The Leiodidae (Coleoptera) of Atlantic Canada: new records, faunal composition, and zoogeography

Th e Leiodidae (the round fungus beetles, the small carrion beetles, and the mammal nest beetles) of Atlantic Canada (New Brunswick, Newfoundland and Labrador, Nova Scotia, and Prince Edward Island) are surveyed. Twenty fi ve species, including Colon (Colon) politum Peck and Stephan, Colon (Myloechus) forceps Hatch, Colon (Myloechus) incisum Peck and Stephan, Colon (Myloechus) schwarzi Hatch, Hydnobius arizonensis Horn, Anogdus dissimilis Blatchley, Anogdus potens (Brown), Cyrtusa subtestacea (Gyllenhal), Leiodes puncticollis (Th ompson), Leiodes rufi pes (Gebler), Agathidium atronitens Fall, Agathidium depressum Fall, Agathidium diff orme (LeConte), Agathidium mollinum Fall, Agathidium oniscoides Palisot de Beauvois, Agathidium pulchrum LeConte, Agathidium repentinum Horn, Agathidium rusticum Fall, Gelae parile (Fall), Anisotoma blanchardi (Horn), Anisotoma discolor (Melsheimer), Anisotoma geminata (Horn), Anisotoma globososa Hatch, and Prionochaeta opaca (Say) are newly recorded in Atlantic Canada. One of these, Hydnobius arizonensis, is newly recorded in Canada. Colon (Myloechus) hubbardi Horn is newly recorded in the Maritime Provinces. Eight species are newly recorded in New Brunswick, 29 in Nova Scotia, two on Prince Edward Island, 12 on insular Newfoundland, and fi ve in Labrador for a total of 56 new jurisdictional records. Catops paramericanus Peck and Cook and Catops simplex Say are newly recorded from mainland Nova Scotia, and records are provided to verify the occurrence of Leiodes impersonata Brown and Leiodes punctostriata Kirby in Nova Scotia, and Leptinillus validus (Horn) in insular Newfoundland. Th ree species, Agathidium hatchi Wheeler, Catops americanus Hatch, and Sciodrepoides watsoni (Spence), are removed from the faunal list of New Brunswick. As a result, 66 species of Leiodidae have now been recorded from Atlantic Canada. Th e name Anisotoma obsoleta (Horn) is revalidated while the name Anisotoma horni Wheeler is newly designated a synonym of A. obsoleta. Th e regional composition and zoogeography of the Leiodidae in Atlantic Canada are examined and species are grouped in six main categories, refl ecting their distribution in the region. Island faunas are examined, particularly in regard to the similarities and diff erences of the faunas of Cape Breton Island, insular Newfoundland, and Prince Edward Island. Lone Shieling, in Cape Breton Highlands National RESEARCH ARTICLE Christopher G. Majka, David Langor / ZooKeys 2: 357-402 (2008) 358 Park, is highlighted as an apparent biodiversity “hot spot” for this family within the region. Finally, the saproxylic component of the fauna (19 species in the Agathiidini) is briefl y discussed, particularly in regard to potentially rare species, and the importance of monitoring populations of saproxylic beetles.


Introduction
Th e Leiodidae (the round fungus beetles, the small carrion beetles, and the mammal nest beetles) is a diverse family of beetles that includes species at one time placed in as many as eight separate families.Th e family is ecologically diverse.For example, in Atlantic Canada it includes Agathidium Panzer 1797, Anisotoma Panzer 1797, and Gelae Miller and Wheeler, 2004 that feed on the plasmodia and fruiting bodies of slime molds (and to a lesser degree on certain fungi); Colon Herbst 1797, Hydnobius Schmidt 1847, Leiodes Latreille 1796, and Liocyrtusa Daff ner 1982 that are known or believed to be associated with subterranean fungi; Colenis Erichson 1845 which is associated with decaying soft fungi; Catops Paykull 1798, Prionochaeta Horn 1880, andSciodrepoides Hatch 1933 that are scavengers found in various kinds of moist decaying matter, particularly carrion; Leptinus Müller 1817 and Platypsyllus Ritsema 1869 that are scavengers and ectoparasites associated with aquatic mammals (particularly beavers); and Nemadus Th ompson 1867 which is a scavenger associated with the nests of harvester ants.
In North America Peck (2001) reported 324 species of Leiodidae.Subsequently Miller andWheeler (2004, 2005), Peck andCook (2002, 2007), and Wheeler and Miller (2005) described 32 additional North American species, increasing the total to 356.Peck (1991) recorded 121 species in Canada, 27 of which were known from Atlantic Canada (New Brunswick, Newfoundland and Labrador, Nova Scotia, and Prince Edward Island).Since then Baranowski (1993), Miller and Wheeler (2005), Peck andCook (2002, 2007), and Peck and Stephan (1996) have added additional records, increasing the known fauna of Atlantic Canada to 42 species.Recent studies on forest beetles in the region, and an examination of reference collections, have made it apparent that the fauna of the Atlantic Provinces is considerably more diverse than has hitherto been known.Th e present study adds 56 new jurisdictional records to the regional fauna.Twenty-fi ve species are added to the fauna overall, one of which is newly recorded in Canada.

Methods and Conventions
A total of 2,971 specimens of Leiodidae from Atlantic Canada were examined; 86 from Labrador, 1,211 from insular Newfoundland,183 from New Brunswick,1,488 from Where there are fewer than 40 specimen records, all are given.Where there are more than 40 specimen records, a summary of specimens is provided and the earliest record is given.Where more recent generic treatments and revisions (i.e., Baranowski 1993) for Leiodes; Miller and Wheeler (2005) and Wheeler and Miller (2005) for Agathidium; Peck and Cook (2002) for Catops and Sciodrepoides;and Wheeler (1979) for Anisotoma) have provided detailed distribution maps and specimen records, except for noteworthy specimens, only new records are listed herein.In the case of the genus Colon, Peck and Stephan (1996) did not include range maps and listed specimen records only for newly described species.Similarly for Colenis, specimen records were not provided by Peck (1998) and the distribution map for C. impunctata did not include most of Nova Scotia.For these two genera, records of all specimens examined are provided since almost all are hitherto unpublished.Th e distribution of species in New Brunswick, insular Newfoundland, Nova Scotia, and Prince Edward Island (and occasionally surrounding areas) is shown in the accompanying distribution maps.Records from Labrador (except from the southernmost points) are not shown.Labrador records for most species are shown in the distribution maps in Baranowski (1993) and Peck and Cook (2002).Th e nomenclature, classifi cation, and taxonomy employed follow Peck (2002).

Leiodes contaminabilis
Leiodes contaminabilis was recorded from New Brunswick by Baranowski (1993) based on the above record (Fig. 5).Th e bionomics of the species are completely unknown (Baranowski 1993).Brown, 1932 NOVA SCOTIA: Victoria Co.: 3 km west of Paquets Lake, Cape Breton Highlands National Park, 17.IX.1984,J.M. Campbell and A. Davies, sifting moss, (3, CNC).Although Leiodes impersonata was reported for Nova Scotia by Peck (1991), it was not recorded from the province by Baranowski (1993) in his revision of the genus Leiodes.Th e above record establishes the presence of the species in Nova Scotia.It was recorded from New Brunswick by Baranowski (1993) (Fig. 5).Most species of Leiodes are believed to feed on subterranean fungi.Th e identity of these fungi is unknown (Baranowski 1993).It is found along small roads or trails in deciduous forests, especially with rather sandy (but not dry) soil and rich vegetation (Baranowski 1993).Baranowski Leiodes neglecta is newly recorded from Nova Scotia.It was recorded from New Brunswick by Baranowski (1993) (Fig. 5).A few specimens have been collected in deciduous or mixed forests; otherwise, the bionomics are completely unknown (Baranowski 1993).

Leiodes punctostriata
Leiodes punctostriata was recorded from Labrador, New Brunswick, and Newfoundland by Baranowski (1993) (Fig. 4).Although it was reported for Nova Scotia by Peck (1991), it was not recorded from the province by Baranowski (1993).Th e above records establish its presence in Nova Scotia.It is probably a forest species found in deciduous forests (Baranowski 1993).Th is Holarctic species is newly recorded in Nova Scotia and in Atlantic Canada (Fig. 5).It is found mainly in woodland sites such as forest meadows, along trails and small forest roads, and in shrubbery along riverbanks (Baranowski 1993).Leiodes strigata is newly recorded in Labrador and Newfoundland (Fig. 4).It was found in meadows along small roads or trails in deciduous forests and also in litter along a creek in a mixed forest (Baranowski 1993).
Th is Holarctic species was recorded from New Brunswick by Baranowski (1993) based on the above record (Fig. 5).It apparently prefers rather sandy habitats, e.g., sandy meadows with low vegetation, clearing, forest plantations, or other open country, and is rarely found in forests (Baranowski 1993).

Agathidium diff orme
Agathidium diff orme is newly recorded in New Brunswick, Nova Scotia, and in Atlantic Canada (Fig. 6).It was collected in fi r, pine, spruce, birch, maple, and poplar forests.Host records include the slime molds Stemonitis sp. and a "red, tubular slime mold" (Miller and Wheeler 2005).All the specimens collected in Nova Scotia were found in red spruce forests.Miller and Wheeler, 2005 NEW BRUNSWICK: Northumberland Co.: Tabusintac, 19.VI.1939, 20.VI.1939 Agathidium fawcettae is newly recorded in New Brunswick.It was reported from Nova Scotia by Miller and Wheeler (2005) (Fig. 6).It was collected throughout the year in coniferous and deciduous forests from a wide variety of litter types.In Nova Scotia, it was found in variously aged deciduous forests (red maple, red oak, birch), in coniferous forests (red spruce, black spruce, hemlock, balsam fi r), and in mixed and coastal forests.It was collected with fl ight-intercept traps, pitfall traps, and by hand collecting.Th ere is one record from leaf litter, and one specimen found in a decomposing red spruce log.Recorded hosts include the slime molds Fulgio septica (L.) Wigg., Physarum viride (Bull.)Pers., Leocarpus fragilis (Dicks.)Rost., Hemitrichia clavata (Pers.)Rostaf., and Badhamia sp.(Miller and Wheeler 2005).

Agathidium fawcettae
Prior to the description of this species, several specimens in this series had been identifi ed as Agathidium exiguum Melsheimer.Th ese two species can only be reliably separated on the basis of the features of male genitalia.All the males examined and micro-dissected as part of this study proved to be A. fawcettae.Accordingly, the present authors conclude that there is no evidence that A. exiguum occurs in the region.Miller and Wheeler (2005) reported A. exiguum in North America east to Ontario and north to New Hampshire.Wheeler, 1977 Agathidium hatchi was reported by Miller and Wheeler (2005) from a series of female specimens collected in northwestern Oregon and a single male specimen apparently collected in Frederickton (sic), New Brunswick (20.III.1961,R.C. Clark, Utah State University).Miller and Wheeler (2005: 21) wrote, that "Th e distribution of the species is unusual in that specimens are known from Oregon and New Brunswick, but from nowhere in between.Th e Oregon specimens (including the type) are females and the New Brunswick specimen is a male.Th erefore, there is some possibility the specimens represent diff erent species ...," and "It is also possible that the New Brunswick specimen is mislabeled."

Agathidium hatchi
Th e preponderance of evidence indicates that the record from New Brunswick is erroneous and resulted from a mislabeled specimen.Several reasons indicate this: a) for most of his career R.C. Clark worked on control of balsam wooly adelgid [Adelges piceae (Ratzeburg)] and he never conducted research on Coleoptera (W.Varty, pers.comm.);b) the weather conditions on 20 March, 1961 Fredericton were fully winter-like.Temperatures ranged from -12.8°C (low) to -2.2°C (high) and there was 64 cm of snow on the ground (National Climate Archive 2007), highly improbable circumstances in which to fi nd a slime mold beetle such as Agathidium; c) if R.C. Clark (who worked for the Canadian Forestry Service) had collected any Agathidium specimens incidentally as part of his research, some specimens would have been deposited in the Atlantic Forestry Centre research collection in Fredericton.Th ere are no such specimens in the collection (G.Smith and J. Sweeney, pers.comm.);d) R.C. Clark lived in Fredericton for over three decades and it is highly unlikely that he would misspell the name of the capital city of New Brunswick as "Frederickton"; e) in the 1960s there were no scientifi c exchanges or research programs between Atlantic Forestry Centre and Utah State University (W.Varty, pers.comm.).

Agathidium pulchrum
Anisotoma blanchardi is newly recorded in Nova Scotia and in Atlantic Canada (Fig. 9).It was collected between May and September in log and forest fl oor litter.Slime mold hosts include Stemonitis fusca, Metatricha sp., and Lycogola sp.(Wheeler 1979).

Anisotoma inops
Anisotoma inops is newly recorded from Newfoundland and Nova Scotia (Fig. 8).It was listed as occurring in New Brunswick by Peck (1991).Adults have been found between May and September (Wheeler 1979).In Nova Scotia, it was found almost exclusively in coniferous forests in red spruce, black spruce, hemlock, white pine, and balsam fi r stands.Specimens were found in a decaying red maple log, in polypore fungi on a red spruce log, in polypore fungi on white birch and balsam fi r, in decaying gill fungi, in Lycoperdon sp.fungi, in a decaying red spruce, and in an "orange ball mushroom."In Newfoundland, it was found associated with decaying red pine.
Anisotoma obsoleta (Horn, 1880)  Anisotoma obsoleta is newly recorded from Newfoundland and Nova Scotia (Fig. 8).It was listed as occurring in New Brunswick by Peck (1991).Adults have been found between April and December in stumps and forest litter.Slime mold hosts include Stemonitis axifera, S. fusca, S. splendens, Fuligo septica, and Comatricha nigra (Wheeler 1979).In Nova Scotia, it was collected in red spruce, hemlock, red maple/red oak/ birch, and mixed coastal forests.Wheeler (1979) proposed the name Anisotoma horni for A. obsoleta (Horn) [described by Horn (1880) in the genus Leiodes and later transferred to Anisotoma by Brown (1937b)] in the belief that the name was preoccupied by "Leiodes obsoletus" (Melsheimer, 1844).Melsheimer (1844), however, described the species as Pallodes obsoletus [which was later transferred to Neocyrtusa by Brown (1937a) and then to Anogdus by Daffner (1988)].Pallodes Melsheimer, 1844, was in turn preoccupied by Pallodes Erichson 1843, a genus in the Nitidulidae.Consequently, the two names were not then, and are not now, in the same genus, hence there is no need for a replacement name.Therefore the original name, Anisotoma obsoleta (Horn), must be reinstated as per Article 59.4 of the Code of Zoological Nomenclature (ICZN 1999).Consequently Anisotoma horni Wheeler is designated as a synonym of A. obsoleta.

Cholevinae
Nemadus brachyderus (LeConte, 1863) In his description of the species, LeConte (1863: 25) published the only locality known to him as "Nova Scotia."Th ere is however, no specimen with this locality in the LeConte Collection in Cambridge, Massachusetts.Th is prompted Peck and Cook (2006) to designate a lectotype.Sanders (1964) also reported the species to be common in over-wintering nests of Camponotus herculeanus (Linnaeus) in New Brunswick.However, Peck and Cook (2006) were unable to examine specimens to confi rm these records.Consequently, the status of this species in both New Brunswick and Nova Scotia should be further investigated.It is a myrmecophile associated with nests of Camponotus noveboracensis (Fitch) and Camponotus herculeanus (Peck and Cook 2006).Fall, 1937 NOVA SCOTIA: Cape Breton Co.: Sydney, UCCB campus, 15.IX.1996,J. N. MacGillivray, (1, NSMC).

Nemadus integer
Nemadus integer was recorded from Nova Scotia by Peck and Cook (2006) based on the above record (Fig. 9).It is a myrmecophile associated with nests of Formica integra Nylander (Peck and Cook 2006).Jeannel, 1936 NOVA SCOTIA: Halifax Co.: Pockwock Lake, 15-30.VI.1997,D.J. Bishop, red spruce forest, fl ight-intercept trap, (1, NSMC).Catops basilaris was reported by Peck and Cook (2002) from Labrador, Newfoundland, New Brunswick, and Nova Scotia including Cape Breton Island (Fig. 11).It is the most widespread and frequently collected species of Catops in North America, primarily collected in forested environments at carrion, but also occasionally on decaying mushrooms, in forest litter, in mammal and wasp nests, and on owl pellets.It is found in deciduous, mixed, and coniferous forests and Sphagnum bog habitats (Peck and Cook 2002).In Nova Scotia, specimens have been collected in coniferous, deciduous, mixed, and coastal forests.(Horn, 1880) Catops egenus was reported from central Labrador by Peck and Cook (2002).It was collected at animal carcasses, on dung, and on decaying mushrooms in coniferous or mixed forests (Peck and Cook 2002).Catops gratiosus was reported from mainland Nova Scotia and Cape Breton Island by Peck and Cook (2002) (Fig 10).It is an infrequently collected species.Most of the collections in the southeastern United States are from caves.Specimens collected from northern or upper elevation areas are from deciduous forests and, in smaller numbers, from coniferous forests and old fi elds where they are found on carrion, decaying fungi, and on mammal dung (Peck and Cook 2002).

Catops luridipennis Mannerheim, 1853
Th is Holarctic species was reported from central Labrador by Peck and Cook (2002).It has been collected on animal carcasses and dung in coniferous forests (Peck and Cook 2002).

Sciodrepoides watsoni
Th is Holarctic species was reported by Peck and Cook (2002) from Newfoundland and New Brunswick (Fig. 12).Th e source of the New Brunswick record (from the Charlotte County area) could not be determined (S.Peck, pers. comm.), nor were vouchers specimens found in any collection consulted.Consequently it is removed from the New Brunswick faunal list.Its status in Newfoundland should also be further investigated since there are no records in the province apart from the specimens listed above.Sciodrepoides watsoni has primarily been collected in forested habitats but it is also tolerant of open old fi elds and grasslands where it is found on carrion and in mammal nests (Peck and Cook 2002).

Platypsyllus castoris
Th is Holarctic species is listed as occurring in Newfoundland by Peck (1991) (Fig. 12).It is associated with beaver as an ectoparasite through their range in North America and Eurasia (Wood 1965).

Regional Composition and Zoogeography
With 25 species of leiodids being newly recorded in Atlantic Canada, and 56 new jurisdictional records reported, it is clear that the knowledge of the composition of the fauna in the region is still incomplete.In particular, few leiodids have been recorded in Prince Edward Island and the leiodid fauna of northern New Brunswick is still poorly known.Nonetheless, some observations on the zoogeography of the fauna in the region are possible, and some distributional patterns are evident.
1) Widespread -Th ere are 17 species that are apparently relatively widespread throughout Atlantic Canada.Th ese include Colon magnicolle, C. hubbardi, Hydnobius substriatus, Leiodes assimilis, L. collaris, L. punctostriata, L. valida, Agathidium mollinum, A. pulchrum, Anisotoma basalis, A. inops, A. obsoleta, Catops a. alsiosus, C. basilaris, C. simplex, Sciodrepoides terminans, and Leptinillus validus.Records from Prince Edward Island are lacking for most of these species; L. collaris and L. validus have not been recorded in Nova Scotia; C. hubbardi and H. substriatus have not been recorded in New Brunswick; and three species (L. valida, A. pulchrum, and L. validus) have only been recorded from a small number of localities.All the species, however, have been found in both the Maritime Provinces and (except for Catops a. alsiosus) insular Newfoundland, and eight have been recorded from Labrador.
2) Maritime Provinces -Th irteen species have been found in the Maritime Provinces that have not been recorded from insular Newfoundland or Labrador.Th ese include Colon asperatum, C. boreale, Leiodes impersonata, L. neglecta, Colenis impunctata, Agathidium atronitens, A. diff orme, A. fawcettae, A. oniscoides, A. rubellum, Nemadus brachyderus, Catops paramericanus, and Prionochaeta opaca.Most of these species have not been found on Prince Edward Island; three species (L.impersonata, C. impunctata, and A. diff orme) have not been recorded on Cape Breton Island; and six species (C. boreale, L. impersonata, L. neglecta, A. atronitens, A. oniscoides, and N. brachyderus) have only been recorded from a small number of localities.Th e pattern of their distribution in the Maritimes appears to be relatively widespread.
3) Newfoundland -Th ree species have been recorded in the region solely from insular Newfoundland: Agathidium repentinum, Platypsyllus castoris, and Anisotoma globososa.Agathidium repentinum and P. castoris (a Holarctic species) are broadly distributed boreal species in Canada and Alaska (Miller and Wheeler 2005;Peck 1991) that may not be present in the Maritime Provinces.Anisotoma globososa is widely distributed in southern Ontario and Quebec and the northern United States from Wisconsin to New Hampshire (Wheeler 1979) and may be present in the Maritime Provinces.Additionally, Sciodrepoides watsoni has only been recorded from Newfoundland (and possibly New Brunswick).Th e status of this species in the region requires further verifi cation and investigation.4) Labrador -Four species have only been recorded in Labrador: Colon politum, Catops egenus, C. luridipennis, and C. luteipes.Except for C. egenus, the other three are

Island Biogeography
Th e number of species of leiodids recorded on Prince Edward Island (two, or 3% of the regional fauna) is extraordinarily low.In comparison, 140 of the 332 native species (42%) of Carabidae (perhaps the best-studied family of Coleoptera in Atlantic Canada) are known from Prince Edward Island (Table 2; Majka et al. 2007b).Although the proportion of native species found on Prince Edward Island varies from family to family, it nonetheless appears that either the leiodid fauna of the island is comparatively much smaller than expected, or that collection eff ort on Prince Edward Island has been insuffi cient to fully determine its leiodid composition, or a combination of both factors.In further comparison, the proportional composition of native leiodids and carabids are similar in the case of Cape Breton Island, Labrador, Nova Scotia, and the Maritime Provinces as a whole (Table 2).Th e proportional composition of leiodids known to occur on insular Newfoundland (37.9%), and in New Brunswick (50.0%) are somewhat smaller in comparison to the proportional representation of carabids in these two jurisdictions (44.6% and 75.0%respectively).Th is is likely also due to insuffi cient collecting eff ort in both areas.Th e proportion of leiodids known from the mainland of Nova Scotia (60.6%) is also somewhat lower.
In the Maritime Provinces the proportion of the leiodid fauna that is Holarctic varies between 8-9% [7-8% in the case of the Carabidae (Majka et al. 2007b)], but on insular Newfoundland the proportion is 13.6% [21.2% in the case of the Carabidae (Majka et al. 2007b)], and in Labrador is 27.3% [36.2% in the case of the Carabidae (Bousquet and Larochelle 1993)].Th e greater proportion of Holarctic species, particularly in Labrador, clearly refl ects its greater proximity to circumboreal environments and landmasses such as the Canadian arctic, Greenland, and Iceland.Th e low proportion of Holarctic leiodids recorded on Cape Breton Island (2.7%) is notable given that the corresponding proportion of Holarctic Carabidae is 12.4% (Table 2 (Majka and McCorquodale 2006;Majka et al. 2007aMajka et al. , 2007b)).In the case of the Leiodidae, the comparative lack of attention with respect to this family in general, their secretive habits, the specialized habitats that some species occupy, and the dearth of collection eff orts in some areas, means that any conclusions about island faunas in the region are premature.

Biodiversity "Hot Spots"
Particularly noteworthy for the region are the large number of species of leiodids collected at Lone Shieling in Cape Breton Highlands National Park, an apparent biodiversity "hot spot" for this family in the region.Twenty-nine species (57% of the Nova Scotia fauna) were collected there, and 36 species (71% of the Nova Scotia fauna) have been found in the Park as a whole, thanks to survey eff orts summarized by Lafontaine et al. (1987).Although the collecting eff ort at this site was outstanding (a dozen experienced researchers from the Biosystematics Research Center in Ottawa, Canada participated over the course of the 1983-1984 fi eld seasons) the site also happens to be within a 350+ year old sugar maple stand which is one of the largest and most pristine old-growth forest stands in the Maritime Provinces (Greenidge 1961).Other rarely collected species of Coleoptera such as Acrotrichis cognata (Matthews) (Ptiliidae) (Majka and Sörensson 2007) and Epuraea parsoni Connell (Nitidulidae) ( Majka and Cline (2006) have been collected at this site as have several species of rare, localized, and disjunct small mammals (Roscoe and Majka 1976).It is possible that the undisturbed old-growth conditions at this site support a large diversity of species that have otherwise been reduced in many areas of Atlantic Canada by the history of forest management practices (Majka 2007).
In comparison, a similar research program in Kouchibouguac National Park in New Brunswick in 1977 recorded only 10 species of leiodids and a more limited survey in 2004-2005 in Kejimkujiuk National Park in Nova Scotia found only seven species of leiodids (C.G.Majka and T. Rossolimo, unpublished data).Th us, the species diversity of the Leiodidae in Cape Breton Highlands National Park appears to be very high, a topic deserving further research.In this regard, it is fortunate that the Lone Shieling site lies within Cape Breton Highlands National Park where it is aff orded a high degree of protection.

Saproxylic Species Diversity and Forest Management
Th e 19 species of leiodids in the Agathiidini are all forest species associated with fungi and/or slime molds and members of the saproxylic invertebrate community, i.e., those organisms that are dependent during some portion of their life cycle upon the dead or dying wood of moribund or dead trees (standing or fallen), upon wood-inhabiting fungi or upon the presence of other saproxylics (Speight 1989).Speight (1989), Grove (2002), and Dudley and Vallauri (2004) are three of many studies that have examined the importance of saproxylic insects in the dynamics of forest ecosystems.In general, upwards of 30% of plant biomass produced annually in forests is in the form of woody tissue and the quantity of plant nutrients recycled annually by saproxylics in forests is roughly 50% of that recycled from the annual leaf fall (Speight 1989).A number of studies have drawn attention to the importance of this group in the trophic dynamics of forests.Siitonen (2001) found that 20-25% of all forest-dwelling invertebrates in Fennoscandia were saproxylic; Martikainen et al. (2000) found that 42% of the 553 species of beetles collected in a spruce forest in Finland were saproxylic ;and Köhler (2000) considered 56% of all forest-dwelling beetle species in forests in the north Rhineland to be saproxylic.Majka (2007) identifi ed 59 "apparently rare" species of saproxylic beetles (collected from ≤ 5 specimens representing ≤ 0.005% of specimens examined in the region.)Th is represented 21% of the 283 species in the 18 saproxylic families, subfamilies, and tribes that were investigated.On that basis, six of the 19 species (32%) including Agathidium depressum, A. oniscoides, A. pulchrum, A. rusticum, Anisotoma errans, and A. globososa would qualify as "apparently rare" (i.e., ≤ 5 specimens have been collected in the region).Th is result is, in all probability due in large measure to the dearth of collecting for many species of leiodids.An increased collecting eff ort is required to discern if some of these species are actually rare, and hence in need of special attention or conservation measures.

Summary
Sixty-six species of Leiodidae are now known from Atlantic Canada, substantially more than the 27 recorded by Peck (1991).Others, undoubtedly, remain to be discovered since many species are secretive, small and inconspicuous, and are poorly sampled by conventional collecting techniques.A better understanding of this family and other forest and saproxylic beetles will add to our understanding of the ecological well-being of our forests.Such baseline faunistic knowledge will, in turn, help us understand how forests and beetles have been aff ected by historical forest management practices, and will help us monitor potential future changes in response to anthropogenic factors such as climate change.

Fig. 11 .
Fig. 11.Distribution of Catops basilaris and Catops simplex in Atlantic Canada.Some locations in Labrador are not shown.

Table 1 .
Leiodidae of the Atlantic Provinces of Canada

Table 2 .
Geographic composition of Atlantic Canada Leiodidae: number and proportion of species .2007b).As the Leiodidae of Cape Breton Island is relatively well surveyed, the low proportion of Holarctic species would appear to be real rather than a collecting artifact.Th e reason for this pattern is unknown.Several studies of Coleoptera in Atlantic Canada have noted an island-related diminution of fauna on Prince Edward Island, Cape Breton Island, and insular Newfoundland 1, Carabidae numbers adapted fromMajka et al. (2007)by the inclusion of Labrador data.al