composition and structure (Majka 2007).

Tetratoma tessellata Melsheimer, 1844

Collection information.

USA: Georgia (new state record*): Clarke Co.: three individuals from two sites. Caught in flight trap from 9 April–6 May 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Occur on fungi growing on dead trees (birch, maple, oak) (Chantal 1985; Majka and Pollock 2006), including a dried oyster mushroom growing on a sugar maple (Webster et al. 2012c).

Throscidae

Aulonothroscus convergens (Horn, 1885)

Collection information.

USA: Georgia: Clarke Co.: 45 individuals from 19 sites. Caught in flight trap from 9 March–9 September 2020.

Distribution.

Eastern United States (Blanchard 1917).

Saproxylic habits.

Emerged from loblolly pine throughout decomposition (Ulyshen and Hanula 2010) and hardwood twigs (Ferro and Nguyen 2016).

Aulonothroscus distans Blanchard, 1917

Collection information.

USA: Georgia (new state record): Clarke Co.: two individuals from two sites. Caught in flight trap from 21 April–6 May 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Emerged in number from dead hardwoods and associated with freshly dead wood (Ferro et al. 2012a).

Conservation.

Significantly higher abundance in secondary (= second-growth) than primary (= old-growth) forests in the southern Appalachian Mountains (Ferro et al. 2012a).

Aulonothroscus parallelus Blanchard, 1917

Collection information.

USA: Georgia (new state record*): Clarke Co.: 25 individuals from 12 sites. Caught in flight trap from 25 March–27 July 2020.

Distribution.

Southeastern United States (previously known from Virginia and unspecified “southern” localities (Blanchard 1917)).

Saproxylic habits.

Unknown; species of Aulonothroscus Horn, 1890 emerged from deadwood and adults occur under bark of dead hardwoods and conifers, including standing dead trees (Howden and Vogt 1951; Ulyshen and Hanula 2010; Ferro et al. 2012a).

Occur under bark of dead oaks (Barron 1971).

Conservation.

Occurrence probability increases in old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2024).

Tenebroides semicylindricus (Horn, 1862)

Collection information.

USA: Georgia: Clarke Co.: a single individual from one site. Caught in flight trap from 14–27 July 2020.

Distribution.

Eastern United States.

Saproxylic habits.

Emerged from water oak, loblolly pine, and sweetgum, and was associated with the crowns of standing dead trees (Ulyshen and Hanula 2009a).

Thymalus marginicollis Chevrolat, 1842

Collection information.

USA: Georgia: Clarke Co.: three individuals from three sites. Caught in flight trap from 21 April–19 May 2020.

Distribution.

Eastern North America, transcontinental in Canada.

Saproxylic habits.

Larvae develop within polypore fungi (e.g., the birch polypore (Fomitopsis betulina (Bull.) B.K. Cui, M.L. Han and Y.C. Dai (Fomitopsidaceae)), and adults occur in association with it as well (Brues 1927; Minch 1952); adults also occur on other fungi, including Daedalea confragosa Berk. (Fomitopsidaceae), Cerrena unicolor (Bull.) Murrill (Cerrenaceae), and Trametes versicolor (L.) Lloyd (Barron 1971).

Zopheridae

Aulonium parallelopipedum (Say, 1826)

Collection information.

USA: Georgia: Clarke Co.: two individuals from two sites. Caught in flight trap from 19 May–16 June 2020.

Distribution.

Eastern United States.

Saproxylic habits.

Emerged from sweetgum and water oak snags (Ulyshen and Hanula 2009a); adults occur on dead logs or under bark, including oak, beech, and elm (Hopkins 1893; Robinson 1918; Hoffmann 1942; Gil 2008); species of Aulonium Erichson, 1845 are likely fungivores that facultatively prey upon bark beetles (Ivie 2002b).

Bitoma quadricollis (Horn, 1885)

Collection information.

Saproxylic habits.

Under bark and in moist, rotting wood of hardwoods (especially oaks and hickory), and occasionally pines (Stephan 1989; Gil 2008); emerged from oaks, loblolly pine, sweetgum logs and hardwood twigs (Ulyshen and Hanula 2009a; Ferro and Nguyen 2016); emerged throughout loblolly pine decomposition and associated with the base and fallen logs of dead pine trees (Ulyshen and Hanula 2009a, 2010).

Conservation.

Significantly associated with young forests (regrown since 1938 and pine dominated) in highly forested landscapes (> 50% forest) in the Piedmont (Traylor et al. 2023a).

Synchita fuliginosa Melsheimer, 1846

Collection information.

USA: Georgia: Clarke Co.: 47 individuals from 26 sites. Caught in flight trap from 21 April–9 September 2020.

Distribution.

Eastern North America.

Saproxylic habits.

Larvae and adults occur under bark and feed on fungal growth, including chestnut blight (Cryphonectria parasitica (Murrill.) Barr (Cryphonectriaceae) (Craighead 1920); emerged from, and occurs under bark of dead elm, maple, oak, pine, hickory, and birch (Savely 1939; Hoffmann 1942; Stephan 1989; Gil 2008), and associated with the wood at the base of dead sweetgums (Ulyshen and Hanula 2009a).

Synchita parvula Guérin-Méneville, 1844

Collection information.

USA: Georgia: Clarke Co.: 567 individuals from 40 sites. Caught in flight trap from 9 March–9 September 2020.

Distribution.

Eastern United States.

Saproxylic habits.

Emerged and collected from oaks (southern red oak and water oak) (Gil 2008; Ulyshen and Hanula 2009a) and associated with dead oaks (Stephan 1989).

Conservation.

significantly associated with old forests (predating 1938 and oak dominated) in the Piedmont (Traylor et al. 2023a).

Discussion

The Coleoptera lists provided here join others in assembling the natural history information of saproxylic invertebrates in the southeastern USA. Hanula (1996) provided a partial list of wood-feeding insects (Blattodea: Termitoidea, Coleoptera, Lepidoptera, and Hymenoptera) for the region, and others assembled similar lists for earthworms (Oligochaeta; Hendrix 1996); mites (Orbatida, Prostigmata, and Mesostigmata; Johnston and Crossley 1996); and millipedes (Diplopoda), centipedes (Chilopoda), and land-snails (various Gastropoda; Caldwell 1996). Snider (1996) compiled an overview of southeastern Collembola occurring in deadwood. Ferro et al. (2012a) provided habitat details and photographs of 71 Coleoptera species associated with various deadwood types and/or forest disturbance categories in Great Smoky Mountains National Park. Ferro et al. (2012a) additionally reviewed the saproxylic literature for the region and wrote a brief history of worldwide saproxylic research. Moreover, published works providing habitat associations and distributional information are available for better known and economically important groups such as Buprestidae (Nelson et al. 2008), Cerambycidae (Lingafelter 2007), and Curculionidae: Scolytinae (Wood 1982). Aside from those referenced above, few resources exist for classifying invertebrate species as saproxylic, which may be a major impediment for deadwood-biodiversity research in the region. Thus, the lists that we present add to the foundation of knowledge about the region’s saproxylic fauna, which will hopefully facilitate further biodiversity research.

An efficient way of gaining knowledge about host range and microhabitat usage for a wide variety of species is emergence experiments, where deadwood pieces are either created or chosen in the field, typically removed from the field after a period of time, and placed into chambers where insects will eventually emerge (methods reviewed in Ferro and Carlton 2011). These types of experiments are useful for inventorying the species using specific types of resources and comparing biodiversity patterns along environmental, spatial, or temporal gradients. For example, studies have examined the saproxylic fauna inhabiting deadwood in response to tree species, decomposition stage, diameter class, vertical stratification, seasonality, sun exposure, flooding, fire exposure, and native/non-native status (Ulyshen and Hanula 2009a, 2010; Ferro et al. 2009, 2012a; Ulyshen et al. 2010, 2012, 2018, 2020; Ferro and Gimmel 2014; Ulyshen 2014; Seibold et al. 2016; Seibold et al. 2018; Vogel et al. 2020; Lettenmaier et al. 2022). Much of the natural history information and species records included in our lists was partially or entirely learned from emergence studies. Such studies are invaluable for providing information about a species’ use of particular habitats. For example, no habitat information was known of the canopy specialist, Tenebroides semicylindricus (Trogossitidae), until it emerged in numbers from dead wood removed from tree crowns (Ulyshen and Hanula 2009a). These studies, however, should be used to complement more thorough investigations of a species’ biology, and documenting the detailed natural history of species should be prioritized when opportunities arise. For example, the fascinating and possibly unique life cycle of Micromalthus debilis (Micromalthidae) would have gone unnoticed were it not for the curiosity and careful study of Barber (1913a, b).

Producing a regional annotated checklist for saproxylic invertebrates would be a great goal for future researchers and would greatly facilitate research in this field. However, we are far from understanding the southeastern fauna well enough to accomplish this, and this is likely the case for other regions as well. From the 293 taxa for which we provide saproxylic habits, 29 species (9.9%) are considered saproxylic solely on taxonomic placement (i.e., no information exists but all congeners with known biology are saproxylic), and 68 species (23.2%) are recorded for the first time in the state of Georgia. Moreover, the extent of natural history known for many species included in our list is limited to a single rearing or emergence event. Thus, very basic biological and distributional information about saproxylic Coleoptera is lacking, and the same applies for other invertebrate groups as well. Filling this information gap is essential for understanding the responses of biodiversity to changing forest conditions and assessing conservation needs.

Acknowledgements

We thank Robert Turnbow and Cecil Smith for early access to their work, An Annotated List of the Coleoptera of Georgia (https://site.caes.uga.edu/ugca/turnbow-and-smith/). We thank two reviewers for their careful reading and comments to improve this manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was funded through Presidential Early Career Award (PECASE) funds awarded to MD Ulyshen and USDA National Institute of Food and Agriculture Hatch project GEO00886 awarded to JV McHugh.

Author contributions

Clayton R. Traylor – Conceptualization, Data curation, Investigation, Methodology, Project administration, Writing – original draft, Writing – review and editing. Michael D Ulyshen – Conceptualization, Funding acquisition, Resources, Writing – review and editing. J. Winston Cornish – Data curation, Investigation, Methodology, Writing – review and editing. Gabriel Tigreros – Data curation, Writing – review and editing. Joseph V. McHugh – Conceptualization, Funding acquisition, Resources, Writing – review and editing.

Author ORCIDs

Clayton R. Traylor https://orcid.org/0000-0001-7798-7198

J. Winston Cornish https://orcid.org/0009-0002-3819-2936

Joseph V. McHugh https://orcid.org/0000-0002-7954-6254

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

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Supplementary material

Supplementary material 1

List of beetle taxa with deadwood associations from experimental studies in the southeastern USA

Clayton R. Traylor, Michael D. Ulyshen, J. Winston Cornish, Gabriel Tigreros, Joseph V. McHugh

Data type: xlsx

Explanation note: Taxa were included if they were collected or emerged from deadwood, or if they were considered saproxylic (see Methods in main text). For taxa collected from deadwood, we provide tree type, collection types (emergent: E, hand-collected: H), and references for each record. For taxa considered saproxylic, we provide references for each study it occurred in. More information on the references can be found in Table 1 of the main text.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

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﻿Abstract

Key words:

﻿Introduction

﻿Methods

﻿Saproxylic definition

﻿List of saproxylic Coleoptera compiled from previous studies

﻿Annotated list of saproxylic Coleoptera

﻿Results

﻿Aderidae

﻿ Cnopus impressus (LeConte, 1875)

Collection information.

Distribution.

Saproxylic habits.

Conservation.

﻿ Ganascus ventricosus (LeConte, 1875)

Collection information.

Distribution.

Saproxylic habits.

﻿ Pseudariotus notatus (LeConte, 1855)

Collection information.

Distribution.

Saproxylic habits.

﻿Anamorphidae

﻿Genus Anamorphus LeConte, 1878

Collection information.

Saproxylic habits.

﻿ Clemmus minor (Crotch, 1873)

Collection information.

Distribution.

Saproxylic habits.

﻿ Micropsephodes lundgreni Leschen and Carlton, 2000

Collection information.

Distribution.

Saproxylic habits.

﻿Anthribidae

﻿ Choragus zimmermanni LeConte, 1876

Collection information.

Distribution.

Saproxylic habits.

﻿ Euparius marmoreus (Olivier, 1795)

Collection information.

Distribution.

Saproxylic habits.

﻿ Eusphyrus walshi LeConte, 1876

Collection information.

Distribution.

Saproxylic habits.

﻿ Euxenus jordani Valentine, 1991

Collection information.

Distribution.

Saproxylic habits.

﻿ Euxenus punctatus LeConte, 1876

Collection information.

Distribution.

Saproxylic habits.

Conservation.

﻿ Goniocloeus bimaculatus (Olivier, 1795)

Collection information.

Distribution.

Saproxylic habits.

﻿ Ischnocerus infuscatus Fåhraeus, 1839

Collection information.

Distribution.

Saproxylic habits.

﻿ Ormiscus saltator LeConte, 1876

Collection information.

Distribution.

Saproxylic habits.

﻿ Piesocorynus moestus (LeConte, 1824)

Collection information.

Distribution.

Saproxylic habits.

﻿ Piesocorynus plagifer Jordan, 1904

Collection information.

Distribution.

Saproxylic habits.

﻿ Toxonotus cornutus (Say, 1831)

Collection information.

Distribution.

Saproxylic habits.

Abstract

Introduction

Methods

Saproxylic definition

List of saproxylic Coleoptera compiled from previous studies

Annotated list of saproxylic Coleoptera

Results

Aderidae

Cnopus impressus (LeConte, 1875)

Ganascus ventricosus (LeConte, 1875)

Pseudariotus notatus (LeConte, 1855)

Anamorphidae

Genus Anamorphus LeConte, 1878

Clemmus minor (Crotch, 1873)

Micropsephodes lundgreni Leschen and Carlton, 2000

Anthribidae

Choragus zimmermanni LeConte, 1876

Euparius marmoreus (Olivier, 1795)

Eusphyrus walshi LeConte, 1876

Euxenus jordani Valentine, 1991

Euxenus punctatus LeConte, 1876

Goniocloeus bimaculatus (Olivier, 1795)

Ischnocerus infuscatus Fåhraeus, 1839

Ormiscus saltator LeConte, 1876

Piesocorynus moestus (LeConte, 1824)

Piesocorynus plagifer Jordan, 1904

Toxonotus cornutus (Say, 1831)

Bostrichidae

Lichenophanes bicornis (Weber, 1801)

Micrapate cristicauda Casey, 1898

Xylobiops basilaris (Say, 1824)

Bothrideridae

Bothrideres cryptus Stephan, 1989

Bothrideres geminatus (Say, 1826)

Carabidae

Mioptachys flavicauda (Say, 1823)

Cerambycidae

Saperda imitans Felt and Joutel, 1904

Cerylonidae

Cerylon castaneum (Say, 1826)

Cerylon unicolor (Ziegler, 1845)

Ciidae

Genus Cis Latreille, 1797

Genus Octotemnus Mellié, 1847

Genus Orthocis Casey, 1898

Cleridae

Chariessa pilosa (Forster, 1781)

Cregya mixta (LeConte, 1866)

Cregya oculata (Say, 1835)

Cymatodera bicolor (Say, 1825)