ISSN 1911-2173

A product of the Biological Survey of Canada & the Entomological Society of Canada

The Cryptophagidae of Canada and the northern United States of America

Georges Pelletier1

Christian Hébert

Natural Resource Canada, Canadian Forest Service, Laurentian Forestry Centre, Sainte-Foy, Québec City, Québec, Canada
1Corresponding author ([email protected])

January 28, 2020: An Editorial Corrigendum has been published for this paper doi:10.3752/cjai.2019.40ed.
March 5, 2020: An Addendum & Corrigendum has been published for this paper doi:10.3752/cjai.2019.40adco.

Abstract

This review of Cryptophagidae of Canada and the Northern United States (US) covers 142 species among the following genera: Caenoscelis (6 species), Antherophagus (4), Cryptophagus (33), Henoticus (4), Henotiderus (2), Micrambe (1), Myrmedophila (1), Pteryngium (1), Salebius (2), Telmatophilus (2), Hypocoprus (1), Atomaria (81), Curelius (1), Ephistemus (2) and Tisactia (1). All species are described morphologically and descriptions include high-definition photographs, geographic distribution data and maps as well as data on seasonal occurrence, habitats and the number of specimens examined. Some already described species that have not yet been recorded in Canada, but that have been found in bordering states with similar habitats, are also included in this publication. An interactive identification key is provided to identify all known species in Canada and Alaska together with a descriptive page for each species.

This publication includes many taxonomic changes, mainly new synonymies. Among Cryptophaginae, the following are considered as new synonyms: Caenoscelis ovipennis Casey, C. macra Casey and C. elongata Casey = C. basalis Casey; Caenoscelis obscura Casey and C. subfuscata Casey = C. cryptophaga Reitter; C. ochreosa Casey = C. ferruginea (C.R. Sahlberg); Caenoscelis angusticollis Casey and C. shastanica Casey = C. parallela Casey; C. macilenta Casey = C. subdeplanata Brisout de Barneville; Cryptophagus pilosus sensu auctorum = C. punctipennis Brisout de Barneville, the true C. pilosus being a nomen dubium. Henoticus mycetoecus is now a synonym of H. pilifer Reitter. Salebius lictor Casey and S. montanus Casey = S. octodentatus (Mäklin).

Most new synonymies are within the genus Atomaria and are as follows: Atomaria (Anchicera) nebulosa Casey = At. (Anc.) apicalis Erichson; At. (Anc.) divisa Casey = At. (Anc.) distincta Casey; At. (Anc.) gonodera Casey, At. (Anc.) riparia Casey, At. (Anc.) subalutacea Casey, At. (Anc.) incerta Casey and At. (Anc.) crypta Casey = At. (Anc.) fuscata Schöenherr; At. (Anc.) ebenina Casey = At. (Anc.) morio Kolenati; At. (Anc.) fallax Casey = At. (Anc.) pusilla (Paykull); At. (Anc.) nanula Casey = At. (Anc.) turgida Ercihson; At. (At.) luculenta (Casey), At. (At.) lucida (Casey) and At. (At.) lucens Grouvelle = At. (At.) nigrirostris Stephens; At (At.) strandi Johnson = At. (At.) coloradensis (Casey); At. (At.) parvicollis (Casey) and A. parva Schenkling = At. (At.) nigricollis (Casey); At. (At.) ochronitens (Casey) = At. (At.) stricticollis (Casey); At. (At.) subangulata (C. Sahlberg) = At. (At.) subdentata (Casey); At. (At.) melas (Casey) = At. (At.) tenebrosa (Casey); At. (At.) cribripennis (Casey) and At. (At.) castanea (Casey) = At. (At.) undulata (Casey). At. (At.) lepidula Mäklin is now considered a nomen dubium, the holotype being lost. Finally, Ephistemus perminutus Casey is a synonym of Curelius japonicus Reitter. One species is restored as a valid species: Atomaria (At.) lineola (Notman), formally synonymized by Johnson (2007) as At. (At.) wollastoni. A. carinula (Casey) was named by Ganglbauer in 1889 and is assigned to a new name: A. neocarinula Pelletier.

This publication includes the descriptions of 22 new species, mostly belonging to the genus Atomaria (provinces or states of types in parentheses): Henoticus varidentatus Pelletier (QC), Atomaria (Anchicera) arcuaticollis Pelletier (QC, AK), At. (Anc.) hudsonica Pelletier (MB), At. (Anc.) neomunda Pelletier (QC, AK), At. (Anc.) nigrirostrisoides Pelletier (QC, AK), At. (Anc.) nigritaria Pelletier (MB, QC), At. (Anc.) ocularia Pelletier (AB), At. (Anc.) parallelicollis Pelletier (QC), At. (Anc.) salicicola Pelletier (AB), At. (At.) albertana Pelletier (AB), At. (At.) calidaria Pelletier (AB), At. (At.) californica Pelletier (CA), At. (At.) cribricollis Pelletier (OR, AB, AK), At. (At.) impressicollis Pelletier (AB, BC, YT), At. (At.) lyricollis Pelletier (MB), At. (At.) ontariensis Pelletier (ON), At. (At.) orbicollis Pelletier (QC, ON), At. (At.) pinicola Pelletier (NB, QC, BC), At. (At.) pseudotsugae Pelletier (BC), At. (At.) puelloides Pelletier (GA), At. (At.) thujaplicatae Pelletier (BC) and At. (At.) visoni Pelletier (AB, BC).

Two genera are recorded for the first time in Canada: Micrambe Thomson, with Micrambe ulicis (Stephens) (QC) (also recorded for the first time in North America) and Tisactia Casey, with Tisactia subglabra Casey (QC, ON).

Including all new species except for the two known only from California and Georgia, 62 species are recorded for the first time in Canada or Alaska. In addition to the new species mentioned above, these species are: Cryptophagus croceus Zimm.(NS, QC, ON, MB, SK, AB), C. denticulatus Heer(QC, ON, MB, AB, BC), C. lycoperdi (Scop.) (QC), C. valens Csy. (QC, ON, MB, SK, BC), Henoticus pilifer Reitter (QC, ON, BC), Hypocoprus latridioides Motsch. (SK, AB), Atomaria (Anchicera) atra (Hrbst.) (NB, QC), A. basalis Eric. (NB, QC, MB, BC, NT, YT, AK), A. brevicollis Csy. (NB, QC, ON, MB, SK, AB, BC, NT), A. gilvipennis Csy.(NS, QC, ON), A. inepta Eric. (QC, BC, NT, AK), A. mesomela Hrbst. (BC), A. morio Kolen.(NB, QC, ON, SK, AB), A. munda Eric. (NB, QC, BC, AK), A. oblongula Csy. (NF, NS, NB, QC, ON, MB, SK, AB, BC, NT, AK), A. ornata Heer (BC), A. peltata Kraatz (BC, AK), A. postpallens Csy. (AB, BC, NT, AK), A. sodermani Sjöb. (QC, NT), A. turgida Eric. (NB, QC, ON, MB, SK, AB, BC, NT, YT, AK), Atomaria (Atomaria) affinis R.F. Sahl. (NS, NB, QC, AB, BC, AK), A. alpina Heer (NS, NB, QC, ON, AB, BC), A. coloradensis (Csy.) (NB, QC, AB, BC, YT, AK), A. constricta (Csy.) (NB, QC, AB, BC), A. crassula (Csy.) (NB, QC), A. dispersa (Csy.) (NB, QC, ON, AB, BC, AK), A. elongatula Eric. (QC, AB, BC), A. fimetarii (F.) (NS, NB, QC, ON, MB, SK, AB, BC, NT), A. impressa Eric. (NB, QC, ON, AB), A. macra (Csy.) (NB, QC, AB, BC, AK), A. neocarinula (Pel.) (AB), A. nigricollis (Csy.) (ON, MB, AB, BC, YT, AK), A. parviceps Notm. (QC, ON, AB), A. patens (Csy.) (NB, QC, ON), A. puella (Csy.) (AB), A. pumilio (Csy.) (NB, QC, ON, MB, SK, BC), A. tenebrosa (Csy.) (NB, QC, ON, AB, AK), A. umbrina (Gyll.) (NB, QC, ON, BC), A. undulata (Csy.) (NB, QC, ON, AB, BC, AK) and A. versa Grouv. (AK).

Excluding all the species mentioned previously, there is at least one new provincial or Alaska state record for the following 61 species: Caenoscelis basalis Csy.(NB, MB), C. ferruginea (C.R. Sahl,) (NS), C. parallela Csy. (AB), C. subdeplanata Bris. de Barn. (NB, BC), Antherophagus convexulus LeC. (SK, AB, AK), A. ochraceus Melsh. (NT), A. pallidivestis Csy. (NB, QC, ON, AB), A. suturalis Mäkl. (SK, AB), Cryptophagus bidentatus Mäkl. (QC, ON, MB, AB, YT), C. cellaris (Scop.) (QC, YT), C. confertus Csy. (ON, MB, YT), C. corticinus Thoms. (MB, AB, YT), C. dentatus (Hrbst.) (QC, ON, MB, YT), C. difficilis Csy. (NB, ON, MB, NT, YT, AK), C. distinguendus (QC, ON, SK, AB), C. fallax Balf.-Br. (QC, ON), C. hebes Csy. (YT), C. histricus Csy.(QC, ON, SK, AB), C. lapponicus Gyll. (QC, ON, AB, NT, YT, AK), C. laticollis Lucas (MB), C. mainensis Csy. (QC, ON), C. obsoletus Reit. (QC, MB, SK, AB, YT), C. peregrinus Wdrf. & Cmbs. (AB), C. quadrihamatus Mäkl. (QC, ON, MB, SK, AB, BC, YT), C. saginatus Sturm (PE, NB, QC), C. scanicus (QC, ON, BC, YT), C. scutellatus Newm. (QC, MB, AB), C. setulosus Sturm (AB, NT), C. stromus Wdrf. & Cmbs. (AB, AK), C. subfumatus Kraatz (NB, MB), C. tuberculosus Mäkl.(QC, ON, MB), C. varus Wdrf. & Cmbs. (QC, ON, MB), Henoticus serratus (AB), Henotiderus lorna (Hatch) (AB), Myrmedophila americana (LeC.) (ON), Pteryngium crenulatum (Eric.) (ON, AB, AK), Salebius octodentatus (Mäkl.) (AB), Atomaria (Anchicera) aleutica Csy. (MB, SK, AB, BC, NT, YT), A. apicalis Eric. (ON, SK, AB, BC, NT, YT, AK), A. distincta Csy. (ON, AB), A. ephippiata Zimm. (AK), A. fulvipennis Mann. (QC, ON, MB, SK, AB, BC, NT, YT), A. fuscata Schnrr. (AB, BC, NT, YT, AK), A. kamtschatica Motsch. (MB, AB, NT, YT), A. laetula LeC. (SK), A. lederi Johns. (NB, QC, ON, AB, BC), A. lewisi Reit. (MB, SK, AB, BC, AK), A. nubipennis Csy. (AB, AK), A. pusilla (Payk.) (ON), A. testacea Steph. (SK, BC), Atomaria (Atomaria) atrata Reit. (QC, ON, AB, AK), A. capitata (Csy.) (NB, QC, AB, AK), A. lineola (Notman) (ON, MB, SK, AB, BC), A. longipennis (Csy.) (QC, AB, AK), A. nigrirostris Steph. (NB, ON, SK, AB, BC, YT), A. planulata Mäkl. (QC, AB, YT), A. quadricollis (Csy.) (QC, ON, AB), A. stricticollis (Csy.) (QC, AB, YT), A. subdentata (Csy.) (NB, QC, ON, MB, SK, AB, AK), A. vespertina Mäkl. (QC, ON, MB, AB) and Ephistemus globulus (NF, LB, NS, QC).

By state or province, these new records total 2 for Newfoundland and Labrador, 1 for Prince-Edward Island, 9 for Nova Scotia, 31 for New Brunswick, 66 for Quebec, 50 for Ontario, 33 for Manitoba, 24 for Saskatchewan, 69 for Alberta, 41 for British Columbia, 17 for the Northwest Territories and Nunavut, 24 for Yukon, and 35 for Alaska.

The following 27 species are considered Holarctic, being naturally distributed in northern Europe, Asia and North America, through Alaska: Caenoscelis ferruginea, Cryptophagus acutangulus, C. bidentatus, C. confertus, C. corticinus, C. lapponicus, C. quadrihamatus, C. tuberculosus, Henoticus serratus, Henotiderus centromaculatus, Salebius octodentatus, Atomaria (Anchicera) apicalis, A. (Anc.) basalis, A. (Anc.) fulvipennis, A. (Anc.) fuscata, A. (Anc.) kamtschatica, A. (Anc.) peltata, A. (Anc.) sodermani, A. (Anc.) turgida, A. (Atomaria) affinis, A. (At.) atrata, A. (At.) elongatula, A. (At.) fimetarii, A. (At.) nigrirostris, A. (At.) subdentata, A. (At.) vespertina and Hypocoprus latridioides.

Cryptophagidae contains up to 34 adventive species accidentally introduced by human activity, many of which live indoors, but also includes some that have successfully colonized natural habitats: Caenoscelis subdeplanata, Cryptophagus cellaris, C. dentatus, C. denticulatus, C. distinguendus, C. fallax, C. laticollis, C. lycoperdi, C. obsoletus, C. punctipennis, C. saginatus, C. scanicus, C. scutellatus, C. setulosus, C. subfumatus, Henoticus pilifer, Micrambe ulicis, Pteryngium crenulatum, Telmatophilus typhae, Atomaria (Anchicera) atra, A (Anc.) lederi, A (Anc.) lewisi, A (Anc.) mesomela, A (Anc.) morio, A (Anc.) munda, A (Anc.) ornata, A (Anc.) pusilla, A (Anc.) testacea, A. (Atomaria) alpina, A. (At.) impressa, A. (At.) linearis, A. (At.) umbrina, Curelius japonicus and Ephistemus globulus.

Résumé

Cette révision des Cryptophagidés du Canada et du nord des États-Unis couvre 142 espèces parmi les genres suivants : Caenoscelis (6 espèces), Antherophagus (4), Cryptophagus (33), Henoticus (4), Henotiderus (2), Micrambe (1), Myrmedophila (1), Pteryngium (1), Salebius (2), Telmatophilus (2), Hypocoprus (1), Atomaria (81), Curelius (1), Ephistemus (2) et Tisactia (1). Toutes ces espèces sont décrites morphologiquement et les descriptions incluent, pour chaque espèce, des photographies à haute définition, des données de répartition géographique avec carte de distribution, ainsi que des données sur l’occurrence saisonnière, les habitats et le nombre de spécimens observés. Quelques espèces n’ayant pas encore été récoltées au Canada, mais vivant dans des habitats similaires aux États-Unis, ont également été incluses dans cette publication. Une clé virtuelle d’identification permet d’identifier toutes les espèces connues au Canada et en Alaska, avec un lien vers leur page descriptive.

Cette publication inclut de nombreux changements taxonomiques, surtout au niveau synonymique. Les espèces suivantes ont été synonymisées: Caenoscelis ovipennis Casey, C. macra Casey and C. elongata Casey = C. basalis Casey; Caenoscelis obscura Casey and C. subfuscata Casey = C. cryptophaga Reitter; C. ochreosa Casey = C. ferruginea (C.R. Sahlberg); Caenoscelis angusticollis Casey and C. shastanica Casey = C. parallela Casey; C. macilenta Casey = C. subdeplanata Brisout de Barneville; Cryptophagus pilosus sensu auctorum = C. punctipennis Brisout de Barneville, le véritable C. pilosus ayant une identité incertaine. Henoticus mycetoecus est maintenant un synonyme de H. pilifer Reitter. Salebius lictor Casey et S. montanus Casey = S. octodentatus (Mäklin).

La plupart de la nouvelle synonymie se trouve dans le genre Atomaria et inclut les espèces suivantes : Atomaria (Anchicera) nebulosa Casey = At. (Anc.) apicalis Erichson; At. (Anc.) divisa Casey = At. (Anc.) distincta Casey; At. (Anc.) gonodera Casey, At. (Anc.) riparia Casey, At. (Anc.) subalutacea Casey, At. (Anc.) incerta Casey et At. (Anc.) crypta Casey = At. (Anc.) fuscata Schöenherr; At. (Anc.) ebenina Casey = At. (Anc.) morio Kolenati; At. (Anc.) fallax Casey = At. (Anc.) pusilla (Paykull); At. (Anc.) nanula Casey = At. (Anc.) inepta Casey; At. (At.) luculenta (Casey), At. (At.) lucida (Casey) et At. (At.) lucens Grouvelle = At. (At.) nigrirostris Stephens; At (At.) strandi Johnson = At. (At.) coloradensis (Casey); At. (At.) parvicollis (Casey) et A. parva Schenkling = At. (At.) nigricollis (Casey); At. (At.) ochronitens (Casey) = At. (At.) stricticollis (Casey); At. (At.) subangulata (C. Sahlberg) = At. (At.) subdentata (Casey); At. (At.) melas (Casey) = At. (At.) tenebrosa (Casey); At. (At.) cribripennis (Casey) and At. (At.) castanea (Casey) = At. (At.) undulata (Casey). At. (At.) lepidula Mäklin est maintenant considéré comme un nomem dubium, l’holotype ayant été perdu. Finalement, Ephistemus perminutus Casey est un synonyme de Curelius japonicus Reitter. L’analyse génétique a permi de restaurer 2 espèces valides que Johnson (2007) avait synonymisées comme étant At. (At.) wollastoni: Atomaria (At.) pumilio et At. (At.) lineola (Notman). A. carinula (Casey) a été auparavant nommé par Ganglbauer en 1889 et un nouveau nom lui a été assigné: A. neocarinula Pelletier.

Cette publication comprend la description de 22 nouvelles espèces, appartenant surtout au genre Atomaria (les provinces et les États sont inclus entre parenthèses): Henoticus varidentatus Pelletier (QC), Atomaria (Anchicera) arcuaticollis Pelletier (QC, AK), At. (Anc.) hudsonica Pelletier (MB), At. (Anc.) neomunda Pelletier (QC, AK), At. (Anc.) nigrirostrisoides Pelletier (QC, AK), At. (Anc.) nigritaria Pelletier (MB, QC), At. (Anc.) ocularia Pelletier (AB), At. (Anc.) parallelicollis Pelletier (QC), At. (Anc.) salicicola Pelletier (AB), At. (At.) albertana Pelletier (AB), At. (At.) calidaria Pelletier (AB), At. (At.) californica Pelletier (CA), At. (At.) cribricollis Pelletier (OR, AB, AK), At. (At.) impressicollis Pelletier (AB, BC, YT), At. (At.) lyricollis Pelletier (MB), At. (At.) ontariensis Pelletier (ON), At. (At.) orbicollis Pelletier (QC, ON), At. (At.) pinicola Pelletier (NB, QC, BC), At. (At.) pseudotsugae Pelletier (BC), At. (At.) puelloides Pelletier (GA), At. (At.) thujaplicatae Pelletier (BC) and At. (At.) visoni Pelletier (AB, BC).

Deux nouveaux genres ont été récoltés pour la première fois au Canada : Micrambe Thomson, incluant Micrambe ulicis (Stephens) (QC) (aussi enregistré pour la première fois en Amérique du Nord) et Tisactia Casey, incluant Tisactia subglabra Casey (QC, ON).

En incluant toutes les nouvelles espèces, 60 espèces sont mentionnées pour la première fois au Canada ou en Alaska. En voici la liste, excluant les deux mentionnées au paragraphe précédent : Cryptophagus croceus Zimm.(NS, QC, ON, MB, SK, AB), C. denticulatus Heer(QC, ON, MB, AB, BC), C. lycoperdi (Scop.) (QC), C. valens Csy. (QC, ON, MB, SK, BC), Henoticus pilifer Reitter (QC, ON, BC), Hypocoprus latridioides Motsch. (SK, AB), Atomaria (Anchicera) atra (Hrbst.) (NB, QC), A. basalis Eric. (NB, QC, MB, BC, NT, YT, AK), A. brevicollis Csy. (NB, QC, ON, MB, SK, AB, BC, NT), A. gilvipennis Csy.(NS, QC, ON), A. inepta Eric. (QC, BC, NT, AK), A. mesomela Hrbst. (BC), A. morio Kolen.(NB, QC, ON, SK, AB), A. munda Eric. (NB, QC, BC, AK), A. oblongula Csy. (NF, NS, NB, QC, ON, MB, SK, AB, BC, NT, AK), A. ornata Heer (BC), A. peltata Kraatz (BC, AK), A. postpallens Csy. (AB, BC, NT, AK), A. sodermani Sjöb. (QC, NT), A. turgida Eric. (NB, QC, ON, MB, SK, AB, BC, NT, YT, AK), Atomaria (Atomaria) affinis R.F. Sahl. (NS, NB, QC, AB, BC, AK), A. alpina Heer (NS, NB, QC, ON, AB, BC), A. coloradensis (Csy.) (NB, QC, AB, BC, YT, AK), A. constricta (Csy.) (NB, QC, AB, BC), A. crassula (Csy.) (NB, QC), A. dispersa (Csy.) (NB, QC, ON, AB, BC, AK), A. elongatula Eric. (QC, AB, BC), A. fimetarii (F.) (NS, NB, QC, ON, MB, SK, AB, BC, NT), A. forticornis (Csy.) (NB, QC, BC), A. impressa Eric. (NB, QC, ON, AB), A. macra (Csy.) (NB, QC, AB, BC, AK), A. neocarinula (Pel.), A. nigricollis (Csy.) (ON, MB, AB, BC, YT, AK), A. parviceps (Notm.) (QC, ON, AB), A. patens (Csy.) (NB, QC, ON), A. puella (Csy.) (AB), A. pumilio (Csy.) (NB, QC, ON, MB, SK, BC), A. tenebrosa (Csy.) (NB, QC, ON, AB, AK), A. umbrina (Gyll.) (NB, QC, ON, BC), A. undulata (Csy.) (NB, QC, ON, AB, BC, AK) et A. versa Grouv. (AK).

En plus des espèces mentionnées précédemment, voici la liste des 61 espèces mentionnées pour la première fois dans une province canadienne ou en Alaska: Caenoscelis basalis Csy.(NB, MB), C. ferruginea (C.R. Sahl,) (NS), C. parallela Csy. (AB), C. subdeplanata Bris. de Barn. (NB, BC), Antherophagus convexulus LeC. (SK, AB, AK), A. ochraceus Melsh. (NT), A. pallidivestis Csy. (NB, QC, ON, AB), A. suturalis Mäkl. (SK, AB), Cryptophagus bidentatus Mäkl. (QC, ON, MB, AB, YT), C. cellaris (Scop.) (QC, YT), C. confertus Csy. (ON, MB, YT), C. corticinus Thoms. (MB, AB, YT), C. dentatus (Hrbst.) (QC, ON, MB, YT), C. difficilis Csy. (NB, ON, MB, NT, YT, AK), C. distinguendus (QC, ON, SK, AB), C. fallax Balf.-Br. (QC, ON), C. hebes Csy. (YT), C. histricus Csy.(QC, ON, SK, AB), C. lapponicus Gyll. (QC, ON, AB, NT, YT, AK), C. laticollis Lucas (MB), C. mainensis Csy. (QC, ON), C. obsoletus Reit. (QC, MB, SK, AB, YT), C. peregrinus Wdrf. & Cmbs. (AB), C. quadrihamatus Mäkl. (QC, ON, MB, SK, AB, BC, YT), C. saginatus Sturm (PE, NB, QC), C. scanicus (QC, ON, BC, YT), C. scutellatus Newm. (QC, MB, AB), C. setulosus Sturm (AB, NT), C. stromus Wdrf. & Cmbs. (AB, AK), C. subfumatus Kraatz (NB, MB), C. tuberculosus Mäkl.(QC, ON, MB), C. varus Wdrf. & Cmbs. (QC, ON, MB), Henoticus serratus (AB), Henotiderus lorna (Hatch) (AB), Myrmedophila americana (LeC.) (ON), Pteryngium crenulatum (Eric.) (ON, AB, AK), Salebius octodentatus (Mäkl.) (AB), Atomaria (Anchicera) aleutica Csy. (MB, SK, AB, BC, NT, YT), A. apicalis Eric. (ON, SK, AB, BC, NT, YT, AK), A. distincta Csy. (ON, AB), A. ephippiata Zimm. (AK), A. fulvipennis Mann. (QC, ON, MB, SK, AB, BC, NT, YT), A. fuscata Schnrr. (AB, BC, NT, YT, AK), A. kamtschatica Motsch. (MB, AB, NT, YT), A. laetula LeC. (SK), A. lederi Johns. (NB, QC, ON, AB, BC), A. lewisi Reit. (MB, SK, AB, BC, AK), A. nubipennis Csy. (AB, AK), A. pusilla (Payk.) (ON), A. testacea Steph. (SK, BC), Atomaria (Atomaria) atrata Reit. (QC, ON, AB, AK), A. capitata (Csy.) (NB, QC, AB, AK), A. lineola (Notman) (ON, MB, SK, AB, BC), A. longipennis (Csy.) (QC, AB, AK), A. nigrirostris Steph. (NB, ON, SK, AB, BC, YT), A. planulata Mäkl. (QC, AB, YT), A. quadricollis (Csy.) (QC, ON, AB), A. stricticollis (Csy.) (QC, AB, YT), A. subdentata (Csy.) (NB, QC, ON, MB, SK, AB, AK), A. vespertina Mäkl. (QC, ON, MB, AB) et Ephistemus globulus (NF, LB, NS, QC).

Pour résumer, nous comptons 2 nouvelles mentions pour Terre-Neuve et Labrador, 9 nouvelles mentions pour la Nouvelle-Écosse, 31 pour le Nouveau-Brunswick, 66 pour le Québec, 50 pour l’Ontario, 33 pour le Manitoba, 24 pour la Saskatchewan, 69 pour l’Alberta, 41 pour la Colombie Britannique, 17 pour les Territoires du Nord-Ouest et le Nunavut, 25 pour le Yukon et 35 pour l’Alaska.

Selon les données de distribution et d’abondance, les 27 espèces suivantes sont considérées comme holarctique, ayant une répartition géographique naturelle dans le nord de l’Europe, de l’Asie et de l’Amérique du Nord, en passant par l’Alaska : Caenoscelis ferruginea, Cryptophagus acutangulus, C. bidentatus, C. confertus, C. corticinus, C. lapponicus, C. quadrihamatus, C. tuberculosus, Henoticus serratus, Henotiderus centromaculatus, Salebius octodentatus, Atomaria (Anchicera) apicalis, A. (Anc.) basalis, A. (Anc.) fulvipennis, A. (Anc.) fuscata, A. (Anc.) kamtschatica, A. (Anc.) peltata, A. (Anc.) sodermani, A. (Anc.) turgida, A. (Atomaria) affinis, A. (At.) atrata, A. (At.) elongatula, A. (At.) fimetarii, A. (At.) nigrirostris, A. (At.) subdentata, A. (At.) vespertina et Hypocoprus latridioides.

Les Cryptophagidés comprennent un grand nombre d’espèces adventices, introduites accidentellement par l’activité humaine. Plusieurs d’entre elles vivent à l’intérieur des bâtiments, mais d’autres ont colonisé avec succès les habitats naturels, dont les 34 espèces suivantes : Caenoscelis subdeplanata, Cryptophagus cellaris, C. dentatus, C. denticulatus, C. distinguendus, C. fallax, C. laticollis, C. lycoperdi, C. obsoletus, C. punctipennis, C. saginatus, C. scanicus, C. scutellatus, C. setulosus, C. subfumatus, Henoticus pilifer, Micrambe ulicis, Pteryngium crenulatum, Telmatophilus typhae, Atomaria (Anchicera) atra, A (Anc.) lederi, A (Anc.) lewisi, A (Anc.) mesomela, A (Anc.) morio, A (Anc.) munda, A (Anc.) ornata, A (Anc.) pusilla, A (Anc.) testacea, A. (Atomaria) alpina, A. (At.) impressa, A. (At.) linearis, A. (At.) umbrina, Curelius japonicus et Ephistemus globulus.

ZooBank Registration: urn:lsid:zoobank.org:pub:2F5486A4-24BA-4694-B1E3-B06412973044

Introduction

Cryptophagidae, also known as hairy fungus beetles, are common beetles often found as adults in the leaf litter, under the bark of trees, logs and stumps. They are easily distinguished by their small size (usually less than 3.5 mm), oval or elongate-oval body, often reddish to yellowish brown, sometimes dark brown to black, usually covered with silky hairs, antennae with a loose 3-segmented club and basal insertions visible from a dorsal view (White 1983), pronotum usually with well-developed lateral carinae, abdominal ventrite 1 longer than the each of the other ventrites, epipleuron distinct in its basal half, and elytra with confused punctation (Leschen and Skelley 2002). They are very secretive beetles, hence the name “Cryptophagidae”, which loosely translated means “feeding in hidden places”. About 165 species belonging to 16 genera have been recorded so far in North America. All genera are included in this publication except Sternodea, which is found in Florida and does not have any described Nearctic species, and Amydropa,with only one described species from Baja California (Leschen and Skelley 2002). Bousquet et al. (2013) listed 73 species in Canada, including 18 Holarctic and 10 introduced species.

The taxonomy of Cryptophagidae is moderately known in Canada and the northern United States of America. Cryptophagus, the nominal genus, was described by Herbst in 1792 with C. scanicus (Linnaeus)as the type species. Cryptophagus cellaris and C. lycoperdi, both described by Scopoli in 1763 andintroduced into North America, were among the first species of the family to be named. During the 19th century, most studies of Cryptophagidae were carried out in Europe, mainly by Erichson in 1846 and Mäklin in 1852-1853. In North America, very few species were described prior to 1900 (LeConte 1869). Casey (1900) made a major contribution to the family; however, the majority of species that he described in his later revision (Casey 1924) have been synonymized. Although he described 114 species in his publications, 66 species fell into synonymy, including 24 species of Cryptophagus Herbst and 21 species of Atomaria Stephan.

No major revision was published on Nearctic species until Woodroffe and Coombs (1961) on Cryptophagus. This excellent and well-illustrated publication has been helpful to link the Palearctic and Nearctic fauna and to separate distinct species based on their geographic variation, which can be significant in that genus. Four of Casey’s species were synonymized in the highly variable C. croceus Zimmermann. Ten introduced species and two Holarctic species were recorded for the first time in North America. They described five new Nearctic species, including the common C. varus Woodroffe & Coombs.

Hatch (1962) revised the family in the Pacific Northwest, stabilizing the names of many species in that region. Bousquet (1989) reviewed the North American genera of Cryptophaginae. Myrmedophila Bousquet was extracted from Cryptophagus as a newly described genus.Poole and Gentili (1996) list all known Nearctic Cryptophagidae. Leschen (1996) revised the genera of the entire family with cladistic phylogenetic analysis. Anchicera Thomson was definitively united with Atomaria as a subgenus, and Glyptophorus Parkwas united with Henoticus ThomsonDownie and Arnett (1996) provided identification keys and diagnoses of all species known in northeastern North America. Among the 56 species of Cryptophagidae recorded, 36 species are still valid, mostly in the Cryptophaginae. Lyubarsky (2012) described three species of Atomaria from the eastern Palaearctic but none of these species fit the description of any Nearctic species.

Cryptophagidae are distributed worldwide. Cryptophagus, Henoticus, Caenoscelis Thomson and Atomaria are widely distributed in North America. Antherophagus Dejean, Henotiderus Reitter, Myrmedophila and Telmatophilus Heer have a northern and a western distribution. Salebius Casey and Hypocoprus Motschulski have been collected so far only in western North America. Pteryngium Reitter appears to be Holarctic. Curelius and Ephistemus have been introduced from Europe or Asia.

Cryptophagidae are found in a wide variety of habitats. Cryptophagus, Henoticus and Atomaria are mostly forest dwellers and are often collected in soil litter and under bark. Henotiderus prefers trees, under the bark, whereas Myrmedophila and Hypocoprus prefer ant nests (Leschen & Skelley, 2002). Many specialized species of Atomaria live in animal burrows, beaver lodges, or in ant and bird nests. Antherophagus adults are found on flowers, feeding on pollen, whereas the larvae of some species live inside bumblebee nests and beehives where they feed on honey. Telmatophilus are common in aquatic habitats, where they feed on Spargania and Typha (Leschen & Skelley, 2002). Many species of Cryptophagus and Atomaria (Anchicera) are found indoors where they feed on stored food products (Bousquet, 1990), or on fungi and mold found within the walls or in basements where humidity is high. Any water problems, often caused by leaking pipes, will usually attract fungi and mold that will attack wood structures and that will in turn attract many silky fungus beetles, mostly in remote areas. Because of their small size and their secretive behavior, they usually remain undetected.

Species in the genera Telmatophilus, Hypocoprus and Tisactia Casey seem to be more active in spring, from April to June, but most species of other genera are most active from June to mid-August. Many species have an extented period of activity that can include fall and winter, remaining active under the snow cover and at the base of tree trunks. Indoors, most species are active year round. Many species that are mostly collected in late spring and summer in Canada are mostly active during winter in the southeastern United States of America.

Methods & Materials

This publication covers all 142 described species that can be found in Canada and the northern United States of America. Types of most species described by Casey (1900) (except for species of Cryptophagus)were examined and photographed by GP, and are redescribed according to the current standards. A list of types and other material examined (with their original name) from various institutions, including synonymies (names in bold are still valid today) and new combinations is provided in the supplemental materials. Types of most species described by Casey (1900) (except for species of Cryptophagus) were examined and photographed by GP, and are redescribed according to the current standards. Unfortunately, it was not possible to examine holotypes of Palearctic species held in European collections. Our concept of Holarctic species follows Woodroffe & Coombs (1961) for Cryptophagus and Johnson (1992) for Atomaria and some species of Cryptophagus that were renamed since Woodroffe & Coombs (1961).

Also, collaborations were made with Richard Leschen and Colin Johnson, two world specialists of the family working on a world checklist. They compiled the Palearctic checklist, including all known species from Europe, Asia and North Africa, (Johnson et al. 2007 – including an unpublished 2014 update). This list gives a broad view of the family, including all species that could potentially be found in North America. From that list, we selected all species with an extensive distribution that could potentially be introduced into North America, and all northern species that could potentially be present in northern Canada and Alaska (Supplemental Materials).

The present publication is available in PDF format along with an online interactive identification key. For each species, a detailed description and image of the habitus, sometimes an image taken in the field, habitat data, some ecological data and a distribution map are provided. Photographs of habitus or morphological structures were taken using a Nikon digital camera DMX1200F through a binocular Nikon SMZ. All photos were processed through a Proscan II system by Prior Scientific (https://www.prior.com/), which stacks all images taken at different layers into one focused image. Each part (head, thorax, elytra, antennae, legs) of all species was photographed individually, in order to improve the resolution, and then joined together using Adobe Photoshop Element software. Images of the following Palearctic species, either introduced or Holarctic, were provided by Lech Borowiec from the University of Wroclaw (see Supplemental Materials for additional related material examined):

Caenoscelis subdeplanata

    •  Brisout de Barneville

Cryptophagus dentatus

    •  (Herbst)

Cryptophagus laticollis 

    • Lucas

Cryptophagus lycoperdi 

    • (Scopoli)

Cryptophagus scutellatus

    •  Newman

Henoticus serratus

    •  Gyllenhal

Telmatophilus typhae

    •  Fallén

Atomaria

    •  (

Anchicera

    • )

apicalis 

    • Erichson

Atomaria

    •  (

Anchicera

    • )

fuscata 

    • (Schönherr)

Atomaria 

    • (

Anchicera

    • )

lewisi

    •  Reitter

Atomaria 

    • (

Anchicera

    • )

mesomela 

    • (Herbst)

Atomaria

    •  (

Anchicera

    • )

pusilla

    •  (Paykull)

Atomaria

    •  (

Anchicera

    • )

testacea 

    • Stephens

Atomaria

    •  (

Atomaria

    • )

elongatula

    •  Erichson

Atomaria

    •  (

Atomaria

    • )

fimetarii 

    • (Fabricius)

Atomaria

    •  (

Atomaria

    • )

linearis

    •  Stephens

Ephistemus globulus 

    (Paykull)

 

Most specimens from the Canadian National Collection in Ottawa were examined. Many specimens from Quebec came from biodiversity projects led by Christian Hébert at the Laurentian Forestry Centre. Claude Chantal, who holds the largest private beetle collection in Quebec, provided a substantial number of specimens from Quebec for study. The Northern Forestry Centre in Alberta (Dave Langor, James Hammond), the University of British Columbia (Karen Needham), the University of Georgia (Richard Hoebeke) and University of Alaska (Derek Sikes) also provided us with numerous specimens. Steve Marshall from the University of Guelph provided a few specimens and some ecological data from Ontario. Reggie Webster also provided a few specimens from New Brunswick.

The genera of Cryptophagidae are well defined taxonomically, thanks to Leschen (1996). Johnson (1992) has developed a useful key to identify most species from central Europe; however, no existing key can help identify all species found in Canada, especially those of the genus Atomaria. We simplified the key of Woodroffe and Coombs (1961) on Cryptophagus,and it should help easily identify nearly all specimens found in Canada and the USA. We hope that this publication will help students, amateurs, technicians and professional entomologists to easily identify most species.

This review was prepared because of the increased interest and research activities (mainly biodiversity projects) related to cryptophagids. We believe that many species can be good ecological indicators for old growth forests as these forests may be subject to more extensive decaying processes involving fungi, the main food source of cryptophagids. Do these beetles, by their activities, limit the growth of mycelium and protect the trees by limiting the spread of tree diseases or, on the contrary, disperse the spores of the fungi and accelerate the decaying process of the forest? The answers to those questions are unknown for the moment but increasing the knowledge of this family and knowing which species can be found in which types of forest, at which stage of maturity or decay, will help to provide some answers in the future.

Interactive keys for species determination

The interactive key was created to facilitate the determination of all species of Cryptophagidae in Canada, Alaska and northern mainland USA (south to Maryland west to Oregon), covering all habitats that can be found in Canada. This key includes all known Nearctic Atomaria. However, a few doubtful species of Atomaria introduced from Europe and previously recorded in North America, but not seen in our material, are not included in this work.This includes the following species:

Atomaria (Atomaria) diluta 

    • Erichson

Atomaria (Atomaria) nigriventris 

    Stephens

 

Atomaria lepidula was described by Mäklin from Alaska; however, the holotype, stored in the Finnish Museum of Natural History, examined and returned by Colin Johnson in 1971, has been lost. According to its original (very short) description in Latin, this species belongs to the subgenus Anchicera, is a uniform reddish brown and is very large in size for the genus (2.4 mm). No specimen with these characters was seen in our material so this name will be considered a nomem dubium.

The interactive key uses external morphological characters that are very useful to identify all species. The most useful diagnostic characters include antennae, mainly the first segment’s (A1) shape and length relative to the second, club A9-A10 shape (subquadrate, slightly to strongly transverse), sometimes the shape of A4-A8, pronotum shape and proportions, elytra shape, proportions and pilosity, and sometimes colour pattern.

Genitalia, which are useful to confirm the identification of species in many beetle families, are not very useful for Cryptophagidae; however, DNA barcoding has been by far a better method to separate species without the risk of destroying specimens (needing only to remove one leg). The DNA barcoding method will be explained in a later section.

Descriptive plates of species

In order to facilitate the identification of the species, all diagnostic characters are in bold, including all major sections of the description: length (L), head, antennae, pronotum, elytra. After the description, each species is compared with the most closely related one or any other species which can be confused.
Both sexes are described together. Species of Cryptophaginae only have slight sexual dimorphism in the tarsal formula, being 5-5-4 in male and 5-5-5 in female. In a few species, such as Caenoscelis antennata, sexescan be also separated by their antennal structure. However, only the presence of tenant setae in claws of males can separate the sexes of Atomariinae for all species. Total length of the species is measured from the tip of the clypeus to the tip of the elytra or abdomen.

Each part of the beetle is described in the following sequence: head (including eyes, mandibles, palpi, clypeus, antennae), thorax (pronotum, sterna, legs, elytra), and abdomen (sternites), describing colouration first, then morphometry and other structural characters. Eyes/head width ratio is calculated by dividing the distance between the internal eye orbits at the level of maximum head width (FW: frons width) by head width from dorsal view (HW) and subtracting the result from 1: 1-(FW/HW). The clypeus is quadrate, without any significant difference between genera or species. Antennae (A) are described in detail, including the shape and relative length of each article compared to the following one (A1-A8) and the relative width of club articles (A9-A11). Antennal description is important to separate species groups and sometimes species.

Pronotum colour, width/length ratio, margin shapes, surface porosity, punctures and pubescence are noted. Width/length ratios of pronotum are measured from maximum width and length at the middle. This ratio can vary from subquadrate (1:1) to twice as wide as long (2:1), with only one species being elongate (Hypocoprus latridioides). Lateral margins can be arcuate, sinuate, straight, subparallel, dentate, crenulate or with a callosity at anterior angles. Pronotum shape and width/length proportion are crucial for the diagnosis of most species. Pronotum is usually glossy, subglossy or rarely opaque (with distinct microsculpture) with strong, moderate or sparse punctures. Pilosity is usually moderately long and sparse but sometimes more dense. Colouration of the sterna (including prosternum, meso- and metaventrite), scutellum and legs is mentioned.

Elytra colour, length/width ratio and elytra/pronotum length ratio are recorded. Elytra often have sides entirely arcuate or straight and divergent at the anterior half or more, rarely subparallel but usually arcuate and convergent at apical half or less. Surface is usually glossy and pilosity is dense, moderate or sparse, long or short, erect, suberect, appressed or prostrate, sometimes both. Abdominal sternite colour is noted. All males have internal genitalia and there is no external dimorphism feature on the abdominal ventrites.

Colour is not very useful to separate most species. Many species have a uniform reddish brown body. Many others have a dark brown head and pronotum combined with yellowish brown elytra. A few species are entirely dark brown to black or entirely yellowish brown. A few species have a constrasting black pattern on paler elytra. Some species such as Atomaria fuscata Schönherr can be very variable in colour, being entirely reddish brown to dark brown. All colour variations are mentioned in the text for each part of the body when necessary. Antennal structure, pronotum and elytra shape have been used to separate most species. Color is mostly used as a confirmation character when there is not much variation within a species except in distinctly patterned species.

Similar species related to the one described are mentioned with all the characters that can help to differenciate between them, starting with the most similar species. All the characters discussed above are usually mentioned in the key.

Distributions are given from eastern Canada to the northwest, including Alaska, down to the southeast and the southwest, including the range outside the geographic scope of this work, both for species occurring in Canada and for species that have not yet been recorded in Canada but that occur in the USA. All distribution maps indicate ecozones (Figure 1), as defined below; however, for the sake of simplicity, some ecozones were grouped together in order to correspond to most species’ distributions.

Arctic zone, northern Canada from coast to coast (including most Arctic Ocean islands) and Alaska, covered mostly by tundra.

Taïga zone, also called Subarctic and sometimes Hudsonian zone, bordering south of the Arctic zone, from coast to coast, covered mostly by open black spruce forests growing sparsely on lichen.

Boreal zone, also called Canadian zone, mostly covered by dense black spruce forests, mostly dominated by balsam fir in the south, extending nearly coast to coast.

Mixed Wood Plains zone, also called Transition zone, mostly covered by deciduous forests dominated by maple and beech, but also by mixed forests with white pine and red oak in more humid areas along the St. Lawrence River and in the Maritime Provinces and states. This area is mostly covered by agricultural lands and many prairie species expand their distribution eastward in that region.

Northern Appalachians zone, includes the Atlantic Maritime zone of Canada, a mountain chain extending from eastern Quebec and southern New Brunswick to western Massachusett, Connecticut, and the Adirondack Mountains of New York; a southern extention of the Boreal zone, dominated by spruce-fir and maple-beech forests.

Southern Appalachians zone, also called Alleghanian zone, a mountain chain extending from Pennsylvania to north Georgia and Alabama, including the Ozark Mountains of Missouri and Arkansas; a southern extention of the Northern Appalachian and Transition zones, covered by maple-beech forests, but with a higher proportion of pine and oak-hickory forests. Spruce-fir forests can be found on the highest mountains.

Southeastern USA Forest Plain zone, also called Carolinian zone, extending south of the Transition and the Alleghanian zones, from New Jersey down to northern Florida and west to Illinois, Arkansas and Texas, dominated by pine and oak-hickory forests.

Southeastern USA Coastal Plain zone, also called Louisianian zone, including the Texas-Louisiana Coastal Plain, bordering the Atlantic Ocean and the Gulf of Mexico seashore south of the Carolinian zone, from New Jersey down to Florida west to the Mississipi Valley and Texas, dominated by vast pine forests and by evergreen hardwood forests dominated by oak.

Temperate Prairie zone, excluding the South Central Semi-Arid Prairie zone, west of the Boreal, Mixed Wood Forest and Southeastern USA Forest Plain zones, from southern Manitoba west to Alberta and south to Missouri, Kansas, Nebraska, and Wyoming, dominated by grasslands and meadows.

Western Cordillera zone, including the cordillera of the western US, from western Alberta to British Columbia, south to Colorado, New Mexico, Utah, Idaho and California; dominated by Ponderosa and lodgepole pines, it is a very diversified area that contains many endemic species, and also extended range of a many northern species.

Marine West Coast Forest zone, a narrow strip along the Pacific Coast from southern Alaska and British Columbia south to northern California; dominated by large conifers such as Douglas-fir, Sitka spruce, western cedar and western white pine.

Other ecozones are recorded, but are only represented by a smaller number of species or have not been adequately sampled.

The distribution pattern of a species does not follow political borders. Our ecozone map system has been especially designed to give a more relevant idea of the likely distribution of a species. When a species is present only in the eastern and the western part of an ecozone, especially in a relatively undisturbed ecozone like the Boreal zone, the probability is very high that it will be eventually collected in between. The main reason why many species have not been reported from northern Ontario, Manitoba and Saskatchewan is because these areas have not been as intensively sampled as in Quebec or Alberta. However, if two specimens have been collected in two distant localities belonging to two different ecozones, the probability that one of these is an accidental introduction is much higher. In that case, we cannot argue for a real continuous distribution.

We include an index of commonness of a species. This mainly reflects the number of specimens seen in collection, which does not necessarily translate to reality in the field, especially for species living in very remote areas or in specialized habitats:

    • Very common: more than 300 specimens seen;

 

    • Common: between 100 and 300 specimens;

 

    • Fairly common: between 50 and 100 specimens;

 

    • Uncommon: between 10 and 50 specimens;

 

    Rare: fewer than 10 specimens.

Table 1 represents a list of all species of Cryptophagidae found in Canada and the northern USA together with their respective ecozones. The results of this table show that the most well diversified ecozones for Cryptophagidae are the Boreal (100 species), Western Cordillera (97 species), Mixedwood Plain (81 species) and Northern Appalachian (76 species). Of course, these ecozones were intensively sampled by three different Canadian Forestry Service centers.

Seasonality is also recorded, mentioning at first the wider time distribution pattern based on extremes of collecting dates, followed by the usual period of adult activity including their peak for common species, all shown in a bar graph. However, for species represented by less than 20 specimens, no bar graph is shown.

Habitats and host trees or plants, when known, are also noted for each species. Most collectors before 1980 did not record any ecological data for most specimens. Since 1992, when biodiversity started being considered in CFS research studies, habitats of many species have begun to be known. Most species living in spruce, mixed fir-birch, and maple forests are now well known. Other species living in grasslands, open shrubby fields and orchards are also relatively well known. However, although some species have been recorded in these habitats, oak and pine forests still need to be more deeply investigated, including freshwater and coastal marshes and bogs. Today, habitats are known for about 85% of the species. Finally, the geographic distribution of the species and the presence of better-known species in the same locality collected on the same date can help us to determine the probable habitats of a determined species, though that information will need to be verified by further research. Except otherwise noted, most habitat data were recorded from specimen labels. Finally, the number of specimens examined is given to indicate the relative abundance of the species in the collections.

Figure 1. Nearctic ecozones.

Checklist

Table 1: Checklist of Cryptophagidae of Canada and Alaska

This list follows Bousquet et al. (2013) with each column representing a state or a province. It includes also all the North American synonymies. All species written in red are new records for Canada. All provinces or states in red are new provincial or state records. For ecozones, shaded cells represent ecozones where the species is well represented, corresponding to its normal distribution while cells in white are where the species is rather scarcely distributed, outside of its normal range. AK – Alaska; YT – Yukon Territory; NT – Northwest Territories (including Nunavut); BC – British Columbia; AB – Alberta; SK – Saskatchewan; MB – Manitoba; ON – Ontario; QC – Quebec; NB – New Brunswick; NS – Nova Scotia; PE – Prince Edward Island; LB – Labarador; NF – Newfoundland. A – Arctic Zone; T – Taïga Zone; B – Boreal Zone; MW – Marine West Coast Forest Zone; WC – Western Cordillera Zone; P – Temperate Prairie Zone; NA – Northern Appalachian Zone; MP – Mixedwood Plain Zone; SA – Southern Appalachian Zone; SE – Southeastern USA Coastal Plain Zone. * indicate Holarctic species. † indicate adventive species.

Province Ecozone
Subfamily CRYPTOPHAGINAE Kirby, 1826
Tribe Caenoscelini Casey, 1900
Genus CAENOSCELIS C.G. Thomson, 1863
C. antennalis (Casey, 1924) BC AB SK MB ON QC NS B WC P NA MP
C. basalis Casey, 1900 MB ON QC NB NS NF B WC P NA MP SA SE
C. ovipennis Casey, 1900 syn. nov.
C. macra Casey, 1900 syn. nov.
C. elongata Casey, 1900 syn. nov.
C. cryptophaga Reitter, 1875 ON QC NS B NA MP SA SE
C. obscura Casey, 1899? syn. nov.
C. subfuscata Casey, 1900 syn. nov.
C. ferruginea (C.R. Sahlberg, 1820)* AK YT NT BC AB SK MB ON QC NS T B MW WC P NA MP SE
C. testacea (Zimmermann), 1869
C. ochreosa Casey, 1900 syn. nov.
C. parallela Casey, 1900 AK BC AB MB B MW WC P
C. angusticollis Casey, 1900 syn. nov.
C. shastanica Casey, 1900 syn. nov.
C. subdeplanata Brisout de Barneville, 1882* BC AB MB ON QC NB B MW WC P NA MP SA SE
C. macilenta Casey, 1900 syn. nov.
Tribe Cryptophagini Kirby, 1826
Genus ANTHEROPHAGUS Dejean, 1821
A. convexulus LeConte, 1863 AK AB SK ON QC NB NS B MW P NA MP SA
A. ochraceus Melsheimer, 1844 NT BC AB SK MB ON QC NB NS PE NF T B MW WC P NA MP SA SE
A. subnitidus Grouvelle, 1911
A. oregonus Casey, 1924
A. pallidivestis Casey, 1900 BC AB ON QC NB B MW WC P NA
A. suturalis Mäklin, 1853 AK BC AB SK T B MW WC
Genus CRYPTOPHAGUS Herbst, 17921
C. acutangulus Gyllenhal, 1827* AK BC AB SK MB ON QC NB NS NF B MW WC P NA MP
C. prominens Casey, 1924
C. restrictus Casey, 1924
C. bidentatus Mäklin, 1853* AK YT BC AB MB ON QC A T B MW WC P
C. punctatissimus Mäklin, 1853
C. cellaris (Scopoli, 1763)† YT BC AB SK MB ON QC A T B MW WC P NA MP
C. rupimontis Casey, 1924
C. angustatus Casey, 1924
C. confertus Casey, 1900* AK YT NT BC AB SK MB ON QC NS PE A T B MW WC P NA
C. archangelicus Sahlberg, 1926
C. corticinus C.G. Thomson, 1863* AK YT BC AB SK MB ON QC A T B WC NA MP
C. depressulus Casey, 1900
C. croceus Zimmermann, 1869 AB SK MB ON QC NS B WC P NA MP SA SE
C. crinitus Zimmermann, 1869
C. fungicola Zimmermann, 1869
C. nodifer Casey, 1900
C. amputatus Casey, 1900
C. laticlavus Casey, 1900
C. densicollis Casey, 1924
C. dentatus (Herbst, 1793)† AK YT BC MB ON QC T B WC NA MP SA SE
Corticaria fumata Marsham, 1802
Cryptophagus quadridentatus Mannerheim, 1843
C. denticulatus Heer, 1841† BC AB MB ON QC WC P MP SE
C. difficilis Casey, 1900 AK YT NT BC AB SK MB ON QC NB LB NF T B WC P NA MP SE
C. contractus Casey, 1924
C. dilutus Reitter, 1874
C. distinguendus Sturm, 1845† BC AB SK ON QC T B WC P NA MP SE
C. keeni Casey, 1924
C. fallax Balfour–Browne, 1953† ON QC NB NS PE B NA MP
C. hebes Casey, 1900 AK YT BC B MW WC
C. lyraticollis Casey, 1900
C. otiosus Casey, 1900
C. histricus Casey, 1900 BC AB SK ON QC LB B WC NA MP SA SE
C. lapponicus Gyllenhal, 1827* AK YT NT BC AB ON QC A T B MW WC NA MP
C. validus Kraatz, 1856
C. laticollis Lucas, 1846† BC AB SK MB NS MW WC P MP SE
C. affinis Sturm, 1845 - Homo.
C. inscitus Casey, 1900
C. cribricollis Casey, 1900
C. lycoperdi (Scopoli) 1763† QC MP
C. mainensis Casey, 1924 ON QC NB NS NF B NA MP
C. maximus Blake, 1928 BC WC
C. obsoletus Reitter, 1879† YT AB SK MB ON QC T B WC P MP
C. peregrinus Woodroffe & Coombs, 1961 BC AB MW WC
C. porrectus Casey, 1900 AK BC B WC
C. punctipennis Brisout de Barneville, 1863† BC SK MB ON QC NB B WC NA MP
C. pilosus sensus auctorum syn. nov.
C. parvinoda Casey, 1900
C. infuscatus Casey, 1900
C. plectrum Casey, 1900
C. cicatricosus Casey, 1900
C. quadrihamatus Mäklin, 1853* AK YT BC AB SK MB ON QC T B WC P NA
C. plagiatus Poppius, 1900
C. saginatus Sturm, 1845† AK BC SK ON QC NB PE B MW WC NA MP
C. scanicus (Linnaeus, 1758)† YT BC ON QC NF T B MW WC MP
C. scutellatus Newman, 1834† AB MB ON QC B MW P NA MP
C. setulosus Sturm, 1845† NT BC AB SK ON QC LB NF T B WC P NA MP SA SE
C. antennatus Casey, 1900
C. grossulus Casey, 1924
C. stromus Woodroffe & Coombs, 1961 AK AB SK MB MW WC P
C. subfumatus Kraatz, 1856† BC AB SK MB ON QC NB B WC P NA MP SE
C. tuberculosus Mäklin, 1852* AK BC AB SK MB ON QC NS NF T B MW WC P MP
C. debilis LeConte, 1858
C. brevipilis Casey, 1900
C. lepidus Casey, 1900
C. boulderanus Casey, 1924
C. instabilis Bruce, 1936
C. valens Casey, 1900 BC SK MB ON QC B WC P NA MP SA SE
C. varus Woodroffe & Coombs, 1961 BC AB SK MB ON QC B MW WC P NA MP
Genus HENOTICUS C.G. Thomson, 1868
Glyptophorus Park, 1929
H. californicus (Mannerheim, 1843) BC MW WC
Cryptophagus inconspicua LeConte, 1863
Henoticus loberinus Sharp, 1900
H. germanicus Reitter, 1906
H. pilifer(Reitter, 1888)† BC ON QC MW NA MP SA SE
H. mycetoecus (Park, 1929) syn. nov.
H. serratus (Gyllenhal, 1808)‡ AK BC AB SK MB ON QC NB NS NF T B MW WC P NA MP SA SE
Paramecosoma denticulata LeConte, 1850
H. varidentatus Pelletier, n. sp. QC MP
Genus HENOTIDERUS Reitter, 1877
Crosimus Casey, 1900
Henoticoides Hatch, 1962
H. centromaculatus Reitter, 1877* AK NT AB SK ON QC NB NS B WC P NA MP SA SE
Crosimus hirtus (Casey, 1900)
C. obesulus (Casey, 1900)
Henoticoides parsonsi Johnson, 1972
H. lorna (Hatch, 1962) AK BC AB B MW WC
Genus MICRAMBE Thomson, 1863
M. ulicis (Stephens, 1830)† QC B
Genus MYRMEDOPHILA Bousquet, 1989
M. americana (LeConte, 1879) AK YT BC AB SK MB ON QC NB B WC P NA
Cryptophagus blumi Blaisdell, 1937
Genus PTERYNGIUM Reitter, 1887
P. crenulatum (Erichson, 1846)† AK BC AB ON QC NB NS B MW WC NA MP
Pteryngium crenatum of authors, not (Gyllenhal, 1808) nor (Fabricius, 1798)
P. malacum Casey, 1900
Genus SALEBIUS Casey, 1900
S. minax Casey, 1900
S. octodentatus (Mäklin, 1852)* AK BC AB MW WC
S. lictor Casey, 1900 syn. nov.
S. montanus Casey, 1900 syn. nov.
S. sexdentatus Casey, 1900
S. tarsalis Casey, 1900
Genus TELMATOPHILUS Heer, 1841
T. americanus LeConte, 1863 BC AB SK MB ON QC NB NS NF B WC P NA MP
T. typhae (Fallén, 1802)† QC NB NS PE B NA MP
C. schonherrii Gyllenhal, 1808 (Cryptophagus)
T. bicolor Reitter, 1913
Subfamily ATOMARIINAE LeConte, 1861
Tribe Hypocopriini Reitter, 1879
Genus HYPOCOPRUS Motschulski, 1839
H. latridioides Motschulski, 1839 AB SK P
H. formicetorum Motschulsky, 1839
H. tenuis Casey, 1900
Tribe Atomariini LeConte, 1861
Genus ATOMARIA Stephens, 1829
(Subgenus Anchicera C.G. Thomson, 1863)
A. aleutica Casey, 1900 AK YT NT BC AB SK MB A T B MW WC P
A. apicalis Erichson, 1846* AK YT NT BC AB SK MB ON QC NB NS NF T B WC P NA MP
A. ovalis Casey, 1900
A. nebulosa Casey, 1924 syn. nov.
A. arcuaticollis Pelletier, n. sp. QC B NA
A. atra (Herbst, 1793)† QC NB NA MP
A. basalis Erichson, 1846* AK YT NT BC MB QC NB T B WC NA
A. brevicollis Casey, 1900 NT BC AB SK MB ON QC NB T B MW WC P NA MP
A. distincta Casey, 1900 AB ON QC NB NS B P NA MP SA
A. divisa Casey, 1900 syn. nov.
A. caseyi (Grouvelle), 1916
A. ephippiata Zimmermann, 1869 AK NT BC AB SK MB ON QC NB NS PE LB NF T B MW WC P NA MP SA SE
A. fulvipennis Mannerheim, 1846* AK YT NT BC AB SK MB ON QC A T B MW WC P
A. fuscata Schöenherr, 1808* AK YT NT BC AB SK MB ON QC NB NS PE LB NF T B MW WC P NA MP SA SE
A. ochracea Zimmermann, 1869
A. saginata Casey, 1900
A. pennsylvanica Casey, 1900
A. gonodera Casey, 1900 syn. nov.
A. riparia Casey, 1900 syn. nov.
A. subalutacea Casey, 1900 syn. nov.
A. incerta Casey, 1900 syn. nov.
A. crypta Casey, 1900 syn. nov.
A. gilvipennis Casey, 1900 ON QC NS T B NA MP SA
A. hudsonica Pelletier, n. sp. MB T
A. inepta Casey, 1900 AK NT BC QC T B MW WC NA MP
A. kamtschatica Motschulsky, 1845* AK YT NT BC AB MB A T B MW WC P
A. laetula LeConte, 1857 BC SK WC P
A. lederi Johnson, 1970† BC AB ON QC NB NS B MW WC P NA MP
A. lewisi Reitter, 1877† AK BC AB SK MB ON QC NB NS B MW WC P NA MP SA SE
curtula Casey, 1900
A. mesomela (Herbst, 1792)† BC MW
A. morio Kolenati, 1846† AB SK ON QC NS WC P NA MP
ebenina Casey, 1924 syn. nov.
A. munda Erichson, 1846† AK BC QC NB B MW WC P NA MP SE
A. neomunda Pelletier, n. sp. AK QC B
A. nigrirostrisoides Pelletier, n. sp. AK B
A. nigritaria Pelletier, n. sp. MB QC P MP
A. nubipennis Casey, 1900 AK BC AB MW WC P
A. oblongula Casey, 1900 AK NT BC AB SK MB ON QC NB NS NF T B WC P NA MP
A. ocularia Pelletier, n. sp. AB P
A. ornata Heer, 1841† BC WC
A. parallelicollis Pelletier, n. sp. QC B
A. peltata Kraatz, 1853* AK BC B
A. postpallens Casey, 1900 AK NT BC AB T MW WC P
A. pusilla (Paykull, 1798)† AK BC ON QC NB NS B MW WC NA MP
A. fallax Casey, 1900 syn. nov.
A. salicicola Pelletier, n. sp. AB SK WC
A. sodermani Sjöberg, 1947* NT QC T MP
A. testacea Stephens, 1830† BC SK QC NB NS MW P NA MP
Silpha ruficornis Marsham, 1802
A. turgida Erichson, 1846* AK YT NT BC AB SK MB ON QC NB T B MW WC P NA MP
A. nanula Casey, 1900 syn. nov.
(Subgenus Atomaria Stephens, 1829)
Agathengis Gozis, 1886
A. affinis R.F. Sahlberg, 1834* AK BC AB QC NB NS B WC P NA MP
A. albertana Pelletier, n. sp. AB B
A. alpina Heer, 1841† BC AB ON QC NB NS B MW WC P NA MP
A. atrata Reitter, 1875* AK BC AB ON QC B MW P MP
melanica Hatch, 1962
A. calidaria Pelletier, n. sp. AB WC
A. californica Pelletier, n. sp. WC
A. capitata (Casey, 1900) AK BC AB QC NB B MW WC NA
A. coloradensis (Casey, 1900) AK YT BC AB QC NB B WC MP SA
A. strandi Johnson, 1967 syn. nov.
A. constricta (Casey, 1900) BC AB QC NB B WC P NA MP SA SE
A. crassula (Casey, 1900) QC NB B NA MP SA
A. cribricollis Pelletier, n. sp. AK AB B WC
A. dispersa (Casey, 1900) AK BC AB ON QC NB T B MW WC NA
A. elongatula Erichson, 1846* AK AB QC B MW MP
A. fimetarii (Fabricius, 1792)* AK NT BC AB SK MB ON QC NB NS T B WC P NA MP
A. forticornis (Casey, 1900)
A. impressa Erichson, 1846† AB ON QC NB B NA MP
A. impressicollis Pelletier, n. sp. AK YT BC AB B WC
A. linearis Stephens, 1830† WC
A. lineola (Notman, 1920) BC AB SK MB ON QC NB NS LB NF B WC NA MP SA
A. wollastoni of authors, not Sharp, 1867
A. longipennis (Casey, 1900) AK BC AB QC B MW WC NA
A. lyricollis Pelletier, n. sp. MB B
A. macra (Casey, 1900) AK BC AB QC NB B MW WC NA MP SA SE
A. neocarinula Pelletier AB P MP
A. carinula (Casey, 1900), preoccupied by A. carinula Rey, 1889
A. nigricollis (Casey, 1900) AK YT BC AB MB ON T B MW WC
A. parvicollis (Casey, 1900), preoccupied by A. parvicollis Rey syn. nov.
A. parva Schenkling, 1923 syn. nov.
A. nigrirostris Stephens, 1830* AK YT BC AB SK ON QC NB NS PE LB NF B MW WC P NA MP SA SE
A. fuscicollis Mannerheim, 1852
A. subnitens (Casey, 1900)
A. luculenta (Casey, 1900) syn. nov.
A. lucida (Casey, 1900), preoccupied by A. lucida Reitter, 1889 syn. nov.
A. lucens Grouvelle syn. nov.
A. ontariensis Pelletier, n. sp. ON MP
A. orbicollis Pelletier, n. sp. ON QC B MP
A. parviceps Notman, 1921 AB ON QC B WC NA MP
A. patens (Casey, 1900) ON QC NB B NA MP SA
A. pinicola Pelletier, n. sp. BC QC NB B WC NA MP
A. planulata Mäklin, 1853 AK YT BC AB QC T B MW WC
A. pseudotsugae Pelletier, n. sp. BC MW
A. puella (Casey, 1900) AB WC P
A. puelloides Pelletier SA SE
A. pumilio (Casey, 1900) BC SK MB ON QC NB B MW WC P NA MP SA SE
A. quadricollis (Casey, 1900) BC AB ON QC B MW WC NA
A. stricticollis (Casey, 1900) YT BC AB QC B MW WC P
A. ochronitens (Casey, 1900) syn. nov.
A. subdentata (Casey, 1900)* AK BC AB SK MB ON QC NB B MW WC P NA MP
A. subangulata J. Sahlberg, 1926 syn. nov.
A. subrecta (Casey, 1900) WC
A. tenebrosa (Casey, 1900) AK AB ON QC NB B WC NA MP
A. melas (Casey, 1900) syn. nov.
A. thujaplicatae Pelletier, n. sp. BC MW
A. umbrina (Gyllenhal, 1827)† BC ON QC NB B WC NA MP
A. undulata (Casey, 1900) AK BC AB ON QC NB B MW WC NA MP
A. cribripennis (Casey, 1900) syn. nov.
A. castanea (Casey, 1900) syn. nov.
A. versa Grouvelle, 1916 AK B WC
A. soror (Casey, 1900), preoccupied by A. soror Gauglbauer
A. vespertina Mäklin, 1853* AK BC AB MB ON QC B MW NA
A. visoni Pelletier, n. sp. BC AB MW WC
Atomaria nomem dubium
A. lepidula Mäklin, 1852 AK
Genus CURELIUS Casey, 1900
Curelius japonicus (Paykull, 1798)† SE
Ephistemus perminutus Casey, 1924 syn. nov.
E. punctatus Blatchley, 1925
Genus EPHISTEMUS Stephens, 1829
E. cactophilus Schwarz, 1899
E. globulus (Paykull, 1798)† BC ON QC NS LB NF MW WC NA MP SE
E. apicalis LeConte, 1863
Genus TISACTIA Casey 1900
T. subglabra Casey, 1900 ON QC MP SA SE

1Key to Nearctic species: Woodroffe and Coombs (1961)

DNA Barcoding

The DNA barcoding methodology used in this publication was designed by Dr. Paul Hebert from the University of Guelph (Ratnasingham and Hebert 2013) and is explained in detail in Milton et al. (2013).We submitted 76 specimens of Cryptophagidae to the Canadian Centre for DNA Barcoding at the University of Guelph. Most of these belonged to the genera Cryptophagus and Atomaria, including 12 paratypes of 8 recently described species, to confirm their identity as distinct species. The Search Data tool was used to search all DNA-barcoded specimens of Cryptophagidae in public records and our project’s databases. In order to build the genetic sequence barcoding, we selected specimens containing sequences from the mitochondrial gene COI-5P with the following criteria:

    Collected in Canada and the US between 1998 and 2018.For Holarctic or adventive species, European specimens available to compare DNA barcoding with Nearctic specimens.Specimens with images that can be identified at the species level included in the BOLD database.Minimum sequence length of 500 bp, giving priority to Full Length Barcode specimens of 640 bp or more.Barcode-compliant specimens.

The resulting list of specimens was classified from the longest sequence (658 bp) to the shortest (500 bp). Specimens can be clustered together and identified with a Barcode Index Number (BIN), which shows a high concordance with species identification. Many specimens wrongly identified in the BOLD database have been corrected and up to 92 species have a valid DNA barcoding corresponding to 65% of Cryptophagidae species of this project. At the bottom of the descriptive page of each species having a BIN, there is a link leading to the BOLD database BIN page.

We produce dendrograms with the Taxon ID tree engine using the nucleotide as a coding marker, the Kimura 2 Parameter model and the BOLD aligner that seems to be more relevant to discriminate species within large genera and to discriminate genera within Cryptophagidae. A dendrogram of clustered species can be seen in the Taxon ID Tree 1 (Supplementary Materials) for representatives of most genera of Cryptophagidae, Taxon ID Tree 2 (Supplementary Materials) for Cryptophagus and Taxon ID Tree 3 (Supplementary Materials) for Atomaria.

The DNA barcoding sequence from the mitochondrial gene COI-5P shows that this method can separated correctly 91% of morphologically distinctive species, including 75% for Cryptophagus and 94% for Atomaria. This reflect the highly intraspecific variability found within Cryptophagus compared with Atomaria.

Taxon ID tree 1 separated most genera well. All Atomaria are grouped togetherand Curelius is distinctly outside of Atomaria.All Cryptophagus are grouped togetherwith Myrmedophila and Henoticus at both ends. Myrmedophila seems to be a synonym of Cryptophagus, following Woodroffe & Coombs (1961) concept,according to the ID tree; however, the DNA sequence was incomplete, with COI-5P of 463 bp, much less than the usual 658 bp. Surprisingly, Salebius is incorporated within Cryptophagus though it is a distinct genus. Antherophagus, Pteryngium, Caenoscelis Henotiderus and Telmatophilus are well separated at the opposite end of the tree.

Taxon ID tree 2 shows a visually easily Nearctic identified species (C. mainensis)in the upper part, followed by three introduced species with toothed laterally prominent callosities without dorsal rim (C. distinguendus to C. scutellatus), followed by a species with callosities strongly produced anteriorly (C. tuberculosus), three species with elytra arcuate throughout and pubescence mostly suberect (from C. setulosus to C. valens), 6 mostly Palaearctic species with elytra mostly oblong and pubescence appressed, followed by very common Nearctic species (C. croceus and C. difficilis) and finally three common Holarctic northern transcontinental species (from C. lapponicus to C. bidentatus). The tree has shown (not illustrated) that many specimens identified as C. jakowlewi in Europe were in fact C. confertus. Of the 2 specimens seen by the first author and identified by Colin Johnson as C. jakowlewi, one was C. confertus and the other C. bidentatus. Esser (1994) showed clearly that all three species were distinct. The true C. jakowlewi has not been collected in North America and C. confertus remains a valid Holarctic species. Among species having two BIN numbers, we can mention C. tuberculosus, C. setulosus, C. dentatus and C. difficilis, all very common and widespread species.

Taxon ID tree 3 shows that most Atomaria species are well discriminated by the analysis, with all Anchicera (from A. mesomela to A. neomunda) being on one side and all Atomaria s. str. (A. nigrirostris to A. subdentata) being on the opposite side. We can see also some distinct groups. In Anchicera,some are more distinct as species with pronotum arcuate at middle and strong contrast black pattern at basal half of elytra (A. mesomela, A. distincta); antennae with A9-A10 transverse (from A. peltata to A. apicalis), within that group, dark species with pronotum borders entirely visible from dorsal view (A. nigritaria, A. peltata), and pronotum with sides subparallel at basal 0.5 (A. turgida, A. apicalis); the A. fuscata complex species group (from A. hudsonica to A. ocularia), antennae with A9-A10 subquadrate and pronotum border entirely visible from dorsal view (from A. kamtschatica to A. arcuaticollis) and pronotum with double-sided borders (from A. ornata to A. neomunda). In Atomaria s. str., we can see at first species with pronotum having strong basal groove (A. nigrirostris, A. impressicollis), with A9-A10 transverse and pronotum with sides not sinuate (A. tenebrosa, A. patens), A9 subquadrate and A10 transverse (A. pumilio, A. wollastoni), elytra with pubescence suberect (A. umbrina), pronotum subquadrate and elytra elongate with sides subparallel (A. linearis), A9 and A10 nearly subquadrate (A. vespertina, A. nigricollis) and A1 short and stout (from A. macra to A. subdentata). The tree also shows that the European concept of A. fuscata might be in fact three distinct species (one not included in this tree) that were synonymized probably by Johnson but this need to be verified. In the present publication, A. fuscata corresponds to the wide Johnson concept of the species (Johnson 1992). It would be very useful to look to all distinct Palearctic BIN species to see the external morphological characters that can separate them. It might prove that A. fuscata (= A. saginata) and A. ochracea are distinct species.

The analysis also shows that the A. wollastoni complex is in fact three species: A. wollastoni which is Palaearctic, A. pumilio and A. lineola, both being Nearctic, previously considered as synonyms of A. wollastoni. The tree also shows that A. pulchra was previously confused with A. vespertina. The analysis confirms the validity of 7 new species described in this publication (all with Pelletier as the author): A. arcuaticollis, A. neomunda, A. nigritaria, A. albertana, A. impressicollis, A. pinicola and A. puelloides. Three new species were discovered and described because of their distinct BINs (all with Pelletier as the author): A. ocularia, A. salicicola and A. calidaria, all from Alberta.

Taxonomy

Family CRYPTOPHAGIDAE Kirby 1837(Mostly from Leschen & Skelley (2002))

Description: Length 0.8 to 5.2 mm, mostly between 1.2 and 3.0 mm. Body robust, oval or elongate-oval, moderately flattened or convex, rarely round; body colour often reddish brown, dark brown with elytra yellowish brown, rarely entirely dark brown to black, usually with moderately long to short, erect to decumbent silky pubescence; glandular ducts present in various parts of the body, probably for the secretion of fluids that impermeabilize the body against the high humidity found in their secretive habitat. Head partially retracted into thorax, longer than wide, usually without antennal groove; eyes usually entire, finely faceted; ocelli absent; genae carinate and projecting ventrally between eye and mentum; clypeus usually subquadrate; mandible with well developed mola, incisor apex and prostheca; maxilla with brushlike galea and lacinia, probably for collecting spores; labium with mentum wider than long; maxillary palpi 3-segmented; labial palpi 2-segmented. Antennae moderately long, usually with a 3-segmented club.Pronotum with distinct lateral carina (except in Hypocoprus); posterior margin usually narrower than anterior margin of elytra, often with depressions at the base; hypopleuron usually without antennal grooves; prosternal process well developed. Scutellum well developed and visible. Mesocoxal cavities closed laterally by metaventrite; metaventrite usually elongate. Legs moderately short and slender in most species; procoxae spherical and separate; trochanter about 25% length of femur; tibia usually with 2 apical spines; tarsi usually 5-5-5 or 5-5-4 in male Cryptophaginae.Elytra usually completely covering the abdomen, without impressed striaepunctation confused, usually not forming distinct rows, (except underneath in many Atomaria species) and apex with double suture; epipleuron usually distinct in anterior half, never reaching the apex.Abdomen with 5 freely articulate ventrites; first ventrite longer than the second ventrite; intercoxal process moderately broad. Male genitalia usually bilaterally symmetrical in most species.Habitat. Most species live in concealed wet habitats with decaying vegetation containing fungi. They are very common in leaf litter, humid soil and the bark of old trees on mold and fleshy fungi where they are microphagous, feeding on the mycelium, hyphae, conidia or spores. A few others are saprophagous. Many species, mostly introduced from Europe, can be found in stored food products, but feed only on mold. Some specialized species are usually collected in bird nests, ant nests and mammal burrows. Four species are found in the nests of bees and wasps. They are probably scavengers. Two species are associated with aquatic plants.Seasonality. They emerge in the spring or summer and some adults can be active up to late fall. It is interesting to note that the Southeastern US species we found in Canada are more active between November and April. It shows that temperature and humidity are very important to stimulate the activity of those species. They are predominantly active during the evening and at night.Collecting methods. Many species living in leaf litter can be collected in great numbers by sifting the litter and using a berlese or with light or pitfall traps. Those living in tree bark can be collected with a Lindgren funnel or with flight interception traps. A few species may be swept or beaten from the foliage. Other specialized species can be collected by hand. Rarely, a few species can be collected on carrion.Similar species in other families often confused with CryptophagidaeMost species often confused with Cryptophagidae belong to Erotylidae in the following small subfamilies (Figure 2):

    • Cryptophilinae Toramini

Toramus pulchellus 

    • (LeConte):Very similar to some

Atomaria (Anchicera) 

    • but pronotum opaque with pair of depressions at base and elytra with pilosity minute and sparse.Cryptophilinae Cryptophilini

Cryptophilus integer 

    • (Heer):Similar to few

Atomaria 

    • but pronotum as wide as elytra at base with side borders visible throughout and two disconnected foveae at base.

Cryptophilus seriatus 

    • Casey:Very similar to some

Henoticus 

    • but pronotum widest slightly in front of middle with serrulation smaller and two disconnected foveae at base and elytra with punctation in dense rows.Loberinae

Loberus 

    • species: Pronotum deeply impressed at basal middle and elytra with seven well-spaced rows of punctures.Cerylonidae Euxestinae

Hypodacne punctata 

    • LeConte:Superficial similarity with

Ephistemus 

    • and

Tisactia 

    but antennal club with antennomeres not loose and elytra without distinct pilosity.

Figure 2. Species in other families similar to Cryptophagidae.

Diagnoses and summaries for Cryptophaginae

Subfamily CRYPTOPHAGINAEFrontoclypeal suture absent; pronotum with lateral carina present; meso-metaventral articulation with a double knob; tarsal formula 5-5-5 in female and 5-5-4 in male; male genitalia with speculum gastrale with long anterior strut, parameres mostly separate.

Tribe CRYPTOPHAGINIAntennae remotely separate at base, inserted under sides of frons; labial palpi 2- segmented with basal palpomere subequal to apical palpomere; pronotum with lateral margin often serrate, dentate, with or without a callosity at anterior corners, without sublateral carinae and with distinct basal pits or grooves.

Antherophagus Dejean 1821Pronotum with sides entire, without callosities at front corners and without lateral tooth at middle; head with a distinct clypeal notch in male. Widely distributed. Associated with flowers and bee or bumblebee nests. Fairly common. Four species in North America, all found in Canada.Cryptophagus Herbst, 1792Pronotum with distinct callosities at front corners and a distinct tooth near middle of sides; pilosity of elytra long, appressed, recurved or suberect. Widely distributed. Associated with forest leaf litter, rotting wood, also in food products stored in wet areas with high moisture, sometimes in ant nests or animal burrows. Very common. 38 species in North America, with 32 species in Canada, including 11 Nearctic, 7 Holarctic and 14 adventive, the majority of the latter group uncommon in natural habitats.

Henoticus Thomson, 1868Body not dorsoventrally compressed; pronotum with sides distinctly serrate, without sublateral lines, serratures strong and recurved, with base deeply bifoveate, foveae connected by wide groove along basal margin; mesoventral process equal in width to mesocoxa; anterior lateral angle of metaventrite with a pubescent pit; elytra reddish to yellowish brown, rarely black, with sparse to moderately dense pubescence; legs with tarsomeres not strongly lobed beneath. Widely distributed. Associated with forest leaf litter, rotting wood, fungi and food products stored in wet areas with high moisture. Common. Four species in North America, all present in Canada, including two Nearctic, one Holarctic and one adventive.

Henotiderus Reitter, 1877Body not dorsoventrally compressed; pronotum with sides distinctly serrate, with sublateral lines, serratures strong and recurved, with base deeply bifoveate, foveae not connected by wide groove along basal margin. Mostly transcontinental in Canada and northern US. Associated with fungi under bark of rotting wood. Fairly common. Two species in North America, both in Canada; one Nearctic and one Holarctic.

Micrambe Thomson, 1863Pronotum sides with callosity covering anterior half (most Palearctic species have a shorter callosity covering anterior third or less), middle tooth lacking. One record in North America (in Quebec, Canada). Associated with Black Spruce trunk burned with fire. Rare. One species introduced from Europe: Micrambe ulicisFirst record of the genus in North America.

Myrmedophila Bousquet, 1989Pronotum with distinct callosities at front corners and a distinct tooth near middle of sides, sides with posterior half subparallel; pronotum and elytra with pilosity very short, appressed, setae shorter than distance between punctures. Transcontinental in Canada and northern US. Associated with ant nests of genus Formica. Uncommon. One native species in North America, occurring in Canada: Myrmedophila americana.

Pteryngium Reitter, 1887Body dorsoventrally compressed; pronotum with sides distinctly serrate, without sublateral lines, serratures minute and not recurved, base with two small basal disconnected foveae; mesosternal process greater in width than mesocoxa; anterior lateral angle of metaventrite with a slight depression; elytra reddish to yellowish brown; legs with tarsomeres not strongly lobed beneath. In transcontinental Canada, Alaska and northern US. Associated mainly with conifer forests, sometimes in hardwood forests, in fungi and polypores. Uncommon. One Holarctic species in North America, in Canada: Pteryngium crenulatum.

Salebius Casey, 1900Pronotum with smooth sides consisting of three distinct lobes. In western North America, from Alaska south to California. Associated with conifer forests, in trees under bark. Fairly common. Two species in North America, one occurring in Canada; one native and one Holarctic.

Telmatophilus Heer, 1841Body entirely dark brown to black, densely clothed with silvery pubescence; pronotum with sides distinctly serrate, without sublateral lines, serratures minute and not recurved; legs with tarsomeres 2 and 3 strongly lobed beneath. In transcontinental Canada and northern US. Associated with aquatic plants of genera Spargania and Typha. Fairly common. Two species in North America, both in Canada; one native and one adventive.

Tribe CAENOSCELINI

Caenoscelis ThomsonAntennae closed together at base, inserted in frons; labial palpi 2-segmented, with basal palpomere wider than apical palpomere; pronotum with lateral margin smooth, with sublateral carinae, and without distinct basal pits or grooves; ventrite V of abdomen with modified setae. Widely distributed, but more common in transcontinental Canada, northern and western USA, extending south in the Appalachian Mountains. Very common. Mostly in forest leaf litter. Six species in North America, all found in Canada, including four natives and two Holarctic species.Subfamily ATOMARIINAEFrontoclypeal suture present; labial palpi 2-segmented with basal palpomere wider than apical palpomere; meso-metasternal articulation with lateral processes; tarsal formula 5-5-5 in both sexes; male genitalia broad, parameres fused.

Tribe HYPOCOPRIINI

Hypocoprus Motschulsky 1839Pronotum nearly cylindrical, longer than wide, with lateral carina lacking; elytra very elongate, about 1.75x as long as wide, not covering the last tergite. In northwestern Canada. Associated with ant nests of genus Formica. Uncommon. One species in North America, found in Canada, Holarctic: Hypocoprus latridioides.

Tribe ATOMARIINI Le ContePronotum with lateral carina present; elytra variable, usually 1.7x as long as wide or less, covering the last tergite. Glandular ducts present on various parts of the body.

Atomaria Stephens 1829Head distinctly visible anterior to the pronotum. Pronotum usually widest from anterior 30 to 85% (except for 2 species). Elytra with sides forming a discontinuous line with the pronotum, distinctly punctate and hairy. Widely distributed. Associated with forest leaf litter, rotting wood, also in food products stored in wet areas with high moisture, sometimes in ant nests or animal burrows. Very common. 80 species in North America, with 75 species in Canada, including 52 Nearctic, 14 Holarctic and 14 adventive, many of the latter group being well established in natural habitats. To facilitate the understanding of this complex genus, all species were separated into different species groups, detailed as follows. This genus is divided into two major subgenera:

Subgenus Anchicera Thomson 1863Elytra with sides arcuate throughout, usually about 1.35-1.5x as long as wide, often with uniform colouration (70% of the species), sometimes patterned (30%); antennae with A1 short and stout (95%), club with A9-A10 subquadrate (61%) or transverse (39%). Pronotum usually with a single side border (90% of the species), with sides distinctly converging posteriorly (84%) or subparallel (16%). Widely distributed. Very common. 35 species in North America, with 34 species in Canada, including 18 Nearctic, 8 Holarctic and 9 adventive. This subgenus can be divided into seven species groups:

Anchicera ornata sp. gr.: Pronotum with double-side borders along most of its length. In transcontinental Canada and northern US. Uncommon. Three species, all adventive.Anchicera pusilla sp. gr.: Pronotum with sides distinctly converging posteriorly; antennal club with A9-A10 transverse. Widely distributed. Very common. Six species, with two newly described Nearctic, two Holarctic and two adventives.Anchicera kamtschatica sp. gr.: Pronotum with sides distinctly converging posteriorly, with both side borders completely visible from dorsal view; antennal club with A9-A10 subquadrate. In transcontinental Canada and Alaska. Common. Four species, with three Nearctic including two newly described species and one Holarctic.Anchicera atra sp. gr.: Pronotum with sides distinctly converging posteriorly, with both side borders visible within posterior 50% only from dorsal view; antennal club with A9 subquadrate, A10 subquadrate or transverse; elytra unicolor, without distinctive pattern. Widely distributed. Very common. Nine species, with six Nearctic including three newly described species, two Holarctic and one adventive.Anchicera mesomela sp. gr.: Pronotum with sides distinctly converging posteriorly, with both side borders visible within posterior 50% only from dorsal view; antennal club with A9-A10 subquadrate; elytra bicolor, with distinctive pattern. Widely distributed. Very common. Five species, with three Nearctic, one Holarctic and one adventive.Anchicera testacea sp. gr.: Pronotum with sides distinctly subparallel at posterior 0.5; antennal club with A9-A10 transverse. Widely distributed. Common. Five species, with one Nearctic, two Holarctic and two adventives.Anchicera nubipennis sp. gr.: Pronotum with sides distinctly subparallel at posterior 0.5; antennal club with A9-A10 subquadrate. Mostly found in western North America. Uncommon. Three species, all Nearctic, including one newly described species.

Subgenus Atomaria Stephens 1829Elytra with sides mostly straight, at least at basal 35%, slightly convergent at basal 35-60% to nearly parallel at basal 65%, usually with uniform colouration (91% of the species) and pubescence decumbent or appressed (88%); antennae with A1 elongated and recurved (70%) or short and stout (30%), club with A9-A10 usually transverse (71%) or subquadrate (29%); pronotum with sides distinctly converging posteriorly (84%) or subparallel (16%). Widely distributed. Very common. 45 species in North America, with 42 species in Canada, including 34 Nearctic, 6 Holarctic and 5 adventive. This subgenus can be divided into 10 species groups:

Atomaria fimetarii sp. gr.: Antennae with A11 distinctly narrower than A10; body cylindrical, very convex, pronotum stout, as broad as elytra. In transcontinental Canada and northern US. Uncommon. One Holarctic species: A. fimetarii.Atomaria nigrirostris sp. gr.: Pronotum with sharply defined basal transverse groove restricted to middle third, sometimes extending to sides. Widely distributed. Mostly uncommon, except the very common A. nigrirostris. Many species specialized in animal burrows or nests. Six species: four Nearctic including two newly described species, one Holarctic and one adventive.Atomaria umbrina sp. gr.: Antennae with A6-A10 strongly transverse. In transcontinental Canada. Uncommon. Two species, including one newly described Nearctic species and one adventive species: A. umbrina.Atomaria puella sp. gr.: Elytra with pubescence sparse and suberected. In central and western North America; one species in southeastern US. Uncommon. Apparently specialized in ant nests. Three species, all Nearctic, including one newly described species in Georgia.Atomaria linearis sp. gr.: Body elongate, pronotum subquadrate, about 1.15x as wide as long, elytra about 1.8x as long as wide. In western US. Rare. Pest species associated with sugar beets. One adventive species introduced into California: A. linearis.Atomaria elongatula sp. gr.: Antennae with A1 short, about 1.3x as long as its apical width, and about 1.2-1.5x as long as A2. Mostly in transcontinental Canada, Alaska, northern and western US, extending south in the Appalachian Mountains. Fairly common. Mostly under the bark of trees, possibly many species specialized in bird nests. Nine species: with seven Nearctic including four newly described species, and two Holarctic.Atomaria crassula sp. gr.: Antennae with A1 elongated and recurved, about 1.5x as long as its apical width and more than 1.5x as long as A2; club with A9-A10 subquadrate. Mostly in transcontinental Canada, Alaska, northern and western US. Fairly common. Mostly under bark of conifers trees. Three species, all Nearctic.Atomaria vespertina sp. gr.: Antennae with A1 elongated and recurved, about 1.5x as long as its apical width and more than 1.5x as long as A2, club with A9 subquadrate to slightly transverse and A10 slightly to distinctly transverse. Mostly in transcontinental Canada, Alaska, northern and western US, extending south in the Appalachian Mountains. Uncommon, except the common A. lineola. Mostly under bark of conifer and hardwood trees. Five species: three Nearctic, including one newly described species, one Holarctic and one adventive.Atomaria alpina sp. gr.: Antennae with A1 elongated and recurved, about 1.5x as long as its apical width and more than 1.5x as long as A2, club with A9-A10 distinctly transverse; pronotum with sides arcuate at anterior 50-70%, widest posterior to middle, not sinuate at middle. Mostly in transcontinental Canada, Alaska, northern and western US, extending south in the Appalachian Mountains. Uncommon. Mostly under bark of conifer and hardwood trees. Six species: five Nearctic, including two newly described species and one adventive.Atomaria affinis sp. gr.: Antennae with A1 elongated and recurved, about 1.5x as long as its apical width, more than 1.5x as long as A2, club with A9-A10 distinctly transverse; pronotum with sides sinuate at middle. Mostly in transcontinental Canada, Alaska, northern and western US extending south in the Appalachian Mountains. Fairly common. Mostly under bark of trees of conifers and hardwood. Ten species: nine Nearctic, including two newly described species, and one Holarctic.

Curelius Stephens, 1829Head mostly concealed under the pronotum; pronotum widest at base; elytra with sides forming a continuous line with the pronotum; body yellowish red, about 1.65-1.75x as long as wide, with punctuations small but distinct. In southeastern US, not yet recorded in Canada. In hardwood trees, mainly oak forests. Very common. One introduced species: Curelius japonicus.

Ephistemus Stephens, 1829Head mostly concealed under the pronotum; pronotum widest at base; elytra with sides forming a continuous line with the pronotum; body reddish to dark brown or black, often paler apically, broadly oval, about 1.5-1.6x as long as wide, with punctation minute, almost indistinct. Widely distributed. Indoor or outdoor species, often associated with lawns, one species associated with cactus. Fairly common. One Nearctic species in southwestern US and one adventive species: Ephistemus globulus.

Tisactia Casey, 1900Head mostly concealed under the pronotum; pronotum widest at base; elytra with sides forming a continuous line with the pronotum; body oblong-oval, about 1.8x as long as wide, with punctuations small but distinct. In southeastern Canada and eastern US. Rare. One Nearctic species: Tisactia subglabra.

Species Keys

Acknowledgments

We thank the following persons for their contribution to the present publication:
Colin Johnson, world specialist who has been working on the family since the 1960s, for providing to us a useful key to identify most European species of Atomaria. This key was tremendously helpful for building the Nearctic key and identifying the main characters that can help separate each species. He provided us with an updated unpublished checklist of Palearctic species, which also included Holarctic species. He put us in contact with the Manchester Museum, where he has worked most of his life.

Richard Leschen, from the New Zealand Arthropods Collection at Landcare Research, for kindly revising this manuscript. He collaborated closely with Colin Johnson and is the main contributor to the unpublished world checklist of Coleoptera. Both men helped build the foundation of the present work.

Floyd Shockley, from the National Museum of Natural History of the Smithsonian Institution, Washington, DC, for providing holotypes and paratypes of 82 species of Cryptophagidae, many being synonymized.

Lech Borowiec, from the University of Wroclaw’s Insect Collection, Poland, for providing images of 55 species of Palearctic Cryptophagidae, 20 of them serving as illustrations in this publication.

Dmitri Logunov, curator of the Manchester Museum in England, for sending specimens of 50 Palearctic species for comparison with Nearctic specimens.

Yves Bousquet and Patrice Bouchard, from Agriculture and Agri-Food Canada’s Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, ON, for providing 5887 specimens.

Claude Chantal, private beetle collector from Varenne, Québec, for providing 1424 specimens.

Colin Favret and Étienne Normandin, from the Collection entomologique Ouellet-Robert of the Centre sur la biodiversité de l’Université de Montréal, Québec, for providing 577 specimens.

Joseph V. McHugh and Richard Hoebeke, from the University of Georgia Collection of Arthropods, Athens, Georgia, for providing 517 specimens.

Derek Sikes, from the University of Alaska Museum Insect Collection, Fairbanks, Alaska, for providing 486 specimens.

Stéphanie Boucher, from the Lyman Entomological Museum, McGill University, Ste-Anne-de-Bellevue, Québec, for providing 474 specimens.

Mario Fréchette and Marie-Chantal Émond, from the Collection d’insectes du Québec, Ministère des Forêts, de la Faune et des Parcs, Québec, Quebec, for providing 350 specimens.

Dave Langor and James Hammond, from the Northern Forestry Centre, Canadian Forest Service, Edmonton, Alberta, for providing 337 specimens.

Karen Needham, from the University of British Columbia’s Spencer Entomological Collection, Vancouver, British Columbia, for providing 252 specimens.

Reggie Webster, collector, from the Canadian Forest Service, Atlantic Forestry Centre, Fredericton, New Brunswick, for providing 126 specimens.

Steve Marshall and S.M. Paiero, from the University of Guelph, Ontario, for providing 106 specimens and one photograph.

Audrey Malachowsky, from the Staten Island Museum, New York, for providing holotypes of two species of Atomaria.

Crystal Maier, from the Field Museum of Natural History, Chicago, Illinois, for providing the paratype of Glyptophorus mycetoecus.

Jaakko Mattila, from the Finnish Museum of Natural History, Helsinki, Finland, for providing specimens of five species of northern Europe, some known to occur in North America.

Jean-Marie Perron, from the Collection d’insectes de l’Université Laval, Québec, Quebec, for providing 96 specimens, mostly from the Léon-Provancher Historical Collection, dating from the 19th century.

Allison Brown, Sean Prosser and Tyler Elliott, from the BOLD team of University of Guelph, Ontario, for producing the DNA barcode of many species of Cryptophagidae, and Megan Milton, also from the BOLD team, for her help with BOLD tools to analyze the genetic sequences of many species.

Philippe Villemaire and André Beaudoin, from the Laurentian Forestry Centre, Canadian Forest Service, Québec, Quebec, for their help with the software ArcGIS, Don Stewart, who helped us prepare the material for the DNA analysis and Isabelle Lamarre, from the same Centre, for kindly revising the introduction of the manuscript.

Frank and Waltraud Koehler, from the Koleopterologie site which has very nice photographs of live Palearctic beetles. We have used photos, with permission of the photographers, from this website: F. Koehler (3), I. Altmann (5), A. de Wilde (1), and J. Dvorak (1). (http://www.koleopterologie.de/gallery/)

Smithsonian Institution, for providing the photographic system, including a high-quality digital camera for taking photos of specimens.

Heather Proctor, Editor-in-Chief of the Canadian Journal of Arthropod Identification, Morgan Jackson and Robert Anderson, responsible for the Coleoptera manuscript revisions.

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Cite

Pelletier, G., Hébert C. 2019. The Cryptophagidae of Canada and the northern United States of America. Canadian Journal of Arthropod Identification 40: 305 pp. doi:10.3752/cjai.2019.40

Supplemental Materials

Additional Materials and Methods