Systematics, Distributions, and Bionomics of the Neoeocatopsgen. Nov. and Nemadus of North America (Coleoptera: Leiodidae: Cholevinae: Anemadini)

Systematics, distributions, and bionomics of the Neoeocatopsgen. nov. and Nemadus of North America (Coleoptera: Leiodidae: Cholevinae: Anemadini)

Authors: Peck, Stewart B., and Cook, Joyce Source: The Canadian Entomologist, 139(1) : 87-117 Published By: Entomological Society of Canada URL: https://doi.org/10.4039/n05-091

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Downloaded From: https://bioone.org/journals/The-Canadian-Entomologist on 26 Jun 2020 Terms of Use: https://bioone.org/terms-of-use Access provided by Carnegie Museum of Natural History 87 Systematics, distributions, and bionomics of the Neoeocatops gen. nov. and Nemadus of North America (Coleoptera: Leiodidae: Cholevinae: Anemadini)

Stewart B. Peck,1 Joyce Cook Department of Biology, Carleton University, Ottawa, Ontario, Canada K1S 5B6

Abstract—This paper completes a review and revision of the tribe Anemadini in North Amer- ica, covering the subtribe Eocatopina (new for North America), with the genus Neoeocatops gen. nov. containing one species, and the subtribe Nemadina, with the genus Nemadus Thomson containing 12 species. Species are forest inhabitants feeding on carrion and other decaying materials, may occur in caves, or are myrmecophiles occupying the nests of formicine ants, mostly of the genera Formica L. and Camponotus Mayr. The following new synonyms are recognized: N. ellipticus Jeannel, 1936 = N. horni Hatch, 1933 and N. obliquus Fall, 1937 = N. triangulum Jeannel, 1936. Lectotypes are designated for Catops brachyderus LeConte, 1863, Catops pusio LeConte, 1859, and Choleva decipiens Horn, 1880. New species are Nemadus browni Peck and Cook, sp. nov., N. criddlei Peck and Cook, sp. nov., and N. falli Peck and Cook, sp. nov.

Résumé—Ce travail complète une revue et une révision de la tribu Anemadini (Coleoptera : Leiodidae : Cholevinae) d’Amérique du Nord; il traite de la sous-tribu Eocatopina (nouvelle pour l’Amérique du Nord) qui contient le genre Neoeocatops gen. nov. avec une espèce et de la sous- tribu Nemadina qui comprend le genre Nemadus Thomson, 1867 avec 12 espèces. Ces espèces vivent dans les forêts et se nourrissent de charogne et d’autres matériaux en décomposition; elles peuvent aussi se retrouver dans les cavernes ou alors être des myrmécophiles vivant dans les nids de fourmis formicinées, surtout des genres Formica L. et Camponotus Mayr. Nous établissons les nouvelles synonymies suivantes : N. ellipticus Jeannel, 1936 = N. horni Hatch, 1933 et N. obliquus Fall = N. triangulum Jeannel, 1936. Nous désignons des lectotypes pour Catops brachyderus LeConte, 1863, Catops pusio LeConte, 1859 et Choleva decipiens Horn, 1880. Les nouvelles espèces sont Nemadus browni Peck et Cook, sp. nov., N. criddlei Peck et Cook, sp. nov. et N. falli Peck et Cook, sp. nov.

[Traduit par la Rédaction]

Peck117 and Cook Introduction usually noticeably smaller than segments 7 and 9. Several of the subfamilies have previously The beetle family Leiodidae is composed of been ranked as separate families. The species of six subfamilies of small-sized beetles with dis- one of these leiodid subfamilies, the Chole- tinctive morphologies and diverse feeding hab- vinae, are mostly generalized scavengers, usu- its. Worldwide, some 340 genera and over 3450 ally occurring on or near decaying animal species are described. They share the unique carcasses. They can be collectively called “the derived character of having some or all of antennal segments 7 to 10 with a nearly en- small carrion beetles”. They have previously closed apical periarticular gutter, and usually often been ranked as a family, under the name with internal vesicles opening into the peri- Leptodiridae or Catopidae. articular gutter (Newton 1998). A very useful Within the subfamily Cholevinae there are six character for quickly recognizing most mem- tribes, four of which occur in North America bers of the family is that antennal segment 8 is from the Mexican Plateau northwards. These

Received 24 October 2005. Accepted 21 September 2006. 1Corresponding author (e-mail: [email protected]).

are Leptodirini, with two described and one un- CASC California Academy of Sciences, San described species in the genus Platycholeus Francisco, California (D. Kavanaugh) Horn in the Pacific Northwest; Ptomaphagini, CCC Claude Chantal Collection, Québec, with many generally distributed species (Peck Quebec 1973, 1977; Perreau 2000; Peck and Skelley CDAE California Department of Agriculture, 2001); Cholevini, with four genera and 22 spe- Sacramento, California (F.G. Andrews) cies (Peck and Cook 2002); and Anemadini. CMNC Canadian Museum of Nature, Gatineau, The tribe Anemadini has been divided into five Quebec (F. Genier and R.S. Anderson) subtribes, of which three occur in North Amer- CNCI Canadian National Collection of In- ica. These are Eunemadina, with Dissochaetus sects, Agriculture Canada, Ottawa, On- Reitter containing three species (Peck 1999); tario (A. Smetana and A. Davies) Eocatopina, here recognized for the first time in CUIC Cornell University, Ithaca, New York North America, with Neoeocatops gen. nov. (V. Pechumen and J. Liebherr) containing one species; and Nemadina, with DENH Department of Entomology, University Nemadus Thomson containing 12 species. This of New Hampshire, Durham, New revision of the subtribes Eocatopina and Nema- Hampshire (D. Chandler) dina completes a review and revision of the EGRC Ed G. Riley Collection, College Sta- Anemadini of America north of Mexico. Addi- tion, Texas tional information on leiodid classification and FMNH Field Museum of Natural History, Chi- world diversity is given in Newton (1998) and cago, Illinois (A.F. Newton) Perreau (2000). FSCA Florida State Collection of Arthropods, Previous publications on the Anemadini in Gainesville, Florida (M.E. Thomas and North America are those by Horn (1880), Hatch P. Skelley) (1933, 1957), Jeannel (1936), Fall (1937), and HAHC Henry and Anne Howden Collection, Peck (1999). Major contributions to knowledge Ottawa, Ontario of the Anemadini fauna of the Palearctic and JCIC Joyce Cook Collection, North Augusta, Oriental regions (with 7 genera and 89 species) Ontario have been made by Jeannel and Szymcza- JJLC Julian J. Lewis Collection, Craw- kowski (references in Perreau 2000). Since the fordsville, Indiana previous publications on the North American JLCC J.L. Carr Collection, Calgary, Alberta species, considerable new material has accumu- (B.F. Carr) (now in CNCI) lated, which is used here to revise and docu- KHSC Karl H. Stephan Collection, Red Oak, ment the systematics, distributions, and Oklahoma (now in FSCA) bionomics of the North American fauna of LACM Los Angeles County Museum, Los An- Anemadini. A review and revision of the North geles, California (C.L. Hogue) American species of the genera Neoeocatops LSUC Louisiana State University, Baton and Nemadus are the purpose and subjects of Rouge, Louisiana (J. Chapin) this paper. MCZC Museum of Comparative Zoology, Harvard University, Cambridge, Mas- Materials sachusetts (A.F. Newton and P. Perkins) This study is based on the examination of MHNG Muséum d’Histoire Naturelle, Geneva, over 3112 specimens from the Nearctic Region, Switzerland (G. Cucodoro) most collected by the first author. Additional MNHN Muséum National d’Histoire Naturelle, specimens were borrowed for study from the Paris, France (Th. Deuve) following collections and curators or private MUIC Mississippi State University, State Col- collectors. Most collection addresses are given lege, Mississippi (T. Schiefer) in full in Arnett et al. (1997). MZLU Zoological Institute, Lund University, AFNC A.F. Newton Collection, Chicago, Illi- Lund, Sweden (M. Sorenson and R. nois Baranowski) AMNH American Museum of Natural History, ODAC Oregon State Department of Agricul- New York, New York (L.H. Herman) ture, Salem, Oregon (R.E. Westcott) BMUW Burke Museum, University of Wash- OSUC Ohio State University Collection, Co- ington, Seattle, Washington lumbus, Ohio (Q. Wheeler)

RTC R. Turnbow Collection, Fort Rucker, Mayr and Formica L. The beetles can be taken Alabama by opening the ant nests and searching in the nest SBMN Santa Barbara Museum of Natural His- galleries and litter. For nests of mound-building tory, Santa Barbara, California (M. ant species, the most productive way to collect Caterino) the beetles seems to be to use boards or flat rocks SBPC Stewart B. Peck Collection, Ottawa, with a rough undersurface laid on top of and Ontario around the nest in the spring. As the spring sun SEMC Snow Entomological Museum, Univer- warms the board or rock, the beetles are appar- sity of Kansas, Lawrence, Kansas ently attracted to the heat and come to cling to the (S. Ashe) underside of the board or rock, where they are SMDV Spencer Entomological Museum, Uni- found when it is lifted. The first mention of this versity of British Columbia, Vancou- method seems to be by Märkel (1841) and ver, British Columbia (S. Cannings) Mannerheim (1843). It seems to be widely used TAMU Texas A&M University, College Sta- in Europe, or at least in England (Walsh 1991). tion, Texas (E.G. Riley) We ourselves have not had success with this UASM Strickland Entomological Museum, method in southeastern Canada. Large samples University of Alberta, Edmonton, Al- from ant nests used in this study are available berta (G.E. Ball) thanks to the special collecting efforts of UCRC University of California at Riverside H.F. Wickham, F. Blanchard, H.C. Fall, and A.F. Entomological Collection, Riverside, Newton. California (D. Yanega) For lectotype and paratype specimens we re- UMIC University of Mississipppi Insect Col- port label data as it appears on the specimen la- lection, University, Mississippi (P.K. bels. We have not edited or altered these data Lago) for uniformity, but have quoted them to aid in UOIC Oregon State University, Eugene, Ore- recognition of type specimens seen by us. Other gon (G. Peters) label data are summarized and reported as gen- USNM United States National Museum of eralizations about distributions, habitats, and Natural History, Washington, D.C. (R. seasonality. Full label data are available from Gordon, D. Furth, W. Steiner) the authors. Not all specimen labels contained WSUC Washington State University, Pullman, full information on locality, date, or habitat. Washington (W.J. Turner) To confirm identifications it is often desirable or necessary to examine the aedeagus of male Especially large collections were gathered in specimens or the spermatheca of female speci- the extensive field-sampling programs of A.F. mens. The sexes are distinguished by the fact Newton and S.B. Peck. Duplicates from SBPC that males have broadly expanded basal pro- have been distributed to CMNC, CNCI, FMNH, tarsal segments, whereas females have all the FSCA, and MCZC. protarsal segments unexpanded (with the same diameter). Specimens were dissected after Methods being relaxed and removed from points or cards. Relaxing was achieved by immersion of Many of the specimens seen during this the specimen for an hour or more in a commer- study were taken at carrion or in carrion-baited cial household ammonia-based window- pitfall traps. Large populations have also been cleaning solution. The specimen was then dis- taken in the guano of insectivorous bats in sected in alcohol. The aedeagus or spermatheca caves in the southeastern United States. The was exposed, examined, and either allowed to next most productive sampling techniques were dry everted from the abdomen tip or glued to the use of flight-intercept traps in forests (Peck the mounting point (some aedeagi) or dehy- and Davies 1980) and nets mounted on a vehi- drated in alcohol and placed in euparal mount- cle driven slowly along forest roads at dusk ing medium on a small acetate-plastic (Peck and Cook 1992). A few specimens have microslide. The identification of individuals is been taken by other methods, and these are in- not always possible. It is sometimes necessary dicated in the habitats section of individual spe- to examine the elytra of both sexes for the pres- cies discussions. ence or absence of microsculpture. External char- Several species of Nemadus live in the nests of acters were examined with a stereomicroscope formicine ants, mostly of the genera Camponotus from 10× to 200× magnification. Illustrations

were made from structures mounted tempo- (LeConte, 1863) (paralleling the widened body rarily in glycerine jelly or on a euparal of Echinocoleus spp.) seems to be a possible microslide, or from dry structures. Details were morphological defensive specialization for life observed with a compound microscope and with ants. The lack of other morphological de- then added to outline illustrations made from fensive specializations suggests that the beetles digital photographs. Wherever possible, illus- are tolerated or ignored by the ants and can trations of the aedeagus include features of the therefore be called synoeketes or symphiles armature of the internal sac, either within the (Hölldobler and Wilson 1990, p. 471). The Pale- median lobe or in an everted position. It was arctic species Nemadus colonoides (Kraatz, not possible to evert the sac in all species. 1851) is also found occasionally in the nests of To determine the higher taxonomic place- ants of the genus Lasius Fabr. as well in the cav- ment of Horn’s “Nemadus” decipiens, which ity nests of birds and elsewhere (Jeannel 1936). did not seem to us to actually belong in Nema- The habit of being an ant-nest associate is best dus, but rather in the subtribe Eocatopina, the developed as a specialization of Nearctic Nema- following Palearctic and Oriental species of that dus species in the parasitus species group. subtribe were examined: Eocatops (s. str.) ophi- Nine North American species of Nemadus in dianus Szymczakowski, 1976; Eocatops (s. str.) the parasitus species group seem to live exclu- pelopsis (Reitter, 1884); and Eocatops (Eone- sively with ants of the subfamily Formicinae, madus) loebli Perreau, 1991. mostly of the genera Camponotus and Formica. Each species of beetle is known mainly from Ecology the nests of a single ant species, but some have been collected occasionally from the nests of All members of the subtribe Nemadina, as two or more ant species. However, each beetle larvae and adults, seem to feed as scavengers, species is usually prevalent in nests of only one usually in cool temperate, moist, and shaded or ant species. Therefore, each beetle species ap- dark environments. Some specimens are fre- pears to show host preference or specificity. quently collected at carrion and carrion-baited The fact that some species of Formica make pitfall traps. One species, Nemadus horni slaves of other species of Formica may account Hatch, 1933, is frequently found in bat guano in for the mixture of ant species with which some caves in the southeastern United States, but it is beetle species have been found. Detailed studies not a cave-limited species. The seasonality and of the beetles’ interactions with ants remain to habitat preferences of this species in four dis- be made. tinct habitats in southern Ontario were documented by Peck and Anderson (1985), and those All adults have fully developed flight wings in forests in New Hampshire and New Jersey and apparently disperse by flight to new ant were documented by Chandler and Peck (1992) nests. The nests of Camponotus spp. are usually and Shubeck et al. (1977), respectively. composed of galleries excavated in the wood of The North American species of Nemadus that rotted logs and stumps, but these ants may also are known to live with ants are assumed to be nest under stones and in soil. The nests of For- general scavengers on wastes, perhaps fecal ma- mica spp. may be under logs and rocks, or nests terial in the ant nests. This habit of occupying may be medium to large mounds of earth, up to ant nests is not unique to Nemadus within the 1.5 m or more across, usually kept free of sur- Leiodidae. At least 13 other genera of leiodids face vegetation, and located in forests or fields. live with ants in various parts of the world Table 1 summarizes the known ant hosts of the (Hölldobler and Wilson 1990, p. 480). Notable beetles and their general nest structure and geo- in North America are four species of Ptoma- graphic distribution. phagus Illiger in the subgenus Echinocoleus When beetles are taken with ants, the ants found with ants of the genus Pogonomyrmex can be identified to subfamily with the use of Mayr in the southern and western United States several entomology textbooks, to genus with (Peck and Gnaspini 1997). It is not known how the use of the keys in Hölldobler and Wilson the ants interact with or react to the beetles. (1990), and to species with the now outdated There are no trichomes or unusual morphologies keys in Creighton (1950), in which some of the to suggest that the ants interact in a positive way species-level nomenclature has changed. Ant with the beetles, such as in an appeasement role. nomenclature used here follows Bolton (1995) Only the widened body of Nemadus brachyderus and Hedlund (2003).

Table 1. General data on ants with which North American species of Nemadus have been collected; data abstracted from Smith (1979) and Hedlund (2003). Distribution limits, using US Name of ant and its Nemadus guest Postal Service abbreviations species Most common nest locations for states and provinces Acanthomyops claviger (Roger), the In woods, pastures, and open MA to ON, south to FL, smaller yellow ant; minor host for fields; occasionally in ex- west to MN, KS, MS N. parasitus posed soil, under stones, in rotting logs and stumps Camponotus chromaiodes Bolton In and under stumps and rotted MA to GA, west to MN and (replacement name for junior logs, with galleries into soil KS homonym C. ferrugineus (Fabr.)), the red carpenter ant; alternate host for N. triangulum C. herculeanus (L.); secondary host Dominant in boreal and alpine NF to AK, south to PA, WI, for N. brachyderus North America; in logs and NM stumps, especially conifers C. noveboracensis (Fitch); principal In forests; in rotted logs and NS to VA, west to BC and host for N. brachyderus; alternate stumps UT host for N. triangulum C. pennsylvanicus (DeGeer), the black In dead and live trees, rotted NB to FL, west to ND and carpenter ant; alternate host for logs and stumps, in buildings NM N. triangulum; minor host for N. parasitus Formica exsectoides Forel, the Alle- Large mounds, in fields, woods, NS and ON, south to GA, gheny mound ant; principal host for clearings west to WI and NM N. gracilicornis; minor host for N. browni, N. parasitus, N. triangulum F. glacialis Wheeler; minor host for Low, grass-covered mounds, in NF south to NY, west to MB N. tenuitarsis open fields and woods and ND F. integra Nylander; principal host for In open forests and woods; in NS to GA, west to SD and N. falli, N. integer; minor host for logs, stumps, under stones, MS N. parasitus debris often on nest F. nitidiventris Emery; minor host for In soil, under stones, base of PQ and ON, south to GA, N. triangulum grass tufts; host of slave west to WY and NM makers F. obscuriventris Mayr; minor host for Woods and grasslands; under PQ and ME south to GA, N. integer logs, some thatching on nest west to MT and UT F. pergandei Emery; minor host for Social parasite of other Formica PQ and NH, south to NC, N. parasitus spp. west to UT F. schaufussi Mayr; minor host for In soil, under stones, at base of ON to GA, west to SD and N. parasitus grass tufts; host of slavers TX F. subsericea Say; principal host for Low mounds in deciduous NB and PQ south to FL, N. myrmecophilus and N. parasitus; woods; in soil and under west to ID and UT alternate host for N. tenuitarsis litter F. aserva Forel; seemingly principal Social parasite of other Formica NF to AK, south to NY, IL, host for N. browni spp. KS, AZ, CA F. ulkei Emery; seemingly host for Large mounds in forest edges or NS to MB, south to OH, IL, N. criddlei and possible minor host clearings; occasionally with CO for N. tenuitarsis plant debris layer Lasius alienus (Foerster), the cornfield Forests, in logs and stumps; un- NS to BC, south to FL, OK, ant; minor host for N. parasitus der stones AZ, CA

Phylogeny and zoogeography early Tertiary (Matthews 1979), we know of no compelling evidence to support this. Because of A phylogenetic analysis of the North Ameri- the somewhat greater number of genera in the can species of Anemadini and their relation- Palearctic (7), we think that generic differentia- ships with the Palearctic and Oriental species is tion could have been centered there, and that it is not possible because of the unavailability of more parsimonious to postulate that the Nearctic comparative specimens and the inadequate liter- fauna was assembled by several dispersal events ature illustrations or descriptions. We attempt to to the Nearctic from the Palearctic, across the suggest possible relationships through the use area known as Beringia, lying between Siberia of informal species groups. These can be tested and Alaska. Dispersal probably occurred at vari- against Palearctic species at a later date. ous times in the Tertiary. This route is supported The genera of Anemadini are mostly either by the paleobotanical record and accounts for the north temperate or south temperate in their dis- strong similarity between the forest vegetation tribution. Of the five subtribes, Eunemadina (16 and insects of the southeastern United States and genera) and Paracatopina (2 genera) have a those of warm temperate southeastern Asia south temperate distribution in Australia, New (Matthews 1979). We think that the evidence sug- Zealand, and southern South America (southern gests that at least four lineages of Palearctic Chile and Argentina), except for Dissochaetus, Anemadini dispersed across Beringia to the which has a seemingly derived distribution, be- Nearctic. Early to late Tertiary colonizations ing widespread in the Neotropics, with three spe- could have been by ancestors of (1) Neoeocatops, cies in the Nearctic. The subtribe Anemadina (4 (2) Nemadus (Laferius), (3) Nemadus pusio and genera) is Palearctic, and Nemadina (2 genera) N. horni of the colonoides species group of and Eocatopina (2 genera) are Holarctic. Nemadus (Nemadus), and (4) the parasitus spe- Anemadina and Nemadina have a few species in cies group of Nemadus (Nemadus). There is no the northern margins of the Oriental Region in compelling evidence of Pleistocene dispersals be- Taiwan, Sumatra, and northern India. None of cause of a lack of shared species in the Nearctic these last three subtribes occurs in the Neotropi- and Palearctic regions. cal, Ethiopian, and Australasian regions. In the absence of any conflicting information, Systematics we accept the hypothesis (Giachino et al. 1998) that the tribe Anemadini originated in a southern Subfamily CHOLEVINAE Kirby, Pangean setting, probably in the part that would become Gondwana. Supporting this is the fact 1837 that the greatest diversity is in the 16 genera and over 81 species of Eunemadina in south temper- Diagnosis ate regions. With the separation of Pangea into a Head with distinct occipital carina, resting northern Laurasia and a southern Gondwana, it against front of pronotum when head is in re- seems likely that the Eunemadina of Australia pose; cervical sclerites present; procoxal cavi- and southern South America and the Paracatopina ties closed behind. of New Zealand originated within these Gond- wanan subdivisions. The Laurasian lineages dif- Tribe ANEMADINI Hatch, 1928 ferentiated into the Anemadina, Nemadina, and Eocatopina. Although there may have been either Diagnosis “trans-Atlantic” vicariance or dispersal of these Mesocoxal cavities contiguous; male genital three subtribes between western Europe and east- segment well developed, tubular; elytra trans- ern North America (Giachino et al. 1998) in the versely striolate.

Key to North American subtribes and genera of Anemadini Couplet 1 adapted from Newton (1998). 1a. Metepisternum with small tongue-like process that overlaps elytron in repose; post-mesocoxal carina bor- dering mesocoxal cavity throughout, not angulate; procoxal cavity closure with broad overlap between notal and sternal processes; antenna not compact, segments 7, 9, and 10 without periapical carina; subtribe Eunemadina ...... Dissochaetus

1b. Metepisternum without tongue-like process; post-mesocoxal carina angularly separated from mesocoxal cavity except at base and apex of coxa; procoxal closure with little overlap between notal and sternal processes; antenna compact and with periapical carina on segments 7, 9, and 10 ...... 2 2a. Paramere of aedeagus with more than 3 setae (Fig. 5); male genital segment (Fig. 6) with pleurosternites with long ventromedial area of overlap; female sternite 8 lacking anterior apophysis (Fig. 7); elytral strigae appearing finer, more closely spaced, about 40 per mm; subtribe Eocatopina . . . Neoeocatops gen. nov. 2b. Paramere of aedeagus with at most 3 setae; male genital segment with pleurosternites without long ventromedial area of overlap; female sternite 8 with anterior apophysis present; elytral strigae appearing coarser, more widely spaced, about 20–30 per mm; subtribe Nemadina ...... Nemadus

Subtribe EUNEMADINA Newton, Perreau; parameres with only 4–5 apical setae 1998 (Fig. 5); male genital segment with pleurosternites overlapping ventromedially along most of their length, median ventral sternite absent Genus Dissochaetus Reitter, (Fig. 6). For comparison of the male genital seg- 1885 [1884] ment with that of Eocatops (s. str.) pelopsis Three species are known in North America Reitter, see Jeannel (1936, Fig. 354). north of Mexico, and these are reviewed in Peck (1999). Description Body (Fig. 1) elliptical, not attenuated poste- Subtribe EOCATOPINA riorly. Total length 2.5–3.8 mm; greatest width 1.3–2.0 mm. Color yellowish brown to dark Jeannel, 1936 brown; head and antennal club darker. Pubes- The subtribe is otherwise composed of 17 cence short, golden, appressed. Epistomal su- species in the Old World, mostly with a distri- ture absent. Eyes well developed. Apical bution in the more semi-arid regions of the segment of maxillary palpus (Fig. 2) conical, southern Palearctic, from Libya, through the slightly longer than penultimate segment. An- Middle East, to Afghanistan (Perreau 2000). tenna (Fig. 3) short, robust, not reaching base Their habitats are poorly known, but some re- of pronotum; segment 3 slightly shorter than cords are from caves, mammal burrows, and segment 2, segment 6 short, segment 8 disc- with ants. One species occurs in Scandinavia like. Pronotum widest behind middle; l:w and lives with Formica ants. (length to width) ratio = 1:1.5–1:1.6; front margin broadly emarginate, sides arcuate, anterior angles obtusely rounded, posterior angles Genus Neoeocatops Peck and broadly rounded. Pronotal punctation rugose, Cook, gen. nov. aligned in irregular rows. Elytra elongate, narrow, widest in basal half, slightly narrowed api- Type species cally; sutural striae impressed, striae 2–5 Choleva decipiens Horn, 1880. weakly indicated; transverse strigae perpendicu- lar to suture, narrowly spaced, 39–40 per mm. Etymology Hind wings fully formed. Mesosternum weakly Formed from neo for new, referring to the carinate. Mesocoxae contiguous. Male protibia New World, and Eocatops, which is the appar- (Fig. 4) short, expanded apically, with 2 long ent sister genus in the Old World. Gender: mas- inner spines, but lacking a true comb of small culine. spines; protarsus (Fig. 4) shorter than protibia; tarsal segments 1–3 short, broad, emarginate Diagnosis apically. Mesotibia and metatibia elongate, nar- The single species placed in Neoeocatops dif- row, with 2 long inner spines and an apical fers from the species of Eocatops (including comb of small spines; mesotibia curved, meta- subgenus Eonemadus Perreau, 1991) in the fol- tibia straight; mesotarsi simple in both sexes. lowing characters: apical segment of maxillary Aedeagus (Fig. 5) with basal piece slightly palpus longer than penultimate segment (Fig. 2); shorter than parameres; ventral lamella short, posterior pronotal angles rounded, not angulate; about one-half length of basal piece; median protibia lacks a true comb of small spines api- lobe broad, flat, nearly as long as parameres, cally (Fig. 4), as also in E. (Eonemadus) loebli narrowing to ogival apex. Internal sac when

Figs. 1–7. Neoeocatops decipiens: 1, outline of body; 2, maxillary palpus; 3, antenna; 4, male protibia and protarsus; 5, aedeagus, dorsal; 6, male genital segment, ventral; 7, female sternite 8. Scale bar = 1.0 mm (Fig. 1), 0.2 mm (Fig. 2), and 0.4 mm (Figs. 3–7).

1 5

7 everted presents 2 elongate rows of broad, flat Territory), red label “Lectotype 2993”, white teeth, with finer setae apically; when inverted, handwritten label “C. decipiens Horn”, and the large teeth overlap in 2 elongate rows. Para- our red lectotype label; seen and dissected. meres elongate, narrow, with 4–5 short apical Horn (1880: 260) stated that the specimens setae. Male genital segment (Fig. 6) longer than were collected by Morrison from Olympia, wide; pleurosternites ventromedially overlap- Washington, which should be considered the ping at length, with numerous small setae api- type locality. Paralectotypes: 5 in Horn col- cally; median ventral sclerite absent; anterior lection and 1 in LeConte collection, MCZC, apophysis broad at base, narrowing gradually to with same white “W.T.” label. apex. Spermatheca not sclerotized. Female Catops decipiens (Horn), Hatch 1928: 201. sternite 8 (Fig. 7) short and broad; anterior apo- Dissochaetus decipiens (Horn), Hatch 1933: physis lacking. 198; Jeannel 1936: 151, 154. Nemadus decipiens (Horn), Hatch 1957: 43. Neoeocatops decipiens (Horn), comb. nov. Material examined (Figs. 1–7, 100) Four hundred sixty-seven specimens. Choleva decipiens Horn, 1880: 259. Lectotype here designated as male in Horn collection, Diagnosis and redescription MCZC, with white label “W.T.” (=Washington Characters as for the genus (above).

Distribution Seasonality The species is distributed in far western Adults have been taken in all months except North America (Fig. 100) from the mountains April. The largest numbers are from July and of coastal and interior British Columbia, south August, followed by February and May in along the Coast and Cascade mountains to the southern California. Sierra Nevada and coastal mountains, to ex- treme southern California. It is known from the Subtribe NEMADINA Jeannel, following provinces and states: British Colum- 1936 bia, Canada; California, Oregon, and Washing- ton, United States. Genus Nemadus Thomson, 1867 Diagnosis Habitats Posterior coxae contiguous, all tarsi with five The species is a scavenger in moist forests to tarsomeres; elytra with glossy surface, setal semi-arid thickets of manzanita (Arctostaphylos bases arranged in transverse strigae; hind mar- spp.; Ericaceae) and chaparral vegetation. It has gin of hind tibia with 2 long inner spines and been collected mostly in unbaited pitfall traps with about 4 longer outer spines; male (with ex- or pitfalls baited with carrion and fermenting panded protarsi) basal mesotarsomere dilated malt solution, and was next most commonly and spongy pubescent beneath; mesosternal ca- taken in flight-intercept traps. There is one re- rina elevated and extending nearly to anterior cord from a blacklight trap. margin of mesosternum.

Key to North American subgenera of Nemadus Nemadus is keyed within the context of the North American genera of Leiodidae in Peck (2000). See Perreau (2000) for included species and references. 1a. Body shape triangular; first segment of male mesotarsus weakly dilated; basal segments of male protarsi weakly dilated ...... Nemadus (Laferius) 1b. Body shape elongate-oval; first segment of male mesotarsus strongly dilated; basal segments of male protarsi strongly dilated ...... Nemadus (Nemadus)

Key to Nemadus species of North America 1a. Body shape triangular (Fig. 8); pronotum width more than twice length, posterior angles slightly produced; elytra strongly attenuated posteriorly; antennae short, not reaching posterior margin of pronotum; male protarsus (Fig. 10) narrower than tibial apex; first mesotarsal segment of male weakly dilated ...... N.(Laferius) brachyderus (LeConte) 1b. Body shape elongate-oval to broadly oval; width of pronotum twice length or less, posterior angles not produced; elytra weakly to moderately attenuated posteriorly; antennae usually reaching posterior margin of pronotum; male protarsi usually as wide as or wider than tibial apex; first mesotarsal segment of male dilated; Nemadus s.str...... 2 2a. Small species, total length usually less than 2.0 mm; antennal segment 8 short (Figs. 16, 23), length less than one-half width; elytra of both sexes with lines of microsculpture between and parallel to transverse strigae; non-myrmecophilous species; the colonoides species group ...... 3 2b. Larger species, total length usually 2.0 mm or more; length of antennal segment 8 about one-half width; males never with elytral microsculpture between transverse strigae; myrmecophilous species; the parasitus species group ...... 4 3a. Body shape elliptical (Fig. 22); male first protarsal segment about as wide as tibial apex (Fig. 24); west coast of North America ...... N.(N.) pusio (LeConte) 3b. Body shape broader, weakly attenuated posteriorly (Fig. 15); male first protarsal segment wider than tibial apex (Fig. 17); eastern North America ...... N.(N.) horni Hatch 4a. Body shape broad anteriorly with elytra attenuated posteriorly (Figs. 37, 53, 92)...... 5 4b. Body shape elongate-oval, weakly attenuated posteriorly (Figs. 29, 45, 61, 69, 76, 84) ...... 7 5a. Body unicolorous; parameres truncate apically (Fig. 95) ...... N. (N.) triangulum Jeannel 5b. Pronotum with dark median area, the fuscous cloud of Fall (1937); parameres narrow apically in dorsal view (Figs. 40, 56) ...... 6

6a. Antennal segment 7 strongly narrowed in basal half (Fig. 54); northeastern United States ...... N. (N.) gracilicornis Fall 6b. Antennal segment 7 weakly narrowed in basal half (Fig. 38); Manitoba and Alberta ...... N. (N.) criddlei sp. nov. 7a. Median lobe of aedeagus triangular in cross section, with median dorsal longitudinal carina (Figs. 48, 64) ...... 8 7b. Median lobe of aedeagus dorsoventrally flat, not carinate dorsally...... 9 8a. Female without microsculpture between elytral strigae ...... N. (N.) integer Fall 8b. Female with microsculpture between elytral strigae ...... N. (N.) falli sp. nov. 9a. Female without microsculpture between elytral strigae; male first protarsal segment narrower than tibial apex (Fig. 86) ...... N. (N.) tenuitarsis Jeannel 9b. Female with microsculpture between elytral strigae; male first protarsal segment as wide as or wider than tibial apex ...... 10 10a. Parameres short, not longer than median lobe of aedeagus (Fig. 32) ...... N. (N.) browni sp. nov. 10b. Parameres longer than median lobe of aedeagus (Figs. 72, 79) ...... 11 11a. Male first protarsal segment wider than tibial apex (Fig. 78); widespread eastern species ...... N. (N.) parasitus (LeConte) 11b. Male first protarsal segment not wider than tibial apex (Fig. 71); Iowa and Kansas only ...... N. (N.) myrmecophilus Jeannel

Subgenus Laferius Perkovsky, Nemadus brachyderus (LeConte), Jeannel 1936: 1994 171.

Nemadus (Laferius) kurbatovi Perkovsky Material examined (Perkovsky 1994) is the type species for the One hundred thirty-three specimens. subgenus and is known only from the type series, from the Zelski Nature Reserve in the far Diagnosis east of the Russian Federation. There is no indi- Nemadus brachyderus can be distinguished cation in the description that it is associated from all other North American species of with ants. It is the only species assigned to this Nemadus by the following combination of char- subgenus other than the following species, acters: body shape broadly triangular (Fig. 8); which Jeannel (1936) had placed in its own antennae short, not reaching posterior margin of species group within Nemadus s. str. pronotum; male first protarsal segment narrower than tibial apex (Fig. 10). Nemadus (Laferius) brachyderus Redescription (LeConte) Body (Fig. 8) broadly oval, strongly attenu- (Figs. 8–14, 101) ated posteriorly. Total length 2.6–2.8 mm; greatest width 1.7–1.8 mm. Color reddish Catops brachyderus LeConte, 1863: 25. Lecto- brown to dark brown; some individuals with type here designated as male in LeConte col- area of darker pigmentation on center of pro- lection, MCZC, with white labels “Mass.”, notum. Pubescence short, silky, golden, ap- “Camb.”, “C. brachyderus Agassiz Lec.”, a pressed. Antenna (Fig. 9) short, not reaching red type label with number 6115, and our red base of pronotum; segment 3 slightly shorter lectotype label; seen and dissected. LeConte than segment 2; segment 7 not constricted in published the only locality known to him as basal half; length of segment 8 greater than Nova Scotia, but there is no specimen with one-half width. Pronotum transverse, widest at this locality in his material. base; l:w ratio = 1:2.1–1:2.25; hind angles Ptomaphagus brachyderus (LeConte), Horn acute, prolonged posteriorly. Elytra widest at 1880: 264. base, strongly attenuated apically; sutural striae Catopomorphus brachyderus (LeConte), Was- impressed, striae 2–9 weakly indicated; trans- mann, 1894: 127. verse strigae moderately widely spaced, 30 Adelops brachyderus (LeConte), Jeannel 1922: per mm; microsculpture of several fine trans- 43; Hatch 1928: 169. verse lines between and parallel to strigae pres- Dissochaetus brachydereus (LeConte), Hatch ent in both sexes. Hind wings fully formed. 1933: 196. Male protarsus (Fig. 10) shorter than protibia;

Figs. 8–14. Nemadus (Laferius) brachyderus: 8, outline of body; 9, antenna; 10, male protibia and protarsus; 11, aedeagus, dorsal; 12, male genital segment, ventral; 13, spermatheca; 14, female sternite 8. Scale bar = 1.0 mm (Fig. 8), 0.2 mm (Fig. 13), and 0.4 mm (Figs. 9–12, 14).

12 14 13

tarsal segment 1 narrower than tibial apex. Habitats Aedeagus (Fig. 11) with basal piece subequal in Ecology: myrmecophile. Most host associa- length to parameres; median lobe flat, about tions (fewer than half the records) are with three-fourths length of parameres, evenly nar- Camponotus noveboracensis (given as C. pictus rowed to acute apex; parameres elongate, Forel by Fall 1937), which ranges from coast to dorsoventrally flattened, with rounded apices. coast, mostly between latitudes 40°N and 48°N. Male genital segment (Fig. 12) longer than There are eight specimens taken with C. herculeanus, which ranges across southern Canada wide; anterior apophysis elongate. Spermatheca and southwards to New Mexico and Pennsylva- (Fig. 13) strongly curved; invagination bearing nia in the United States. The beetles were ap- sclerotized teeth. Female sternite 8 (Fig. 14) tri- parently collected by opening and searching the angular with elongate anterior apophysis. nests of the host ants in pine logs and stumps. Sanders (1964) reported the beetles to be com- Distribution mon in overwintering nests of C. herculeanus in New Brunswick, but we cannot confirm the re- The species is distributed from southeastern cord. The beetles apparently disperse to ant Canada and the New England states, westward nests by flight, but in what season is not appar- through the Great Lakes Region, to the Black ent. One specimen is from a nest of the rodent Hills of South Dakota and southern Manitoba Microtus pennsylvanicus (Ord), one was taken (Fig. 101). It is known from the following prov- by sweeping, and one was taken in a malt- inces and states: Manitoba, Nova Scotia, On- syrup-baited pitfall trap. Most labels do not in- tario, and Quebec, Canada; Massachusetts, dicate microhabitat or collecting method. Maine, Michigan, Minnesota, New Hampshire, New York, South Dakota, Vermont, and Wis- Seasonality consin, United States. Adults of the species have been taken mostly

in late spring – early summer and late fall – early Diagnosis winter. The numbers taken per month are as Nemadus horni can be distinguished from all follows: April, 1; May, 47; June, 14; July, 3; other North American species of Nemadus by August, 8; September, 5; October, 16; Novem- the characters of the colonoides species group ber, 18; and December, 12. and the characters given in the key.

Subgenus Nemadus Thomson, Redescription 1867 Body (Fig. 15) elongate-oval, weakly attenuated posteriorly. Total length 1.8–2.1 mm; Jeannel (l936) placed the species now in the greatest width 1.0–1.2 mm. Color medium subgenus Nemadus into two species groups, the brown to nearly black; pronotum with area of colonoides and parasitus groups, and gave a darker pigmentation in paler individuals; key of characters to diagnose them. Since then, antennal club darker than basal segments. Pu- many additional Palearctic species have been bescence of moderate length, golden, appressed described without associating them with a spe- to subappressed. Antenna (Fig. 16) of moderate cies group. The catalogue of Perreau (2000) did length, reaching base of pronotum; segment 3 not use species groups for Nemadus. The re- shorter than segment 2; segment 7 not con- sults of our study and examination of the North stricted in basal half; segment 8 short, length American fauna support Jeannel’s species- less than one-third width. Pronotum widest near group designations, but we cannot comment on base; l:w ratio = 1:1.7; hind angles obtuse, not the placement of the more recently described prolonged posteriorly. Elytra widest in anterior Palearctic species. half, weakly narrowed apically; sutural striae impressed; transverse strigae moderately widely colonoides species group spaced, 30 per mm; microsculpture of several fine lines between and parallel to strigae pres- The colonoides species group is defined by ent in both sexes. Hind wings fully formed. the following characters: small body size, total Male protarsus (Fig. 17) subequal in length to length usually less than 2.0 mm; antennal seg- protibia; tarsal segment 1 wider than tibial ment 8 short, length less than one-half width; apex. Aedeagus (Fig. 18) with basal piece lon- elytra of both sexes with lines of micro- ger than parameres; median lobe flat, about sculpture between and parallel to transverse two-thirds length of parameres, abruptly nar- strigae; parameres narrowly conchoidal (shell- rowing to small apical projection; parameres shaped). It has not been determined whether the narrow, conchoidal, with rounded apices. Male two North American species are sister species, genital segment (Fig. 19) slightly longer than or if they are more closely related to Palearctic wide; anterior apophysis moderately long. species. Spermatheca (Fig. 20) moderately curved; invagination with crescent-shaped rugosities. Nemadus (Nemadus) horni Hatch Female sternite 8 (Fig. 21) short and broad with short anterior apophysis. (Figs. 15–21, 102) Nemadus horni Hatch, 1933: 194. Type in Distribution USNM, seen. Type locality: Framingham, The species is widely distributed in eastern Massachusetts. Jeannel 1936: 169; Fall 1937: North America from southeastern Canada 337; Salgado 1999: 37; Peck and Skelley southwards to Florida and westward to Mis- 2001: 141. souri and eastern Texas (Fig. 102). It is known Ptomaphagus pusio LeConte, Horn 1880: 264, from the following provinces and states: On- misidentification. tario and Quebec, Canada; Alabama, Arkansas, Nemadus ellipticus Jeannel 1936: 170. Type District of Columbia, Florida, Georgia, Iowa, Il- male in MNHN, seen. Type locality: Cin- linois, Indiana, Kentucky, Louisiana, Massachu- cinnati, Ohio. New synonomy. setts, Maryland, Maine, Michigan, Missouri, North Carolina, New Hampshire, New Jersey, Material examined New York, Ohio, Pennsylvania, South Carolina, One thousand one hundred fifty-eight speci- Tennessee, Texas, Virginia, Vermont, and Wis- mens. consin, United States.

Figs. 15–21. Nemadus (Nemadus) horni: 15, outline of body; 16, antenna; 17, male protibia and protarsus; 18, aedeagus, dorsal; 19, male genital segment, ventral; 20, spermatheca; 21, female sternite 8. Scale bar = 1.0 mm (Fig. 15), 0.2 mm (Fig. 20), and 0.4 mm (Figs. 16–19, 21).

18 17

19 20 15 21

For the use of workers who inventory the scavenger commonly taken in the eastern fauna of caves, we provide the following re- deciduous forest biome on small dead mammals, cords of caves known to have contained popula- in dung- and carrion-baited pitfall traps, in tions of Nemadus horni: flight-intercept traps, in forest litter, and in moist CANADA. Ontario: Leeds Co.: Stonehouse guano of insectivorous bats. It has been collected Cave. UNITED STATES. Alabama: Blount less commonly by car nets, in rotted soft fungi, Co.: Bangor Cave. Calhoun Co.: Lady Cave, in tree holes, and in the nests of rabbits and the Weaver Cave. Colbert Co.: Gist Cave, Wolf vole M. pennsylvanicus, the dens of woodchucks Den Cave. Conecuh Co.: Turks Cave. DeKalb and foxes, and the burrows of Geomys pocket Co.: Bartlett Cave, Cook Cave, Fort Payne gophers in Alabama and Georgia (Peck and Cave, Lykes Cave. Jackson Co.: House of Hap- Skelley 2001). It is not myrmecophilous. Cave piness Cave, Saltpeter Cave. Limestone Co.: and higher altitude (montane) populations are Pope Cave. Marshall Co.: Terrill Cave. Morgan more prevalent in the southern part of the spe- Co.: Talucah Cave. Talladaga Co.: Kymulga cies range. Cave, Dulaney Cave. Florida: Alachua Co.: Warren’s Cave. Jackson Co.: Judges Cave, Seasonality Gerads Cave. Marion Co.: Metfords Cave. Adults have been collected in southern Can- Georgia: Dade Co.: Morrison Cave, Newsome ada from April to December, but they are active Gap Cave. Decatur Co.: Climax Cave. Walker mostly from May to September. The species Co.: Cove Cave, Cave Spring Cave, Pettijohn was taken only in May and August at carrion Cave. Illinois: Monroe Co.: Fogelpole Cave. baits in low numbers in a deciduous forest in Indiana: Clark Co.: Cave Springs Cave, Oak southern Ontario (Peck and Anderson 1985). It Hill Cave. Crawford Co.: Little Wyandotte also occurred in low numbers during summer in Cave. Orange Co.: Cave’s Cave, Tucker Dam flight-intercept traps in forests in New Hamp- Quarry Cave. Kentucky: Trigg Co.: Cool shire (Chandler and Peck 1992). In forests and Spring Cave. Woodford Co.: Weber’s Cave. caves in the southeastern United States, adults Tennessee: Bedford Co.: Reese Cave. Bradley are active in all months of the year, but in for- Co.: Quarry Cave. Butler Co.: Reese Cave. Cof- ests they are more active in the cooler months fee Co.: Jarrells Cave. Dickson Co.: Columbia than in the hotter, drier summer months. Caverns. Franklin Co.: Caney Hollow Cave, Dry Cave Sink. Robertson Co.: Double Cave. Texas: Wheeler Co.: Big Mouth Cave. Vir- Nemadus (Nemadus) pusio ginia: Lee Co.: Sweet Potato Cave, Gilley (LeConte) Cave. (Figs. 22–28, 103) Habitats Catops pusio LeConte, 1859: 282. Lectotype The species is the most frequently collected here designated as female in LeConte collec- member of the genus. It is seemingly a generalist tion, MCZC, with gold disc and red type

Figs. 22–28. Nemadus (Nemadus) pusio: 22, outline of body; 23, antenna; 24, male protibia and protarsus; 25, aedeagus, dorsal; 26, male genital segment, ventral; 27, spermatheca; 28, female sternite 8. Scale bar = 1.0 mm (Fig. 22), 0.2 mm (Fig. 27), and 0.4 mm (Figs. 23–26, 28).

23 25 24

26 22 27 28

label with number “3155” and our red parallel to strigae present in both sexes. Hind lectotype label. Seen. Published type locality: wings fully formed. Male protarsus (Fig. 24) Punto de los Reyes (Point Reyes), California. subequal in length to protibia; tarsal segment 1 Ptomaphagus pusio, Horn 1880: 264; misidenti- subequal in width to tibial apex. Aedeagus fication of material of Nemadus horni from (Fig. 25) with basal piece longer than Michigan and Lake Superior region. parameres; median lobe flat, nearly as long as Blatchley 1910: 282; misidentification of ma- parameres, evenly narrowed to acute apex; terial from Indiana. parameres narrow, weakly conchoidal, with Nemadus pusio (LeConte), Hatch 1933: 195, rounded apices. Male genital segment (Fig. 26) 1957: 43; Jeannel 1936: 169; Fall 1937: 337; longer than wide; anterior apophysis elongate, Salgado 1999: 37. narrow. Spermatheca (Fig. 27) weakly curved. Female sternite 8 (Fig. 28) short and broad with Material examined short anterior apophysis. One hundred thirty-one specimens. Distribution Diagnosis The species is distributed in far western Nemadus pusio can be distinguished from all North America from northern Queen Charlotte other North American Nemadus species by the Islands of British Columbia, southwards along characters of the colonoides species group and the Coast and Cascade ranges to coastal Cali- the characters given in the key. fornia at Santa Barbara, and inland in the Sierra Nevada to Sequoia National Park (Fig. 103). It Redescription is known from the following provinces and Body (Fig. 22) elliptical, not attenuated pos- states: British Columbia, Canada; California, teriorly. Total length 1.5–1.8 mm; greatest Oregon, and Washington, United States. width 0.8–0.9 mm. Color yellowish brown to dark brown; pronotum and scutellum often darker; antennal club darker than basal seg- Habitats ments. Pubescence of moderate length, golden, The species is seemingly a generalist scaven- appressed. Antenna (Fig. 23) of moderate ger in moist wooded habitats. No habitat or length, barely reaching base of pronotum; seg- food preference is indicated by label data. ment 3 distinctly shorter than segment 2; seg- Specimens have been collected in low numbers ment 7 not constricted in basal half; segment 8 in a chicken nest, in nests of Neotoma spp. short, length about one-third width. Pronotum (woodrats) (Walters and Roth 1950), in pocket widest before base; l:w ratio = 1:1.7; hind an- gopher burrows, in ground nests of the wasp gles obtuse, not prolonged posteriorly. Elytra Vespula pensylvanica Rohwer (Hymenoptera: widest anterior to middle, rounded apically; Vespidae), in hardwood and conifer leaf litter, sutural striae impressed; transverse strigae mod- at blacklights, in carrion- and human-dung- erately widely spaced, 30 per mm; micro- baited pitfall traps, in flight-intercept traps, and sculpture of several fine lines between and by sweeping of grasses.

Seasonality Canada, Ottawa, from 1927 to 1967 (Cody et Adults have been taken in all months except al. 1986). September. The largest numbers of records are from January (28), May (27), and June (27). Diagnosis Nemadus browni can be distinguished from parasitus species group all other North American species of Nemadus by the characters of the parasitus species group The parasitus species group is defined by the and the characters given in the key. following characters: body size larger than in the colonoides group, total length usually Description greater than 2.0 mm; length of antennal seg- Body (Fig. 29) oval, weakly attenuated poste- ment 8 greater than one-half width; males al- riorly. Total length 1.8–2.2 mm; greatest width ways lacking elytral microsculpture; parameres 1.1–1.4 mm. Color reddish brown, pronotum broadly conchoidal (shell-like) in shape; myr- with area of darker pigmentation. Pubescence mecophilous species. Summaries of host ant of moderate length, golden, subappressed. An- nest characteristics and distributions are in Ta- tenna (Fig. 30) short, not reaching base of pro- ble 1. The group is presently recognized to oc- notum; segment 3 distinctly shorter than cur only in North America. These species can segment 2; segment 7 not constricted in basal be difficult to identify if both male and female half; length of segment 8 less than one-half specimens and host ants are not available for width. Pronotum widest before base; l:w ratio = study. 1:1.9–1:2.0; hind angles subquadrate, not prolonged posteriorly. Elytra widest in basal half, Nemadus (Nemadus) browni Peck weakly attenuated apically; sutural striae im- and Cook, sp. nov. pressed; transverse strigae moderately widely spaced, 27 per mm; microsculpture of several (Figs. 29–36, 104) fine lines between and parallel to strigae present in female only; male elytra shining. Hind Type material wings fully formed. Male protarsus (Fig. 31) Holotype: male (CNCI). CANADA. Mani- subequal in length to protibia; tarsal segment 1 toba: Riding Mt. Pk., 2.VI.38, W.J. Brown. wider than tibial apex. Aedeagus (Figs. 32, 33) Paratypes (48). CANADA. Ontario: Ottawa, 5 with basal piece longer than parameres; median (CNCI). Quebec: Gatineau Park, Ramsay lobe flat, as long as parameres, abruptly nar- L[ake] area, 13.VI.1971, J.M. Cambell, 2 rowed to small apical projection; parameres (CNCI). Cascapedia, 21.VI.1933, W.J. Brown, short, broad, conchoidal, with rounded apices. 1 (CNCI). Manitoba: Riding Mt. Pk., 4.VI.38, Male genital segment (Fig. 34) about as long as W.J. Brown, 3 (CNCI). Riding Mt. Pk., 5.VI.38, wide; anterior apophysis moderately long. W.J. Brown, 25 (CNCI, SBPC). UNITED Spermatheca (Fig. 35) moderately curved; STATES. Maine: Cumberland Co., V.9.1916, invagination bearing strong setae. Female A. Nicolay, 1 (USNM). Same data except sternite 8 (Fig. 36) triangular with short anterior VI.14.1916, 1 (USNM). Minnesota: Marble, apophysis. 3.VI.1935, K. Cooper, 2 (USNM). New Hamp- shire: no other data, 1 (MCZC). Pittsburg, Distribution 30.V.1937, C.A. Frost, sifting at sap, 1 (CNCI). This species is distributed in the southern Same data, 2 (MCZC). Antrim, 23.VI.1933, part of eastern and central Canada and the adja- C.A. Frost, under lawn grass, 1 (MCZC). South cent northern tier of states of the eastern United Dakota: Hill City, Coll. Hubbard & Schwarz, 3 States (Fig. 104). It is known from the follow- (USNM). ing provinces and states: Quebec, Ontario, and Manitoba, Canada; Maine, New Hampshire, Etymology Minnesota, and South Dakota, United States. The epithet browni, a patronym, is in recognition of the extensive field collecting across Habitats Canada and publications on beetles of W.J. Ecology: myrmecophile. Adults have a prefer- Brown (1902–1977). He worked as Canada’s ence for the nests of Formica aserva (= Formica first professional coleopterist with the Canadian subnuda) (32 specimens from six collecting National Collection of Insects, Agriculture events in Manitoba and Quebec), and a single

Figs. 29–36. Nemadus (Nemadus) browni: 29, outline of body; 30, antenna; 31, male protibia and protarsus; 32, aedeagus, dorsal; 33, aedeagus, lateral; 34, male genital segment, ventral; 35, spermatheca; 36, female sternite 8. Scale bar = 1.0 mm (Fig. 29), 0.2 mm (Fig. 35), and 0.4 mm (Figs. 30–34, 36).

31 32 34

33 29

35 36

specimen was associated with F. exsectoides in was employed as a Canadian government ento- Maine. A few specimens were collected by sift- mologist from 1902 onwards (Criddle 1973; ing at sap flows at tree bases. Roughley 2000; Wheeler 2005).

Seasonality Diagnosis Adults have been collected only in the Nemadus criddlei can be distinguished from months of May and June. all other North American species of Nemadus by the characters of the parasitus species group Nemadus (Nemadus) criddlei Peck and the characters given in the key. and Cook, sp. nov. Description (Figs. 37–44, 104) Body (Fig. 37) oval, moderately attenuated posteriorly. Total length 2.0–2.2 mm; greatest Type material width 1.3 mm. Color light reddish brown, pro- Holotype: male (CMNC). CANADA. Al- notum with area of darker pigmentation. Pubes- berta: Tp. [township] 29, Rge. [range] 5, W. 5 cence of moderate length, golden, subappressed. Mer.[idian], 14.X.1961, B.F. & J.L. Carr, ant Antenna (Fig. 38) of moderate length, reaching nest sifting. Paratypes (14). CANADA. Al- base of pronotum; segment 3 subequal to seg- berta: Tp. 20, Rge. 3, W. 5 Mer., 19.IV.1964, ment 2 in length; segment 7 not constricted in B.F. & J.L. Carr, ant nest tramping, 3 (SBPC). basal half; length of segment 8 greater than one- Manitoba: Aweme, 1.V.1916, N. Criddle, 2 half width. Pronotum widest before base; l:w ra- (CNCI). Aweme, 2.V.1916, N. Criddle, 2 tio = 1:1.9–1:2.0; hind angles subquadrate, not (MCZ), 6 (CNCI, SBPC). Aweme, 3.V.1916, N. prolonged posteriorly. Elytra widest at base, Criddle, 1 (CNCI). moderately attenuated apically; sutural striae impressed; transverse strigae widely spaced, 22 Etymology per mm; microsculpture of several fine lines be- The epithet criddlei, a patronym, is in recogni- tween and parallel to strigae present in female tion of the meticulous pioneering beetle collecting only; male elytra shining. Hind wings fully in the early 1900s of Norman Criddle (1875– formed. Male protarsus (Fig. 39) longer than 1933) at Aweme, near Treesbank, Manitoba. He protibia; tarsal segment 1 wider than tibial apex.

Figs. 37–44. Nemadus (Nemadus) criddlei: 37, outline of body; 38, antenna; 39, male protibia and protarsus; 40, aedeagus, dorsal; 41, aedeagus, lateral; 42, male genital segment, ventral; 43, spermatheca; 44, female sternite 8. Scale bar = 1.0 mm (Fig. 37), 0.2 mm (Fig. 43), and 0.4 mm (Figs. 38–42, 44).

40 39

37 41

44 43

Aedeagus (Figs. 40, 41) with elongate basal Seasonality piece; median lobe flat, about two-thirds length Adults have been collected in April, May, of parameres, evenly narrowed to small apical and October. projection; parameres broad, conchoidal, with rounded apices. Male genital segment (Fig. 42) Nemadus (Nemadus) falli Peck and longer than wide; anterior apophysis moderately Cook, sp. nov. long. Spermatheca (Fig. 43) strongly curved; invagination bearing sclerotized teeth. Female (Figs. 45–52, 104) sternite 8 (Fig. 44) triangular with short anterior apophysis. Type material Holotype: male (CMNC). UNITED STATES. Ohio: Hocking Co., 3 mi. W 33 on Distribution 16, 3.V.1975, L.E. Watrous, Q.D. Wheeler, in The species is known only from the southern nest of Formica. Paratypes (85). UNITED regions of the Canadian provinces of Alberta STATES. District of Columbia: 2 (MCZC), 1 and Manitoba (Fig. 104). (CNCI), collection of Frederick Allen Eddy, 2 (MCZC), W.D. Dietz Coll., 3 (MCZC), Henry Ulke Beetle Coll., 7 (CMNH). Brookland, Habitats 17.VIII.1896, in nest of Formica integra, Coll. Ecology: myrmecophile. Known only from T. Pergande, 1 (USNM). Wash.[ington], Ulke, 1 ant nests. A single specimen was taken in the (CNCI). Massachusetts: no precise locality, nest of Formica ulkei, the only record of a spe- collection of Frederick Allen Eddy, 1 (MCZC). cific host association. We think it notable that Maryland: Plummers I., 19-4-08, W.L. Park (1929, 1935), in one of the more intensive McAtee, 4 (USNM), 1 (CNCI). Same data, 17- studies of inhabitants of the nests of F. ulkei, 5-08, 1 (USNM). New Jersey: Hillsdale, seemingly did not find Nemadus spp. with these 12.VII.1936, Quirsfeld, 1 (MCZC). Ohio: ants in northeastern Illinois. Cinci.[nnati], V-10-16, 2 (MCZC). Same data

Figs. 45–52. Nemadus (Nemadus) falli: 45, outline of body; 46, antenna; 47, male protibia and protarsus; 48, aedeagus, dorsal; 49, aedeagus, lateral; 50, male genital segment, ventral; 51, spermatheca; 52, female sternite 8. Scale bar = 1.0 mm (Fig. 45), 0.2 mm (Fig. 51), and 0.4 mm (Figs. 46–50, 52).

48 47

49 45

52 51 50

as holotype, 10 (OSUC), 21 (SBPC). Pennsyl- Description vania: Pittsburg, VIII, 1 (CMNH). Virginia: Body (Fig. 45) oval, weakly attenuated poste- Vienna, 11.XII, J.C. Bridwell, ex log, 16 riorly. Total length 2.2–2.4 mm; greatest width (USNM). Franklin Co., Blue Ridge Pkwy. mi 1.3–1.4 mm. Color reddish brown; pronotum 154, Smart View, 2560′, 31.V–20.VIII.1981, S. with area of darker pigmentation. Pubescence Peck, intercept traps, 4 (SBPC). of moderate length, golden, suberect. Antenna (Fig. 46) reaching base of pronotum; segments 2 and 3 subequal in length; segment 7 not con- Etymology stricted in basal half; length of segment 8 about The epithet falli, a patronym, is in recogni- one-half width. Pronotum widest before base; tion of the contributions of Henry C. Fall l:w ratio = 1:1.8–1:1.9; hind angles weakly ob- (1862–1939) to leiodid and other beetle taxon- tuse, not prolonged posteriorly. Elytra widest at omy. He was a teacher in southern California base, weakly attenuated apically; sutural striae who retired to Tyngsboro, Massachusetts, and impressed; transverse strigae moderately widely placed his extensive beetle collection of over spaced, 27 per mm; microsculpture of several 100 000 specimens in the Museum of Compara- fine lines between and parallel to strigae pres- tive Zoology, Harvard University. He described ent in female only; male elytra shining. Hind 1484 beetle species in 144 entomological pa- wings fully formed. Male protarsus (Fig. 47) pers, and his last paper, published in 1937, was subequal in length to protibia; tarsal segment 1 on Nemadus. His life and publications are sum- wider than tibial apex. Aedeagus (Figs. 48, 49) marized in Darlington (1940). with basal piece longer than parameres; median lobe about three-fourths length of parameres, triangular in cross section, with median dorsal Diagnosis carina, apex elongate, narrow. Parameres broad, Nemadus falli can be distinguished from all conchoidal, with rounded apices. Male genital other North American species of Nemadus by segment (Fig. 50) about as long as wide; ante- the characters of the parasitus species group rior apophysis moderately long. Spermatheca and the characters given in the key. (Fig. 51) strongly curved; invagination bearing

setae. Female sternite 8 (Fig. 52) triangular between and parallel to strigae present in fe- with short anterior apophysis. male only; male elytra shining. Hind wings fully formed. Male protarsus (Fig. 55) longer Distribution than protibia; tarsal segment 1 wider than tibial This species is distributed in the eastern apex. Aedeagus (Figs. 56, 57) with basal piece United States from southern New England and longer than parameres; median lobe flat, about the mid-Atlantic states to Ohio (Fig. 104). It is two-thirds length of parameres, evenly nar- known from the following states: District of rowed to small, elongate projection; parameres Columbia, Massachusetts, Maryland, New Jer- broad, conchoidal, with emargination on inner sey, Ohio, Pennsylvania, and Virginia. margin before rounded apex. Male genital segment (Fig. 58) longer than wide; anterior apo- Habitats physis elongate. Spermatheca (Fig. 59) weakly Ecology: myrmecophile. Adults are associ- curved, invagination bearing strong setae. Fe- ated with Formica integra. male sternite 8 (Fig. 60) triangular with short anterior apophysis. Seasonality Adults have been collected in the months of Distribution March, April, May, July, August, and Decem- The species occurs in the northeastern United ber. States (Fig. 105). It is known from the following states: Connecticut, Massachusetts, New Nemadus (Nemadus) gracilicornis Hampshire, New Jersey, and New York. Most Fall of the specimens were collected by H.C. Fall in Massachusetts. (Figs. 53–60, 105) Nemadus gracilicornis Fall, 1937: 339. Type Habitats male in MCZC, seen and dissected. Type lo- Ecology: myrmecophile. There are 58 speci- cality: Cornwall, Connecticut. mens associated with Formica exsectoides as the host. There are no other collecting data other Material examined than these associated with ants. One hundred forty-one specimens. Seasonality Diagnosis Adults have been mostly collected with ants Nemadus gracilicornis can be distinguished in the spring in April and May: March, 2; April, from all other North American species of 66; May, 40; June, 5; September, 2; and Octo- Nemadus by the characters of the parasitus spe- ber, 1. cies group and the characters given in the key.

Redescription Body (Fig. 53) oval, moderately attenuated Nemadus (Nemadus) integer Fall posteriorly. Total length 2.1–2.4 mm; greatest (Figs. 61–68, 106) width 1.3–1.4 mm. Color reddish brown, pronotum with area of darker pigmentation. Pubes- Nemadus integer Fall, 1937: 338. Type female cence of moderate length, golden, appressed to in MCZC, seen and dissected. Type locality: subappressed. Antenna (Fig. 54) elongate, nar- Tyngsboro, Massachusetts. row, usually extending slightly beyond base of pronotum; segment 3 subequal to segment 2 in Material examined length; segment 7 constricted in basal half; length of segment 8 greater than one-half width. Two hundred nine specimens. Pronotum widest before base; l:w ratio = 1:1.9– 1:2.1; hind angles subquadrate, not prolonged Diagnosis posteriorly. Elytra widest at base, moderately Nemadus integer can be distinguished from attenuated apically; sutural striae impressed; all other North American species of Nemadus transverse strigae widely spaced, 21–22 by the characters of the parasitus species group per mm; microsculpture of several fine lines and the characters given in the key.

Figs. 53–60. Nemadus (Nemadus) gracilicornis: 53, outline of body; 54, antenna; 55, male protibia and protarsus; 56, aedeagus, dorsal; 57, aedeagus, lateral; 58, male genital segment, ventral; 59, spermatheca; 60, female sternite 8. Scale bar = 1.0 mm (Fig. 53), 0.2 mm (Fig. 59), and 0.4 mm (Figs. 54–58, 60).

56 55

53 57

60 59

Redescription elongate, narrow. Parameres broad, conchoidal, Body (Fig. 61) oval, weakly attenuated poste- with rounded apices. Male genital segment riorly. Total length 2.2–2.5 mm; greatest width (Fig. 66) slightly longer than wide; anterior 1.3–1.4 mm. Color reddish brown to black; pro- apophysis moderately long. Spermatheca notum with area of darker pigmentation in paler (Fig. 67) strongly curved; invagination bearing individuals. Pubescence of moderate length, sclerotized teeth. Female sternite 8 (Fig. 68) tri- golden, suberect. Antenna (Fig. 62) of moderate angular with short anterior apophysis. length, barely reaching base of pronotum; segment 3 shorter than segment 2; segment 7 not Distribution constricted in basal half; length of segment 8 The species is distributed in southeastern greater than one-half width. Pronotum widest Canada and eastern United States (Fig. 106). It before base; l:w ratio = 1:1.8–1:1.9; hind angles is known from the following provinces and weakly obtuse, not prolonged posteriorly. Elytra states: Nova Scotia, Canada; Massachusetts, widest at base, weakly attenuated apically; Maine, New Hampshire, New Jersey, and North sutural striae impressed; transverse strigae mod- Carolina, United States. Most of the specimens erately widely spaced, 25 per mm; elytra shin- were collected by H.C. Fall in Massachusetts. ing, microsculpture absent in both sexes. Male protarsus (Fig. 63) subequal in length to proti- Habitats bia; tarsal segment 1 wider than tibial apex. Ecology: myrmecophile. Adults of the beetle Aedeagus (Figs. 64, 65) with basal piece longer have a preference for the nests of Formica than parameres; median lobe about three- integra; 124 specimens were associated with fourths length of parameres, triangular in cross F. integra, 2 were associated with F. obscu- section, with median dorsal carina, apex riventris, and 1 was associated with F. rubicunda

Figs. 61–68. Nemadus (Nemadus) integer: 61, outline of body; 62, antenna; 63, male protibia and protarsus; 64, aedeagus, dorsal; 65, aedeagus, lateral; 66, male genital segment, ventral; 67, spermatheca; 68, female sternite 8. Scale bar = 1.0 mm (Fig. 61), 0.2 mm (Fig. 67), and 0.4 mm (Figs. 62–66, 68).

63 64

61 65

66 67

Emery, 1893. All specimens for which collecting die through the summer and autumn. We pre- data are available were associated with ants. sume that the eggs are laid in spring and early summer, that the larvae develop and pupate in Seasonality mid or late summer, and that adults emerge in Adults have been taken most often in the autumn. The only observations of larvae of the spring in April and May, in the following beetles in ant nests are from July by A.F. New- months and numbers: April, 125; May, 52; ton, and they support this conclusion. There June, 4; July, 9; September, 2; and October, 3. was no reproduction in the captive specimens. Adults have been taken with ants in the follow- Adults obviously disperse by flight to new ant ing months and numbers: April, 119; May, 38; nests, but data on specimens taken from nests June, 2; July, 7; September, 1; and October, 2. do not clearly indicate whether this dispersal is in the autumn or spring. Behavior, longevity, and reproduction Adults of N. integer were collected by S.B. Peck at Ashby and in the Middlesex Fells, both near Boston, Massachusetts, to observe behav- Nemadus (Nemadus) iour and to obtain larvae. They were kept in myrmecophilus Jeannel moist plaster-bottomed petri dishes, in constant darkness at 14 °C, and fed moist baker’s yeast. (Figs. 69–75, 105) Of those kept with ants, none were observed to interact with the ants. Of 22 adults collected in Nemadus myrmecophilus Jeannel, 1936: 170. October, 1 died the next April, 11 in May, 6 in Type in MNHN, seen. Type locality: Iowa July, 2 in August, and 2 in October. Of 7 adults City, Iowa. collected in May, all were dead by October. Ptomaphagus parasitus (LeConte), Wickham This suggests that late-autumn adults are re- 1896: 370, 1900: 3, misidentification. cently emerged and that they pass the winter, Ptomaphagus pusio, Wickham 1892: 321, mis- reproduce the next spring or early summer, and identification.

Figs. 69–75. Nemadus (Nemadus) myrmecophilus: 69, outline of body; 70, antenna; 71, male protibia and protarsus; 72, aedeagus, dorsal; 73, male genital segment, ventral; 74, spermatheca; 75, female sternite 8. Scale bar = 1.0 mm (Fig. 69), 0.2 mm (Fig. 74), and 0.4 mm (Figs. 70–73, 75).

72 71

69 75

73 74

Material examined parameres; median lobe flat, about two-thirds Thirty-seven specimens. length of parameres, evenly narrowed to small apical projection; parameres broad, conchoidal, with rounded apices. Male genital segment Diagnosis (Fig. 73) about as long as wide; anterior Nemadus myrmecophilus can be distinguished apophysis moderately long. Spermatheca from all other North American species of (Fig. 74) strongly curved; invagination bearing Nemadus parasitus by the characters of the spe- sclerotized teeth. Female sternite 8 (Fig. 75) tri- cies group and the characters given in the key. angular with short anterior apophysis.

Redescription Distribution Body (Fig. 69) elliptical, slightly narrowed This species is known only from the type lo- posteriorly. Total length 2.1–2.2 mm; greatest cality, Iowa City, Iowa, except for a single spec- width 1.2–1.3 mm. Color reddish brown; imen from an unspecified locality in Kansas pronotum with area of darker pigmentation. Pu- (Fig. 105). bescence of moderate length, golden, subappressed. Antennae (Fig. 70) of moderate Habitats length, barely reaching base of pronotum; seg- Ecology: myrmecophile. Adults are associ- ment 3 slightly shorter than segment 2; segment ated with the nests of Formica subsericea. 7 weakly constricted basally, length of segment 8 less than one-half width. Pronotum widest be- Seasonality fore base; l:w ratio = 1:1.7–1:1.9; hind angles Based on specimen label data, this species subquadrate, not prolonged posteriorly. Elytra was collected only in April and May, as verified widest in basal half, slightly narrowed apically; by Wickham (1892, 1896, 1900). sutural striae impressed; transverse strigae moderately widely spaced, 25 per mm; microsculpture of several fine lines between and parallel to Nemadus (Nemadus) parasitus strigae present in female only; male elytra shin- (LeConte) ing. Hind wings fully formed. Male protarsus (Figs. 76–83, 107) (Fig. 71) subequal in length to protibia; tarsal segment 1 subequal in width to tibial apex. Catops parasitus LeConte, 1853: 282. Lectotype Aedeagus (Fig. 72) with basal piece longer than here designated as male in LeConte collection,

Figs. 76–83. Nemadus (Nemadus) parasitus: 76, outline of body; 77, antenna; 78, male protibia and protarsus; 79, aedeagus, dorsal; 80, aedeagus, lateral; 81, male genital segment, ventral; 82, spermatheca; 83, female sternite 8. Scale bar = 1.0 mm (Fig. 76), 0.2 mm (Fig. 82), and 0.4 mm (Figs. 77–81, 83).

79 78

80 76

82 83 81

MCZC, with blue label “C. parasitus/N.Y. (Fig. 77) of moderate length, barely reaching March. Lec.”, red type label with number base of pronotum; segment 3 shorter than seg- 3156, and our lectotype label; seen and dis- ment 2; segment 7 not constricted in basal half; sected. Type locality: New York. Murray 1856: length of segment 8 greater than one-half 460. width. Pronotum widest before base; l:w ratio = Ptomaphagus parasitus (LeConte), Horn 1880: 1:1.7–1:1.8; hind angles subquadrate, not pro- 264; Hamilton 1888: 162; Schwarz 1889: longed posteriorly. Elytra widest in basal half, 242; Blatchley 1910: 281. weakly attenuated apically; sutural striae im- Catopomorphus parasitus (LeConte), Wasmann pressed; transverse strigae moderately widely 1894: 127. spaced, 27–28 per mm; microsculpture of sev- Adelops parasitus (LeConte), Jeannel 1922: 43; eral fine lines between and parallel to strigae Hatch 1928: 169. present in female only; male elytra shining. Hind Nemadus parasitus (LeConte), Hatch 1933: wings fully formed. Male protarsus (Fig. 78) 192; Jeannel 1936: 170; Fall 1937: 338; subequal in length to protibia; tarsal segment 1 Salgado 1999: 37. wider than tibial apex. Aedeagus (Figs. 79, 80) with basal piece longer than parameres; median Diagnosis lobe flat, short, abruptly narrowed to small api- Nemadus parasitus can be distinguished from cal projection; parameres broad, conchoidal, all other North American species of Nemadus with rounded apices. Male genital segment by the characters of the parasitus species group (Fig. 81) about as long as wide; anterior apo- and the characters given in the key. physis moderately long. Spermatheca (Fig. 82) moderately curved; invagination bearing sclero- Redescription tized teeth. Female sternite 8 (Fig. 83) triangu- Body (Fig. 76) oval, weakly attenuated poste- lar with short anterior apophysis. riorly. Total length 2.0–2.2 mm; greatest width 1.1–1.2 mm. Color reddish brown, pronotum darker in some individuals. Pubescence of mod- Material examined erate length, golden, subappressed. Antenna Three hundred forty-eight specimens.

Figs. 84–91. Nemadus (Nemadus) tenuitarsis: 84, outline of body; 85, antenna; 86, male protibia and protarsus; 87, aedeagus, dorsal; 88, aedeagus, lateral; 89, male genital segment, ventral; 90, spermatheca; 91, female sternite 8. Scale bar = 1.0 mm (Fig. 84), 0.2 mm (Fig. 90), and 0.4 mm (Figs. 85–89, 91).

86 87

88 84

90 91

Distribution in a blacklight trap, and by sweeping sweet fern The species is distributed across the eastern (Comptonia spp.; Myricaceae). United States, through New England and south to Virginia and Alabama (Fig. 107). We have Seasonality seen specimens from the following states: Ala- Adults have been taken most frequently in bama, Connecticut, District of Columbia, Mas- April and May and in the following months and sachusetts, New Hampshire, New Jersey, New numbers: March, 20; April, 131; May, 69; June, York, Pennsylvania, and Virginia. Most of the 3; July, 4; August, 10; September, 1; October, specimens were collected by H.C. Fall in Mas- 1; and December, 32. Adult beetles have been sachusetts. Fall (l937, p. 338) indicated records taken with ants in the following months and from Colorado and Texas but we have not been numbers: March, 15; April, 75; May, 33; Au- able to confirm these. gust, 5; and December, 32 (in Alabama).

Habitats Nemadus (Nemadus) tenuitarsis Ecology: myrmecophile. There is an apparent Jeannel preference for the nests of Formica subsericea. (Figs. 84–91, 105) Adults of the beetles have been taken with the following ant species in the following numbers: Nemadus tenuitarsis Jeannel, 1936: 170; F. subsericea, 122; F. integra,6;F. pergandei,5; Salgado 1999: 37. Type male in MNHN, F. schaufussi,4;F. exsectoides,1;Acanthomyops seen. Type locality: Jeannette, Pennsylvania. claviger,1;Lasius alienus,1;Camponotus pennsylvanicus, 1. Some labels indicate Material examined F. fusca L., 1758 as the host but no specimens of Eighty-seven specimens. this ant species were verified, and it is likely that these records refer to F. subsericea. Specimens Diagnosis were also taken in low numbers in baited pitfall Nemadus tenuitarsis can be distinguished traps, in flight-intercept traps, in malaise traps, from all other North American species of

Figs. 92–99. Nemadus (Nemadus) triangulum: 92, outline of body; 93, antenna; 94, male protibia and protarsus; 95, aedeagus, dorsal; 96, aedeagus, lateral; 97, male genital segment, ventral; 98, spermatheca; 99, female sternite 8. Scale bar = 1.0 mm (Fig. 92), 0.2 mm (Fig. 98), and 0.4 mm (Figs. 93–97, 99).

94 95

92 96

98 99

Nemadus by the characters of the parasitus spe- (Fig. 89) slightly longer than wide; anterior cies group and the characters given in the key. apophysis moderately long. Spermatheca (Fig. 90) moderately curved; invagination bear- Redescription ing strong setae. Female sternite 8 (Fig. 91) tri- Body (Fig. 84) elliptical, slightly narrowed angular with short anterior apophysis. posteriorly. Total length 2.0–2.3 mm; greatest width 1.1–1.3 mm. Color reddish brown; prono- Distribution tum with area of darker pigmentation. Pubes- This species occurs in eastern Canada and cence of moderate length, golden, subappressed northeastern United States from New York west to suberect. Antenna (Fig. 85) of moderate to Illinois and Wisconsin (Fig. 105). It is known length, reaching base of pronotum; segment 3 from the following provinces and states: Que- slightly shorter than segment 2; segment 7 bec and Ontario, Canada; Illinois, New York, weakly constricted basally; length of segment 8 Ohio, Pennsylvania, and Wisconsin, United more than one-half width. Pronotum widest be- States. fore base; l:w ratio = 1:1.7–1:1.9; hind angles subquadrate, not prolonged posteriorly. Elytra Habitats widest in basal half, slightly narrowed apically; sutural striae impressed; transverse strigae mod- Ecology: myrmecophile. This species has erately widely spaced, 27–28 per mm; elytra been collected in association with Formica sub- shining, microsculpture absent in both sexes. sericea (three records), F. ulkei (one record), Hind wings fully formed. Male protarsus and F. glacialis (one record). It has also been (Fig. 86) subequal in length to protibia; tarsal taken in pan traps and a flight-intercept trap. segment 1 narrower than tibial apex. Aedeagus (Figs. 87, 88) with basal piece longer than Seasonality parameres; median lobe flat, about three-fourths Adults have been taken most often in April length of parameres, evenly narrowed to small and May, in the following months and numbers: apical projection; parameres broad, conchoidal, February, 3; March, 2; April, 11; May, 22; June, with rounded apices. Male genital segment 1; July, 4; August, 5; and October, 4.

Figs. 100–101. 100, collection localities of Neoeocatops decipiens; 101, collection localities of Nemadus (Laferius) brachyderus.

Figs. 102–103. 102, collection localities of Nemadus (Nemadus) pusio; 103, collection localities of Nemadus (Nemadus) horni (open circles, cave records; black dots, epigean (non-cave) records).

Fig. 104. Collection localities of Nemadus (Nemadus) browni (black dots), Nemadus (Nemadus) criddlei (stars), and Nemadus (Nemadus) falli (asterisks).

Fig. 105. Collection localities of Nemadus (Nemadus) gracilicornis (black dots), Nemadus (Nemadus) myrmecophilus (asterisks), and Nemadus (Nemadus) tenuitarsis (black dots in circles).

Figs. 106–107. 106, collection localities of Nemadus (Nemadus) integer with Formica integra as host ant; 107, collection localities of Nemadus (Nemadus) parasitus.

Fig. 108. Collection localities of Nemadus (Nemadus) triangulum with several host ants, mostly Camponotus noveboracensis and C. pennsylvanicus.

Nemadus (Nemadus) triangulum Distribution Jeannel The species is distributed in southeastern Canada and northeastern United States south- (Figs. 92–99, Fig. 108) wards to Virginia, Alabama, and Missouri (Fig. 108). Most of the specimens were col- Nemadus triangulum Jeannel, 1936: 171. Type lected by H.C. Fall in Massachusetts. We have male in MNHN, seen. Type locality: Cincin- seen specimens from the following provinces nati, Ohio. and states: Nova Scotia, Ontario, and Quebec, Nemadus obliquus Fall 1937: 339. Type female Canada; Alabama, Arkansas, Connecticut, in MCZC, seen and dissected. Type locality: Georgia, Illinois, Kentucky, Massachusetts. Tyngsboro, Massachusetts. New synonomy. Maryland, Maine, Missouri, North Carolina, New Hampshire, New Jersey, New York, Ohio, Pennsylvania, Tennessee, and Virginia, United Material examined States. Two hundred thirty-seven specimens. Habitats Diagnosis Ecology: myrmecophile. Adults of the beetles Nemadus triangulum can be distinguished have been taken with the following ant species from all other North American species of in the following numbers: Camponotus nove- Nemadus by the characters of the parasitus spe- boracensis, 19; C. pennsylvanicus, 12; C. chro- cies group and the characters given in the key. maiodes,5;F. exsectoides,2;F. nitidiventris,1. Specimens have also been taken frequently in litter of logs, stumps, and leaves, in tree holes, and Redescription with flight-intercept traps. Body (Fig. 92) oval, moderately attenuated posteriorly. Total length 1.9–2.3 mm; greatest Seasonality width 1.2–1.4 mm. Color reddish brown; prono- Most adult beetles have been taken in May. tum lacking area of darker pigmentation. Pubes- Adults have been taken with ants in the follow- cence short, silky, golden, appressed. Antenna ing months and numbers: April, 7; May, 14; (Fig. 93) of moderate length, barely reaching June, 11; July, 2, August, 2; September, 2; and base of pronotum; segment 3 slightly shorter October, 1. Total monthly numbers are March, than segment 2; segment 7 not constricted in 2; April, 29; May, 49; June, 40; July, 17; Au- basal half; length of segment 8 greater than gust, 19; September, 5; October, 3; November, one-half width. Pronotum widest before base; 2; and December, 3. l:w ratio = 1:1.9–1:2.0; hind angles weakly obtuse, not prolonged posteriorly. Elytra widest in Acknowlegements anterior half, moderately attenuated posteriorly; sutural striae impressed; transverse strigae mod- The field and laboratory work for this study erately widely spaced, 30 per mm; micro- were partially supported by operating and dis- sculpture of several fine lines between and covery grants to S.B.P. from the Natural Sci- parallel to strigae present in female only; male ences and Engineering Research Council of elytra shining. Hind wings fully formed. Male Canada. The many collection curators and pri- protarsus (Fig. 94) subequal in length to proti- vate collectors who loaned or contributed mate- bia; tarsal segment 1 slightly wider than tibial rial are all gratefully thanked. A.F. Newton has apex. Aedeagus (Figs. 95, 96) with basal piece provided much help over the years. Dr. Th. longer than parameres; median lobe flat, about Deuve (MNHN), M. Perreau, and Hume two-thirds length of parameres, evenly narrowed Douglas were especially helpful in arranging to small apical projection; parameres broad, the loan of the Jeannel types. Jiri Vavra conchoidal, with truncate apices. Male genital (Ostrava), G. Cucodoro (Geneva), and Stefano segment (Fig. 97) about as long as wide; ante- Zoia (Milan) helped obtain comparative speci- rior apophysis moderately long. Spermatheca mens of Eocatops. Jarmila Kukalova-Peck gen- (Fig. 98) moderately curved; invagination bear- erously assisted on many long field trips. Stefan ing sclerotized teeth. Female sternite 8 (Fig. 99) Cover (MCZC) provided the identifications of broadly triangular with short anterior apoph- all the ants associated with the beetle speci- ysis. mens.

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