Slime-Mold Beetles Of The Genus Agathidium Panzer In North And Central America, Part Ii. Coleoptera: Leiodidae

SLIME-MOLD BEETLES OF THE GENUS AGATHIDIUM PANZER IN NORTH AND CENTRAL AMERICA, PART II. COLEOPTERA: LEIODIDAE KELLY B. MILLER Department of Entomology, Cornell University Current address: Department of Integrative Biology Brigham Young University Provo, Utah 84042 ([email protected]) QUENTIN D. WHEELER Department of Entomology, Cornell University Current address: Department of Entomology The Natural History Museum, Cromwell Road South Kensington SW7 5BD, United Kingdom ([email protected]) BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 291, 167 pp., 380 figures Issued March 24, 2005 Copyright q American Museum of Natural History 2005 ISSN 0003-0090 F RONTISPIECE . Male Agathidium vaderi, new species, dorsal habitus. Original by Byron Alexander. 3 CONTENTS Abstract ....................................................................... 5 Introduction .................................................................... 5 Agathidium concinnum Species Group ............................................. 5 Key to A. concinnum Species Group .............................................. 8 A. akallebregma Miller and Wheeler, n.sp. ...................................... 11 A. mollinum Fall, 1934b ...................................................... 13 A. angulare Mannerheim, 1852 ................................................ 14 A. hatchi Wheeler, 1977 ...................................................... 19 A. concinnum Mannerheim, 1852 .............................................. 21 Agathidium pulchrum Species Group ............................................. 24 Key to A. pulchrum Species Group .............................................. 24 A. difforme (LeConte, 1850) .................................................. 32 A. maculosum Brown, 1928 ................................................... 39 A. pulchrum LeConte, 1853 ................................................... 41 A. amae Miller and Wheeler, n.sp. ............................................. 45 A. laetum Fall, 1934b ........................................................ 46 A. athabascanum Fall, 1934b ................................................. 47 A. columbianum Fall, 1934b .................................................. 49 A. rusticum Fall, 1934b ...................................................... 49 A. repentinum Horn, 1880 .................................................... 50 A. marae Miller and Wheeler, n.sp. ............................................ 52 A. rotundulum Mannerheim, 1852 ............................................. 53 A. aristerium Wheeler, 1987 .................................................. 54 A. atronitens Fall, 1934b ..................................................... 55 A. oregonense Miller and Wheeler, n.sp. ........................................ 58 A. picipes Fall, 1934b ........................................................ 59 A. hamulum Miller and Wheeler, n.sp. ......................................... 61 A. politum LeConte, 1866 .................................................... 62 Agathidium compressidens Species Group ........................................ 63 Key to A. compressidens Species Group .......................................... 64 A. fenderi Hatch, 1957 ....................................................... 64 A. vesperpressidens Hatch, 1957 ............................................... 67 A. compressidens Fall, 1934b ................................................. 68 Agathidium iota Species Group .................................................. 69 A. iota Miller and Wheeler, n.sp. .............................................. 69 Agathidium oniscoides Species Group ............................................ 70 Key to A. oniscoides Species Group ............................................. 72 Agathidium kimberlae Subgroup ................................................. 85 A. vaderi Miller and Wheeler, n.sp. ............................................ 87 A. kimberlae Miller and Wheeler, n.sp. ......................................... 89 Agathidium aztec Subgroup ..................................................... 91 A. bituberculum Miller and Wheeler, n.sp. ...................................... 91 A. oedema Miller and Wheeler, n.sp. ........................................... 93 A. gomezae Miller and Wheeler, n.sp. .......................................... 94 A. hidalgoense Miller and Wheeler, n.sp. ....................................... 95 A. skoliosternum Miller and Wheeler, n.sp. ..................................... 96 A. erythromelas Miller and Wheeler, n.sp. ...................................... 97 A. rhamphastes Miller and Wheeler, n.sp. ...................................... 99 A. megoniscoides Miller and Wheeler, n.sp. .................................... 100 A. grumum Miller and Wheeler, n.sp. ......................................... 101 A. triangularum Miller and Wheeler, n.sp. ..................................... 102 4 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY A. lobosternum Miller and Wheeler, n.sp. ...................................... 103 A. potosii Miller and Wheeler, n.sp. ........................................... 103 A. popocatepetlae Miller and Wheeler, n.sp. ................................... 105 A. hyle Miller and Wheeler, n.sp. ............................................. 106 A. stenomma Miller and Wheeler, n.sp. ........................................ 107 A. tribulosum Miller and Wheeler, n.sp. ....................................... 108 A. tribulograndum Miller and Wheeler, n.sp. ................................... 109 A. invisitatum Miller and Wheeler, n.sp. ....................................... 110 A. multidentatum Miller and Wheeler, n.sp. .................................... 112 A. sejunctum Miller and Wheeler, n.sp. ........................................ 113 A. grandidentatum Miller and Wheeler, n.sp. ................................... 114 A. andersoni Miller and Wheeler, n.sp. ........................................ 115 A. disgregum Miller and Wheeler, n.sp. ....................................... 116 A. oaxacaense Miller and Wheeler, n.sp. ...................................... 117 A. oculeum Miller and Wheeler, n.sp. ......................................... 118 A. recurvatum Miller and Wheeler, n.sp. ....................................... 119 A. impensum Miller and Wheeler, n.sp. ........................................ 120 A. cheneyi Miller and Wheeler, n.sp. .......................................... 121 A. tenangoense Miller and Wheeler, n.sp. ...................................... 122 A. rumsfeldi Miller and Wheeler, n.sp. ........................................ 123 A. hirsutum Miller and Wheeler, n.sp. ......................................... 124 A. cortezi Miller and Wheeler, n.sp. ........................................... 125 A. tumidiventre Miller and Wheeler, n.sp. ...................................... 126 A. aztec Miller and Wheeler, n.sp. ............................................ 127 A. iridescens Miller and Wheeler, n.sp. ........................................ 128 Agathidium dentigerum Subgroup ............................................... 130 A. dentigerum Horn, 1880 ................................................... 130 A. stephani Miller and Wheeler, n.sp.. ......................................... 131 A. akrogeneios Miller and Wheeler, n.sp. ...................................... 133 A. framea Miller and Wheeler, n.sp. .......................................... 134 A. appalachium Miller and Wheeler, n.sp. ..................................... 137 A. pocahontasae Miller and Wheeler, n.sp. ..................................... 138 A. carolinense Miller and Wheeler, n.sp. ...................................... 139 A. gallititillo Miller and Wheeler, n.sp. ........................................ 140 A. divaricatum Miller and Wheeler, n.sp. ...................................... 142 A. bushi Miller and Wheeler, n.sp. ............................................ 143 A. georgiaense Miller and Wheeler, n.sp. ...................................... 144 Agathidium microphthalmum Subgroup .......................................... 145 A. chauliodoum Miller and Wheeler, n.sp. ..................................... 145 A. microphthalmum Miller and Wheeler, n.sp. .................................. 147 A. nimbosilva Miller and Wheeler, n.sp. ....................................... 148 Agathidium oniscoides Subgroup ............................................... 149 A. oniscoides Palisot de Beauvois, 1817 ....................................... 149 A. rubellum Fall, 1934b ..................................................... 154 A. exiguum Melsheimer, 1844 ................................................ 156 A. fawcettae Miller and Wheeler, n.sp. ........................................ 161 Species of uncertain status ..................................................... 163 A. difficile Matthews, 1887 .................................................. 163 A. mexicanum Hendrichs, 1979 ............................................... 164 Checklist of Agathidium Species of North and Central America .................... 164 Acknowledgments ............................................................ 166 References ................................................................... 166 5 ABSTRACT The Agathidium concinnum,A. pulchrum,A. compressidens,A. iota, and A. oniscoides groups are revised. The following new species are described: A. akallebregma, n.sp., A. akrogeneios, n.sp., A. amae, n.sp., A. andersoni, n.sp., A. appalachium, n.sp., A. aztec, n.sp., A. bituberculum, n.sp., A. bushi, n.sp., A. carolinense, n.sp., A. chauliodoum, n.sp., A. cheneyi, n.sp.,A.cortezi,n.sp., A.disgregum,n.sp.,A.divaricatum,n.sp.,A.erythromelas,n.sp.,A. fawcettae,n.sp.,A.framea,n.sp., A.gallititillo,n.sp.,A.georgiaense,n.sp.,A.gomezae,n.sp., A. grandidentatum, n.sp., A. grumum, n.sp., A. hamulum, n.sp., A. hidalgoense, n.sp., A. hirsutum, n.sp., A. hyle, n.sp., A. impensum, n.sp., A. invisitatum, n.sp., A. iota, n.sp., A. iridescens, n.sp., A. kimberlae, n.sp., A. lobosternum, n.sp., A. marae, n.sp., A. megoniscoides, n.sp., A.microphthalmum,n.sp.,A.multidentatum,n.sp., A.nimbosilva,n.sp.,A.oaxacaense,n.sp., A. oculeum, n.sp., A. oedema, n.sp., A. oregonense, n.sp., A. pocahontasae, n.sp., A. popocatepetlae, n.sp., A. potosii, n.sp., A. recurvatum, n.sp., A. rhamphastes, n.sp., A. rumsfeldi, n.sp.,A.sejunctum,n.sp.,A.skoliosternum,n.sp.,A.stenomma,n.sp.,A.stephani,n.sp.,A. tenangoense,n.sp.,A.triangularum,n.sp.,A.tribulograndum,n.sp.,A.tribulosum,n.sp., A. tumidiventre, n.sp., A. vaderi, n.sp., A. vesperpressidens, n.sp. The following new synonyms are proposed: A. assimile Fall, A. municeps Fall, and A. falli Hatch 5 A. angulare Mannerheim, new synonyms; A. alticola Fall 5 A. athabascanum Fall, new synonym; A. contiguum Fall, A. varipunctatum Hatch, and A. striolum Hatch 5 A. picipes Fall, new synonyms; and A. californicum Horn and A. alutaceum Fall; 5 A. exiguum Melsheimer, new synonyms. Lectotypes are designated for the following species: A. angulare,A. californicum,A. concinnum Mannerheim, A. dentigerum Horn, A. difficile Matthews, A. effluens Mannerheim, A. exiguum, A. globatile LeConte, A. laetum Fall, A. pulchrum LeConte, A. rotundulum Mannerheim, A. rubellum Fall, A. ruficorne LeConte, and Phalacrus difformis LeConte. INTRODUCTION This is the second of two parts of a taxonomic revision of the species of the genus Agathidium Panzer (Coleoptera, Leiodidae) of North and Central America. A general introduction to the revision, a detailed account of the external morphology of a representative species, keys to described New World genera of Agathidiini and species groups within the scope of this study, and a summary of host associations, primarily with Myxomycetes (slime-molds), were presented in part I (Wheeler and Miller, in press), along with taxonomic treatments of the A. revolvens,A. sexstriatum, and A. brevisternum species groups. Below are presented the remaining species in the A. compressidens,A. concinnum,A. iota,A. pulchrum, and A. oniscoides species groups, and a checklist for all Agathidium species of North and Central America. Materials and methods are the same for this portion of the revision as for part I. For measurements: OHW (ocular head width) 5 greatest width of head across eyes; MDL (median head length) 5 head length from base of labrum to posterior margin of cranium; PNW (pronotal width) 5 greatest width of pronotum; PNL (pronotal length) 5 length of pronotum along midline; ELW (elytral width) 5 greatest width of combined elytra; SEL (postscutellar elytral length) 5 length of elytra along suture from posterior margin of scutellum to apex; MTL (metasternal length) 5 length of metasternum along midline; MTW (metasternal width) 5 greatest width of metasternum; TBL (total body length) 5 MDL 1 PNL 1 SEL. AGATHIDIUM CONCINNUM SPECIES GROUP D IAGNOSIS : The group is characterized by the well-developed postocular temporum which extends the lateral margin of the head posteriorly beyond the eye between 0.5 and 1.0 times the eye length (figs. 1–5). The anterior portion of the mesosternum is relatively short, much shorter than the posterior portion which is moderately to strongly concave. The humeral angles of the elytra are relatively angulate. D ISCUSSION : This species group appears to correspond to the subgenus A.(Cyphoceble) Thomson based on the long temporum. In North America the group contains five species that fall into two subgroups. The first includes 6 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 1. Agathidium angulare, dorsal habitus. A. akallebregma,A. angulare, and A. mollinum which share the presence of a male mandibular horn (e.g., figs. 17–20), moderately elongate body shape, and strongly concave posterior portion of the mesosternum. The second group includes A. hatchi and A. concinnum which are large species (very large in the case of A. hatchi) and are characterized by the lack of a male mandibular horn, a broad, depressed body form (e.g., fig. 2), and weakly concave posterior portion of the mesosternum. Of course, given the lack of a cladistic analysis it is impossible to say which of these states is derived. However, given the presence of a prominent 2005 7MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Fig. 2. Agathidium concinnum, dorsal habitus. left mandibular horn in at least some of these species (certainly an apomorphy within the Agathidiini), they may be related to the members of the A. brevisternum and A. pulchrum species groups ( 5 subgenus A. (Neoceble)auctorum). Some species of the A. pulchrum group have a relatively long temporum, even approaching half the length of the eye. The gradational nature of this feature in Nearctic Agathidium further suggests that these species groups may not each be monophyletic. These species occur in far western and 8 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 3–16. Agathidium concinnumand A. pulchrum-group species. 3–15. Heads, dorsal aspect: 3, A. akallebregma.4, A. hatchi.5, A. concinnum.6, A. laetum.7, A. athabascanum.8, A. columbianum. 9, A. rusticum.10, A. repentinum.11, A. rotundulum.12, A. atronitens.13, A. oregonense.14, A. hamulum.15, A. politum.16, A. rusticum, left antenna, dorsal aspect. Bars 5 0.5 mm, except fig. 16 bar 5 0.25 mm. northern North America east to the northeastern part of the continent. KEY TO A. CONCINNUM SPECIES GROUP 1. Anterior clypeal margin strongly excavated, anterolateral margins of frons extending prominently beyond anterior margin of clypeus (figs. 1, 3, 18, 20); posterior portion of mesosternum strongly concave, nearly vertical posteriorly; horn often present on male left mandible (figs. 17–20) ........... 2 —Anterior clypeal margin not excavated, margin extending to or beyond level of anterolateral margins of frons (figs. 2, 4, 5); posterior por- 2005 9MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Figs. 17–30. Agathidium concinnumand A. pulchrum-group species, male heads showing left mandibular horn: 17, 18, A. mollinum:17, left lateral; 18, dorsal. 19, 20, A. angulare:19, left lateral; 20, dorsal. 21, 22, A. pulchrum:21, left lateral; 22, dorsal. 23, 24, A. marae:23, left lateral; 24, dorsal. 25, 26, A. aristerium:25, left lateral; 26, dorsal. 27, 28, A. atronitens:27, left lateral; 28, dorsal. 29, 30, A. picipes:29, left lateral; 30, dorsal. Bars 5 0.5 mm. tion of mesosternum concave, but not strongly so, not approaching vertical posteriorly; horn absent on male left mandible .......... 4 2(1). Anterior surface of head irregular, frons swollen and expanded around margins of clypeus, especially laterad of clypeus . . ........ A. akallebregma, new species — Anterior surface of head evenly curved over surface of frons and clypeus, frons not particularly swollen or only slightly so laterad of clypeus ................ 3 3(2). Size smaller (TBL , 2.5 mm) ; male mandibular horn when fully developed long, extending from medial surface of mandible in very strong C-shaped curve to left and back over mandible (figs. 17, 18); median lobe of aedeagus in lateral aspect slender and relatively straight, apical por- 16 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Head very broad (figs. 1, 20) (MDL/OHW 5 0.62–0.73), convex, without transverse, medial crease; temporum prominent, about 0.6 to 1.0 times eye length when viewed in dorsal aspect (figs. 1, 20); head finely and sparsely to moderately densely punctate, some specimens with interspersed micropunctules, surface between punctures shiny, a few specimens with indistinct, fine microreticulation; eyes large, rounded, finely faceted; frontoclypeal suture obsolete medially; clypeal margin strongly excavate (figs. 1, 20); anterior margin of labrum entire; antennomere ratios: length I:II:III 5 2.3:1.0:1.4, width VII:VIII:IX 5 1.1:1.0:2.5. Pronotum broad (PNL/PNW 5 0.55–0.57), laterally not strongly produced, anterolateral angle subquadrate to rounded, posterolateral angle distinctive, but rounded; punctation similar to that of head. Elytra moderately elongate (SEL/ELW 5 1.01–1.20); punctation coarser than that of pronotum, some specimens relatively densely punctate, others more sparse, some specimens with numerous micropunctules between larger punctures; sutural stria about three-fifths length of elytron on most specimens. Mesosternum strongly concave posteriorly, with prominent lobe extending ventrally between mesocoxae, anterior portion less than one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum broad medially (MTL/MTW 5 0.33–0.35), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres moderately broad and with moderately large ventral field of spatulate setae; mandibular horn, when fully developed, very long, curving anteriorly, then posteriorly over front of head (figs. 19, 20), subcircular throughout in cross-section, punctate and setose along its length, apical fovea prominent with large cluster of setae, some specimens with mandible variously thickened, others without modification; metafemur moderately slender, unmodified; metasternal fovea small, round, anterior, moderately large brush of fine, dense setae. Median lobe in lateral aspect large, moderately robust, strongly curved basally with slight constriction, lateral margins evenly converging, apical portion flat, slender, curved dorsad, apex abruptly expanded (fig. 36); in ventral aspect moderately broad, lateral margins subparallel, apical portion triangular, apex pointed, orifice large (fig. 35); operculum long and moderately broad, apically narrowly emarginate, inconspicuous and hyaline (fig. 35); lateral lobes long, slender, curved basally, apices rounded with 2 long setae (figs. 35, 36). Female tarsi 5-4-4. D ISTRIBUTION : This species is widespread across northern North America from Alaska to New Hampshire and south in western North America to Arizona (fig. 42). Records from Indiana and Arkansas suggest that the species may be even more widespread than records indicate. S PECIMENS E XAMINED : ‘‘W.T’’ [probably Washington Territory] (1, MCZC). CANADA: Alberta: Demmitt, 3 km W 90 km NW Grand Prairie, 12 Jun 1984, poplar forest, FIT, S and J Peck (1, PECK); 3.8 mi N Cadomin, 4 Aug 1985, lodgepole pine forest, FIT, RS Anderson (40, CNCI); Sibbold Flats Rec. Area, 6 Sep 1981, FIT, RS Anderson (10, CNCI); George Lake 50 km NW Edmonton, 11 Jun 1984, spruceaspen forest, FIT, S and J Peck (2, CNCI). British Columbia: Pink Mountain 252 km S Ft Nelson, Rt 97, 12 Jun 1984, aspen forest, FIT, S and J Peck (14, PECK); Lorna, 17 Aug 1934, R Hopping (1, CASC); Mt Robson P Park, Berg Lake Trailhead, 11 Jul 1984, RS Anderson (10, PECK); Bear Lake, 20 Aug 1969, sifting, BF and JL Carr (1, CARR); 37 km W Ft Nelson, Ak Hwy, 12 Jun 1984, aspen-spruce forest, FIT, S and J Peck (12, PECK); Mt Robson P Park, Berg Lake Trailhead, 11 Jul 1984, RS Anderson (44, CNCI); 14 km E Coal River, 160 km E Watson Lake, YT, 14 Jun 1984, spruce-alder forest, malaise FIT, S and J Peck (12, PECK); 5.6 mi E E Border Glacier Natl Park on #1, 11 May 1984, Columbia forest, RS Anderson (3, CNCI); Mount Robson Prov. Park, Berg Lake Trailhead, 4 Jul 1984, cedar/hemlock forest litter, RS Anderson (1, PECK); Kootenay Natl Park, 17 Jul 1985, on Stemonitis, SL Stephenson (3, AMNH); Pink Mt, 252 km S Ft Nelson, rt 97, 5 Jun 1984, aspen forest, FIT, S and J Peck (10, CNCI); Stanley, 19 Jul 1932, N Graham (2, CASC); Manning Prov. Park, 19 Jun 1979, alpine meadow, pitfall traps, CD Dondale (20, CNCI); Lorna, 19 Jul 1926, H Richmond (1, CASC). Manitoba: Grass River Prov. Park, 100 rd km SE FlinFlon, 7 Jun 1984, pine-aspen forest, FIT, S and J Peck (4, CNCI); Grass Riv. Prov. Park, 100 rd KM SE FlinFlon, 7 Jun 1984, pineaspen forest, FIT, S and J Peck (2, PECK). Ontario: Stittsville, 5 Sep 1976, M Sanborne (1, PECK); 25 km W Ignace, 75 km E Dryden, 5 Jun 2005 17MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II 1984, fir-maple forest, FIT, S and J Peck (1, PECK); Kemptville, 2 Oct 1984, BRI staff (1, CNCI); Gloucester, 15 Sep 1984, M Sanborne (1, CNCI); Chaffey Locks QUSB, 5 Oct 1980, forest, malaise FIT, S Peck (16, CNCI); Smooth Rock Falls 56 km NW Cochrane, 3 Jun 1984, mixed spruce-birch, evening car netting, S and J Peck (1, PECK); Haley Sta. 15 km NW Refrew, 2 Nov 1979, mixed forest, malaise, S and J Peck (1, CNCI); Lake Superior Prov. Park, Algoma Co., 9 Sep 1980, R Baranowski (1, LUND); 25 km W Ignace, 75 km E Dryden, 5 Jun 1984, fir-maple forest, FIT, S and J Peck (1, CNCI); Stittsville, DA Smith’s woods, 18 Oct 1979, forest, malaise FIT, S Peck (3, CNCI). Quebec: Hull, Gatinaeu Park, nr Pinks Lake, 30 Jun 1979, for. Mal. Trough, S Peck, A Davies (5, PECK); Gatineau Park nr Pinks Lake, 1 Oct 1979, forest, malaise, S Peck, A Davies (10, CNCI); Masham, Lac Philippe, Gatineua, 13 Dec 1981, on Brefeldia maxima, S Peck (21, CNCI); Hull, Gatinaeu Park, nr Pinks Lake, 24 Jun 1979, for. Mal. Trough, S Peck, A Davies (7, PECK); Masham lac Philip, Gatineau, 13 Sep 1981, reared from larvae, on Brefeldia maxima, S Peck, A Newton (3, FMNH). Saskatchewan: Jay Jay Lake Rt 102, 130 km NE Candle Lake, 8 Jun 1984, pine-fir forest, FIT, S and J Peck (1, CNCI); Jay Jay Lake, Rt 102, 130 km NE Candle Lake, 8 Jun 1984, pine-fir forest, FIT, S and J Peck (4, PECK). Yukon: Dawson City, km 45 Kondike Hwy, 9 Jul 1985, cut willow, Equisetum, FIT, SA Marshall (1, PECK); Engineer Creek Dempster Hwy, km 194, 19 Jun 1984, spruce-aspen forest, car top, S and J Peck (1, PECK); Ross River, 16 Jun 1984, aspen-willow, river terrace, malaise FIT, S and J Peck (9, PECK); Dempster Jct 40 km E Dawson, 19 Jun 1984, willow-spruce forest, malaise FIT, S and J Peck (26, PECK); Dempster Hwy, Tombstone Mtn Camp, 5 Jul 1985, willow shrub, FIT, SA Marshall (2, PECK); km147 Dempster Hwy, 6 Jul 1985, fen willows-spruce dung pan trap, SA Marshall (1, PECK); Simpson Lake, 81 km N Watson Lake, 15 Jun 1984, pine-willow forest, malaise FIT, S and J Peck (9, PECK); Moose Creek, 14 km NW Stewart Cross., 18 Jun 1984, stream willow thicket, malaise FIT, S and J Peck (20, PECK); Dempster Jct 40 km E Dawson, 19 Jun 1984, willow spruce forest, malaise FIT, S and J Peck (1, CNCI); Engineer Creek, Dempster Hwy km 194, 19 Jun 1984, willow spruce forest, malaise FIT, S and J Peck (1, CNCI); Engineer Creek, Dempster Hwy, km 194, 19 Jun 1984, aspenspruce forest, FIT, S and J Peck (38, PECK); Ross River, 16 Jun 1984, aspen willow river terrace, malaise FIT, S and J Peck (6, CNCI). UNITED STATES: Alaska: Chatanika Hwy 6, Chatanika River, 13 Aug 1984, floodplain birch forest litter, Berlese, S and J Peck (2, PECK); Houston, 24 Jun 1984, spruce-birch forest with moss, FIT, S and J Peck (8, PECK); Kenai Pen, E Skilak Rd Jct Rt 1, 25 Jun 1984, aspen-spruce forest, malaise FIT, S and J Peck (17, PECK); Nenana, 13 mi NE Rt 3, mi 318, 27 Jul 1984, birch-spruce forest, malaise FIT, S and J Peck (9, PECK); 15 mi N Fox 30 mi N Fairbanks, 30 Jul 1984, mixed birch woods, malaise FIT, S and J Peck (14, PECK); 11 mi S Anderson Jct Rt 3 mi 270, 23 Jun 1984, alder, poplar, spruce, FIT, S and J Peck (40, PECK); Chena Ridge Rd, 5 mi W Fairbanks, 22 Jun 1984, poplar forest, malaise FIT, S and J Peck (33, PECK); Houston, 24 Jun 1984, spruce-birch forest with moss, malaise FIT, S and J Peck (33, PECK); Kenai Pen, Tern Lake, Jct, rts 1 & 9, 25 Jun 1984, poplar, spruce, alder, malaise FIT, S and J Peck (23, PECK); Tok, 21 Jun 1984, spruce-poplar, FIT, S and J Peck (9, PECK); Chena River Rec. Area 30 mi E Fairbanks, 22 Jun 1984, spruce forest, malaise FIT, S and J Peck (5, PECK); 11 mi S Anderson Jct Rt 3 mi 270, 23 Jun 1984, alder, poplar, spruce, malaise FIT, S and J Peck (2, PECK); Big Delta, 21 Jun 1984, spruce-willow, malaise FIT, S and J Peck (43, PECK); Seward, Salmon Creek, 26 Jun 1984, mature poplar-spruce, malaise FIT, S and J Peck (12, PECK); Kenai Pen. Tern Lake Jct Rts 1 & 9, 25 Jun 1984, poplar-spruce-alder, malaise FIT, S and J Peck (4, PECK); Chena Ridge Rd, 5 mi W Fairbans, 27 Jul 1984, birch-spruce forest, FIT, S and J Peck (17, CNCI); Seward, Salmon Creek, 26 Jun 1984, mature poplar-spruce forest, malaise FIT, S and J Peck (9, CNCI); Alaska Hwy 12 mi N Tok, 14 Jul 1985, carrion trap, SA Marshall (1, PECK); Ellito Hwy mi 27.8 White Mtn Trail, 16 Jul 1985, mushroom trap, SA Marshall (1, PECK); 15 mi N Fox 30 mi N Fairbanks, 30 Jul 1984, mixed birch woods, malaise FIT, S and J Peck (1, CNCI); Chena Ridge Rd, 5 mi W Fairbanks, 27 Jul 1984, birch-spruce forest, malaise FIT, S and J Peck (1, CNCI); Bonanza Creek Extension Forest nr Fairbanks, 12 Jul 1989, on Stemonitis sp., SL Stephenson (6, CUIC); Chena River Rec. Area, 30 mi E Fairbanks, 22 Jun 1984, spruce forest, malaise FIT, S and J Peck (16, PECK); Chena Hot Springs, mi 50 Hot Springs Road, 28 Jul 1984, open spruce-moss taiga, malaise FIT, S and J Peck (1, PECK); campus woods, Univ. Alaska, Fairbanks, 30 Jul 1989, on Fuligo septica, SL Stephenson (19, CUIC). Arizona: (1, MCZC). Arkansas: Montgomery Co.: Crystal Rec. Area S campground, 27 Sep 1991, deciduous forest, C Carlton (1, LSAM); Polk Co.: Caney Creek Wild. Area 3.5 mi N Bard Springs, 20 Jan 1992, beech-maple, Berlese, C Carlton (1, LSAM); Caney Creek WA Sugar Creek, 4 Oct 1993, C Carlton (1, LSAM); Scott Co.: Hogan Mt. 18 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Turkey Area, 19 Oct 1991, deciduous forest, C Carlton (2, LSAM). California: Mohawk, Jun 1925, Fenyes (1, MCZC); Sugar Pine, Fenyes (1, CASC); Sta Cruz Mts (5, CASC); colr. (7, MCZC); Mohawk, Jun (1, CASC); Mohawk (3, CASC); Tuolumne Meadows, 7 Jul 1916 (1, MCZC); Amador Co.: 1 mi W Pine Grove, 24 Jun 1975, leaf litter, mixed conifer hardwood forest, A Newton, M Thayer (1, MCZC); Butte Co.: Garland Rd, 2 mi E Hwy 32, 27 Feb 1986, on Polyporus sp. on Quercus kelloggii, F Andrews, A Hardy, T Eichlin, D Mayhew (1, FGAC); 5 mi NE Butte Mdws, Cherry Hill Cpgd, 7 May 1976, FG Andrews (2, FGAC); Fresno Co.: Shaver Lake Heights, 26 Oct 1972, under bark, pine stump, AJ Gilbert (2, FGAC); Nevada Co.: 14.2 mi SW Alleghany, 4 Apr 1985, under Pinus bark, Andrews, Hardy (1, FGAC); Placer Co.: Carnelian Bay, Lake Tahoe, 24 Jun 1964, DF Veirs (1, MCZC); Plumas Co.: 14 Jun 1913 (1, MCZC); Butte Valley Dam, 30 Jun 1975, 4000 9 , under bark conifer, A Newton, M Thayer (2, MCZC); Sierra Co.: Yuba Pass, 20 Jul 1983, 6700 9 , F Andrews, A Hardy (2, FGAC); Tehama Co.: 10 mi SE Mineral, 4 Sep 1961, JF Lawrence (1, MCZC); Trinity Co.: 5 mi SE Peanut, 21 May 1973, under bark, Pinus ponderosa, J Doyen (2, EMEC); Tulare Co.: Sequoia Natl Park, Halstead Creek, 23 May 1984, 7000 9 , R Baranowski (1, LUND). Colorado: Silverton, Jul 1934, Carpenter (1, MCZC); Grand Co.: 21 Aug 1983, 9600 9 , on cream-colored plasmodium, SL Stephenson (2, CUIC); Gunnison Co.: 17 Aug 1983, 10,700 9 , on orangeyellow plasmodium, SL Stephenson (13, CUIC); 17 Aug 1983, 10,700 9 ,onFuligo septica, SL Stephenson (33, CUIC). Indiana: Indianapolis, 14 Jul 1910 (1, MCZC). Montana: Flathead Co.: Swan Lake, 2 Jul 1989, on Stemonitis axifera, SL Stephenson (1, CUIC); 27 Jun 1985, 6700 9 ,onComatricha sp., SL Stephenson (2, CUIC); Lake Co.: 29 Jun 1985, on Stemonitis sp., SL Stephenson (10, CUIC). New Hampshire: Subalpine, Mt Washington, 4 Jul 1896 (3, MCZC); N Conway, Jun (1, MCZC); Mt Washington, 8 Sep 1926, Darlington (1, MCZC); Randolph, Sep 1904, JDS (1, MCZC); Mt Chocoura, 24 Jun 1932 (1, MCZC); Carr. Co.: The Bowl, 2.5 mi NW Wonalancet, 18 Sep 1985, FIT, DS Chandler (6, CNCI); 1 mi N WonalancetEParkSpringBr, 18Sep1985,FIT, DS Chandler (2, CNCI); Coos Co.: Norton Pool, 3miNEEastInletDam,22Aug1986,FIT,DS Chandler (13, CNCI); Straf Co.: Hubbard Brook Exp For., Bear Brook, 6 Jul 1983, rotten wood, DS Chandler (1, DENH). New York: Tompkins Co.: Ithaca, 17 Oct 1976, on Myxomycetes, T Hlavac (1, MCZC). Oregon: 5 mi NE Newberg, 14 Nov 1968, light trap, Anderson and Goeden (1, RLWE); Grants Pass, 2 Oct 1965, K Goeden (2, RLWE); Benton Co.: Mary’s Peak, 17 May 1983, 3500 9 , Ribes litter, DS Chandler (1, DENH); Mary’s Peak, 20 Jun 1970, on polyporous fungus, RL Westcott (1, RLWE); Clackamas Co.: 10 mi N 10 mi E Gout Camp, Mt Hood rd 5–31, 30 Aug 1972, 5500 9 , hemlock litter, Benedict (6, PECK); Clackamas Co.: 1.5 mi S jct US 26 Ore. 35, 11 Jul 1975, litter, mixed conifer forest, Berlese, A Newton, M Thayer (1, MCZC). Utah: Garfield Co.: Aquarius Plateau, Bluebell Knoll, Mar 1986–Sep 1986, pitfall trap, AJ Gilbert (23, FGAC); Wasatch Co.: Lost Lake Camp, Uinta Mountains, 29 Aug 1940, 7800 9 , HP Chandler (1, CASC). Washington: Pierce Co.: Mt Rainier Natl Park 4.7 mi W Longmire, 26 Jul 1975, 2200 9 ,onFuligo septica, A Newton, M Thayer (2, MCZC); Yakima Co.: 8 mi SW Tieton RS, Snoqualmie NF Bear Creek, 11 Jun 1973, WJ Turner (17, WSUC). D ISCUSSION : This species has been described several times based on supposed degree of angulation of the lateral corners of the pronotum, type of dorsal punctation, and presence or absence of a male mandibular horn. We are unable to find any significant differences in degree of angulation between any of the many specimens examined, including the type specimens of A. angulare, A. municeps, and A. assimile. There are some differences in the type of punctation, however. Dorsal punctation of the elytra and, to a lesser extent, the pronotum and head vary from relatively coarse and sparse to very fine and dense. Specimens from the eastern portion of the range (such as from where A. municeps was described) are often much more finely and densely punctate. However, male genitalia and other characters such as the nature of the metasternum and male metasternal fovea agree in all the populations studied throughout the range of the species, and punctation tends to be quite variable within as well as among populations, suggesting that this character is not useful for delimiting species. Therefore, we have synonymized A. angulare,A. municeps, and A. assimile.A. angulare has priority. Interestingly, although Hatch advised Fall that A. temporale was preoccupied (Fall, 1934a), and Hatch knew of Fall’s (1934a) replacement of A. temporale Fall with A. municeps (Hatch, 1936), he still erected a new replacement name, A. falli (Hatch, 1957), although by this time it was unnecessary. 2005 19MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II This species has been collected during every month. The species occurs in a broad variety of forest types and has been collected in a number of trap types including flight intercepts, pitfalls, and ‘‘carrion traps’’. Elevation records are from 2200 ft (Washington) to 10,700 ft (Colorado). Host records from specimen labels include: Brefeldia maxima, Comatricha sp., Fuligo septica,Stemonitis axifera,Stemonitis sp., Polyporus sp., and from ‘‘cream-colored’’ and ‘‘orange-yellow’’ plasmodia. Agathidium hatchi Wheeler Figures 4, 37, 38, 43 Anisotoma fenderi Hatch, 1957: 32. Agathidium hatchi Wheeler, 1977: 137 (new combination, replacement name for Agathidium fenderi (Hatch, 1957, Anisotoma), preoccupied by Agathidium fenderi Hatch, 1957). T YPE M ATERIAL : Holotype, / in OSAC, examined by the second author (Wheeler, 1977). T YPE L OCALITY : United States, Oregon, Ocean Park. D IAGNOSIS : This species differs from others by its very large size (TBL . 3.5 mm), clavate antennae (antennomeres 5–11 gradually widening) (fig. 4), unmodified male metafemur (without teeth) and very distinctive median lobe which is long, slender, irregularly sinuate, and has the apex bifid in ventral aspect (figs. 37, 38). D ESCRIPTION : Body very large (TBL 5 3.77–4.14 mm), broad (PNW/TBL 5 0.53– 0.55), laterally broadly rounded, somewhat contractile. Head, pronotum, and elytra red-brown; venter red-brown, darker on metasternum; antennae brown, club dark brown; palpi brown; legs yellow-brown. Head broad (fig. 4) (MDL/OHW 5 0.66– 0.67), flattened, without transverse, medial crease; temporum very prominent, about length of eye when viewed in dorsal aspect, extending laterad beyond level of eye at posterior end (fig. 4); head finely and densely punctate, surface between punctures with conspicuous microreticulation of small cells; eyes large, rounded, finely faceted; frontoclypeal suture prominent through entire length; clypeal margin relatively straight, extending to level of anterolateral margins of frons (fig. 4); anterior margin of labrum entire; antenna clavate, club not abrupt (fig. 4), antennomere ratios: length I:II:III 5 1.8:1.0:1.6, width VII:VIII:IX 5 1.0:1.0:1.2. Pronotum broad (PNL/PNW 5 0.43–0.48), laterally not strongly produced, anterolateral angle moderately rounded, posterolateral angle more angulate than anterolateral angle; punctation and surface microsculpture similar to that of head. Elytra broad, robust, laterally broadly rounded (SEL/ELW 5 1.08–1.19); coarsely and densely punctate, punctures not in series, surface between punctures shiny and smooth; sutural stria about two-thirds length of elytron. Mesosternum slightly concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion about one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum broad medially (MTL/ MTW 5 0.30–0.31), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres moderately expanded and with large ventral field of spatulate setae; left mandible unmodified; metafemur slender, unmodified; metasternal fovea moderately large, transverse, linear, anterad of middle, with prominent series of fine, dense setae. Median lobe in lateral aspect long, slender, irregularly sinuate, with distinct prominence on dorsal surface at base of apical portion which is robust, slightly curved and apically pointed (fig. 38); in ventral aspect very slender, lateral margins subparallel, apically prominently divided into two robust, apically pointed rami, apical orifice elongate (fig. 37); operculum absent (fig. 37); lateral lobes broad, broadly fused along dorsal and ventral margins forming a sheath around median lobe, apical portions broad, tapered to rounded apices, ending far short of apex of median lobe, without apical setae (figs. 37, 38). Female tarsi 5-4-4. D ISTRIBUTION : This species is known from Oregon and New Brunswick (fig. 43). S PECIMENS E XAMINED : CANADA: New Brunswick: Fredrickton, 20 Mar 1961, RC Clark (1, Utah State). UNITED STATES: Oregon: 5 MI NE Newberg, 2 Nov 1968, black light trap, Anderson and Goeden (1, Utah State). 20 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 43. Geographic distribution of Agathidium concinnum-group species: A. concinnum 5v ;A. hatchi 5m . 2005 21MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II D ISCUSSION : This species was first described in Anisotoma but was transferred to Agathidium by Wheeler (1977). Hatch (1957) had already described a species of Agathidium named fenderi, and thus Wheeler replaced the transferred homonymous name with A. hatchi. Superficially the species does resemble many members of Anisotoma in being exceptionally large, broad, and dorsoventrally flattened. The unusual male genitalia are also reminiscent of some members of Anisotoma and are quite different from most other North American Agathidium. However, the prolonged temporum and supraocular carina (fig. 4), anteriorly shortened clypeus (fig. 4), and lack of serial striae on the elytra together suggest the species best belongs in Agathidium following Wheeler (1977). The antennae are unusual in that antennomeres 5– 8 are gradually broadened such that the antenna does not have an abrupt 3-segmented club (fig. 4). In other respects, the species seems to resemble A. concinnum in general shape, relatively shallow concavity of the posterior portion of the mesosternum, and prolonged and protruding temporum. The distribution of this species is unusual in that specimens are known from Oregon and New Brunswick, but from nowhere in between. The Oregon specimens (including the type) are females and the New Brunswick specimen is a male. Therefore, there is some possibility the specimens represent different species and that our male-female association is incorrect. However, since the specimens agree closely in all characters examined and numerous other species have transcontinental distributions, we regard conspecificity of the specimens to be a reasonable hypothesis until additional material can be discovered. It is also possible that the New Brunswick specimen is mislabeled. The species has been collected in March and November. One specimen was collected in a black light trap. Agathidium concinnum Mannerheim Figures 2, 5, 39, 40, 43, 118 Agathidium concinnum Mannerheim, 1852: 370; Horn, 1880; Leng, 1020; Fall, 1934b; Hatch, 1957. Agathidium castaneum: Attributed to ‘‘Me´ne´trie´s in litteris’’ by Mannerheim (1852). (Published in synonymy with A. concinnum and never made available by subsequent use as a valid name.) NOMEN NUDUM . Agathidium effluens Mannerheim, 1853: 202; Horn, 1880 (synonymized with A. concinnum); Leng, 1920. T YPE M ATERIAL :Agathidium concinnum: lectotype (designated here to clarify assignment of this name to this species), / in MCZC labeled ‘‘46/Type 7944 [number handwritten, red label]/A. concinnum Sitkha Mannh. [handwritten]’’. Mannerheim had multiple individuals of this species, but was not clear how many. We have selected the female in MCZC labeled as the type as the lectotype. Agathidium effluens: lectotype (designated here to clarify assignment of this name to this species), / in MCZC labeled ‘‘Kenai [handwritten]/A.effluens RA. Mann [handwritten]/concinnum8 [handwritten]’’. Mannerheim was not clear whether multiple specimens were involved during this description, so we have selected a single specimen from the MCZC as the lectotype. T YPE L OCALITY :Agathidium concinnum: United States, Alaska. Agathidium effluens: United States, Alaska, Kenai Peninsula. D IAGNOSIS : This species is characterized by the laterally protruding temporum in a bulge beyond the lateral margin of the eye (figs. 2, 5), the broad, depressed body form, the metafemur with a small, medial tooth on the posterior margin in most male specimens (fig. 118), and the median lobe of the aedeagus in lateral aspect very robust, with broad basal portion and subbasal constriction, apically tapered to a sharp, curved point (fig. 40) and in ventral aspect broad, with the apical portion broadly triangular and sharply pointed (fig. 39). The apical orifice is large with prominent, truncate rami visible (fig. 39). D ESCRIPTION : Body large (TBL 5 2.51– 3.30 mm), broad (PNW/TBL 5 0.48–0.49), laterally broadly rounded, moderately contractile. Head pronotum and elytron light brown to dark red-brown; venter light brown, darker on metasternum; antennae brown, club dark brown; palpi and legs brown. Head broad (figs. 2, 5) (MDL/OHW 5 0.60–0.67), flattened, without transverse, medial crease; temporum very prominent, 22 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY about one-half length of eye when viewed in dorsal aspect, extending prominently lateral to eye (figs. 2, 5); head finely and moderately densely punctate, smooth and shiny on surface between punctures; eyes large, rounded, finely faceted; frontoclypeal suture obsolete medially; clypeal margin curved, extending slightly beyond anterolateral margins of frons (figs. 2, 5); anterior margin of labrum entire; antennomere ratios: length I:II:III 5 2.4:1.0:2.1, width VII:VIII:IX 5 1.0:1.0:2.0. Pronotum broad (PNL/PNW 5 0.49–0.50), not strongly produced laterally, anterolateral angle subquadrate, posterolateral angle moderately angulate; punctation and surface microsculpture similar to those of head. Elytra broad, laterally broadly rounded (SEL/ELW 5 1.02–1.11); coarsely and densely punctate, punctures not in series, surface between punctures shiny and smooth; sutural stria about one-half length of elytron on most specimens. Mesosternum moderately and broadly concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion about one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum relatively broad medially (MTL/ MTW 5 0.24–0.29), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres slightly expanded and with small ventral field of spatulate setae; left mandible unmodified; metafemur slender, many specimens with distinct, medial, small tooth (fig. 118); metasternal fovea minute, medial with small brush of fine, dense setae. Median lobe in lateral aspect very robust, very broad basally with prominent subbasal constriction, medially curved and expanded, apically tapered to sharply pointed, slightly dorsally directed apex (fig. 40); in ventral aspect broad, apical portion very broadly triangular, tapered to slightly acuminate, pointed apex, orifice extremely large (fig. 39); operculum comprised of two long, broad, flat structures, each ramus apically rounded to subtruncate (fig. 39); lateral lobes moderately broad basally, more slender submedially, apically expanded and directed ventrad, apices slightly constricted and pointed with 2 prominent setae (figs. 39, 40). Female tarsi 5-4-4. D ISTRIBUTION : This species is widespread in western North America from Alaska south to Arizona and east to Colorado and New Mexico (fig. 43). S PECIMENS E XAMINED : ‘‘W.T.’’ [probably Washington Territory] (1, MCZC). CANADA: British Columbia: NEAR Mabel Lake at Squaw Valley, 11 Aug 1982, R Baranowski (2, LUND); Mesachie, 30 Jun 1980, H and A Howden (8, CNCI); Stanley, 21 Jun 1932, W Mathers (1, CASC); Pender harbor, 20 Jun 1929, RT Turner (1, CASC); Lorna, 29 Jul 1926, H Richmond (4, CASC); Merrit, 22 Aug 1925, Pinus ponderosa, WM Mathers (1, MCZC); Lorna, 28 Jul 1926, Picea engelmanni, H Richmond (1, MCZC); Lorna, 20 Jul 1924, R Hopping (2, MCZC); Lorna, 23 Jul 1924, R Hopping (2, MCZC); Lorna, 28 Jul 1926, Picea engelmanni, B Hopping (1, CASC); Monashee Mtn Near Cherryville, 11 Aug 1982, 1600 m, M Sorensson (4, LUND); Stanley (1, CASC); Terrace, ME Hippisley (1, MCZC); Terrace (1, MCZC); Terrace, Feb 1960, ME Clark (1, MCZC); Terrace, ME Clark (1, MCZC); near Mabel Lake at Squaw Valley, 5 Aug 1982, R Baranowski (2, LUND); Monashee Mtn Near Cherryville, 10 Aug 1982, 1400–1600 m, M Sorenson (3, LUND); Manning Park, E gate, 24 Jul 1984, lodgepole pine, D Miller (1, CNCI); Manning Prov. Park, 27 Nov 1992, 1220 m, spruce or fir log, red or brown rotten wood, R Baranowski (1, LUND). UNITED STATES: Alaska: E Fork Toklat River, McKinley Park, 17 Aug 1958, Lindroth (1, MCZC). Arizona: state only (5, MCZC); White Mts, 4 Jul 1933 (2, CASC); Flagstaff, San Francisco Mts, 13 Aug 1934, 8000 9 , D Rockefeller (2, AMNH). California: state only (14, MCZC); Miami, Jun (1, MCZC); Sta Cruz Mts (1, CASC); Tehachapi, Wickham (1, MCZC); colr. (2, MCZC); Stanford Univ, 4 Apr 1955, PS Bartholomew (2, CASC); Tahoe Canyon, 27 Jun, A Fenyes (2, MCZC); Miami, Jun (1, CASC); Pomona, 30 May 1896 (1, MCZC); Yosemite, 20 May 1934, Bryant (1, CASC); McClaw, A Fenyes (1, CMNH); Los Gatos (1, MCZC); Tehachapi, Wickham (1, MCZC); North Fork, 31 May 1920, Dietrich (2, MCZC); Tahoe Canyon, Jun (1, CASC); Manzanita Lake, Lassen Natl Park, 23 May 1941, CD Michener (1, EMEC); Wawona, Jun (1, CASC); Sugar Pine, 6 Jul (1, CASC); Miami, Jun (1, CASC); Alpine Co.: 4 mi W Markleville, 30 Sep 1970, grass duff, Berlese, FG Andrews (1, FGAC); Butte Co.: 5 mi NE Butte Mdw Cherry Hill Cpgd, 7 May 1976, caught flying, FG Andrews (2, FGAC); El Dorado Co.: 0.7 mi E Pacific House, 28 Apr 1992, screening flume, F Andrews, T Eichlin (1, FGAC); 7 mi S Yellow- 2005 23MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II jacket Camp, 28 Jun 1980, CY Kitayama (4, EMEC); China Flat CG, 2 mi S Kyburz, 21 Jun 1979, 4300 9 , W and AR Hardy (1, FGAC); 5 mi S Meyers, 29 Jun 1975, 7200 9 , mixed conifer forest, litter, Berlese, A. Newton, M. Thayer (1, MCZC); Fresno Co.: 5 mi SW Big Creek, 9 Jul 1962, JF Lawrence (2, MCZC); Kern Co.: 2 mi W Woffard Hts, 29 Apr 1964, Doyen (1, EMEC); Mariposa Co.: Fish Camp, 6 mi NW, 8 Jul 1962, JF Lawrence (1, EMEC); 2 mi NE Darrah, 17 Nov 1975, oak litter, Berlese, AJ Gilbert (1, FGAC); Mono Co.: Convict Lake (near Mammoth Lakes), 13 Jun 1984, 8000 9 , R Baranowski (1, LUND); Inyo Hat For. HM Hall Nat Area, 28 Aug 1973, 10,000–10,400 9 ,Pinus murrayana forest, pitfall trap, R Papp (3, EMEC); Nevada Co.: Sagehen Creek, 28 Aug 1983, debris, log in stream, F Andrews, A Hardy (1, FGAC); Placer Co.: Carnelian Bay, Lake Tahoe, 24 Jun 1964, DF Veirs (1, EMEC); Plumas Co.: Lake Almanor, 23 Apr 1971, pine litter, FG Andrews (1, FGAC); San Bernardino Co.: Hardin Flats, 13 Jun 1964, R Scott (1, FGAC); Santa Clara Co.: colr. (1, MCZC); Santa Cruz Co.: county only (1, MCZC); Boulder Creek, 7 May 1968, A and A Gillogly (1, FGAC); Shasta Co.: Burney Falls, 18 Jun 1954, RO Schuster (2, FGAC); Sierra Co.: Yuba Pass, 8 Jul 1976, moss along small stream, FG Andrews (1, FGAC); 10 mi E Sierra City, N Yuba River, 28 Jun 1976, 6200 9 , L and N Herman (1, AMNH); Siskiyou Co.: county only (1, CASC); 3 mi E McCloud, Elk Creek, 30 Jun 1976, 3200 9 ,L and N Herman (1, AMNH); 5.4 mi SE Seiad Valley, O’Neil Creek, 3 Jul 1976, 1500–2000 9 , L and N Herman (1, AMNH); Tehama Co.: Snoqualmie Gulch 10 mi SE Manton, 1 Dec 1986, 2850 9 , leaf litter, DS Chandler (1, CNCI); Toulumne Co.: 10 mi E Buck Meadows, 16 Jun 1984, 3500 9 , R Baranowski (1, LUND); Tulare Co.: Sequoia Natl Park., 10 May 1979, R Baranowski (1, LUND); Sequoia Natl Park, Halstead Creek, 23 May 1984, R Baranowski (13, LUND); Ash Mountain, 17 Mar 1984, flume forebay, R Haines (1, FGAC); Tuolumne Co.: 30.5 NE Strawberry near Deadman Creek, 18 Jul 1976, 8000 9 , L and N Herman (3, AMNH). Colorado: Breckenridge, 15 Jul 1896, 9600–10,000 9 , Wickham (1, MCZC); above Ouray, mineral point trail, 1 Jul 1889, 9000– 10,000 9 , Wickham (1, MCZC); Veta Pass, Jun 1924 (1, MCZC); Grand Co.: SE Rabbit Ears Pass, 21 Jun 1975, 9000 9 , squirrel middens under conifers, A. Newton, M. Thayer (2, MCZC). Idaho: Boise Co.: N Centerville, 14 Sep 1974, AD Allen (1, PECK); Lemhi Co.: continental divide 2.4 mi airline SW of Bannock Pass, 24 Aug 1962, 8500 9 , HB Leech (2, CASC); continental divide 2.4 mi airline SW Bannock pass, 24 Aug 1969, HB Leech (16, CASC). Montana: Gallatin Co.: Earthquake Lake, 28 Jun 1966, J Haddock (1, EMEC); Madison Co.: Hidden Lake Bench, Oct– May 1989, 7400 9 , FIT, DL Gustafson (1, MTEC). Nevada: state only (13, MCZC); White Pine Co.: Snake Range, 6 mi W Baker, 1984–1985, pitfall trap, D Giuliani (3, FGAC). New Mexico: San Miguel Co.: Gallinas, 6 Aug 1992, JF Cornell (2, CUIC). Oregon: state only (3, MCZC); Albany (1, MCZC); Portland, 22 Jun (3, OSUC); Corvallis, 12 Jun 1925, EP VanDyke (1, CASC); Clarks, 10 Jan 1961, H Foster (1, RLWE); nr Plains, 1 Apr 1956, fir slab wood, K Goeden (5, RLWE); Benton Co.: nr Corvallis, Mary’s Pass, May 1966, rotary net trap (3, CUIC); Corvallis, 23 Jun 1974, from nest of Neotoma fuscipes, D Carslon (2, FGAC); Clakamas Co.: Salmon River nr Zigzag, 27 Aug 1982, R Baranowski (3, LUND); Harney Co.: Steens Mt N side Loop Rd, 12 May 1972, aspen-juniper duff, E Benedict (1, CNCI); Steens Mt, Fish Lake Rd, 19 Jul 1971, 6000 9 , juniperaspen duff, HE Benedict (1, PECK); 15 mi E Frenchglen Lily Lake, 29 Jul 1971, aspen duff, E Benedict (6, CNCI); Klamath Co.: Kimball St. Park. Wood River Spring, 2 mi N Fort Klamath, 20 Jun 1978, J Schuh, L and N Herman (1, QDWC); PoLake Co.: 0.5 mi S Buell, 9 Jan 1981, Neotoma nest 15 9 off ground, PJ Johnson (1, FMNH); 1 mi N Suver Jct, old Hwy 99-W, 23 Mar 1980, fecal material, nest of Neotoma sp., PJ Johnson, J LaBonte (2, FMNH); Buell, 27 Sep 1981, fecal chamber of Neotoma fuscipes,PJ Johnson (2, FMNH); 3 mi S Airlie, 23 Mar 1980, Neotoma nest, PJ Johnson (2, FMNH); Yamhill Co.: Muddy Valley, 2 Feb 1982, Neotoma nest, fence post, PJ Johnson (1, FMNH). Utah: Uintah Mts, Tryol Lake, 26 Aug 1920, 9800 9 , HP Chandler (1, CASC); Alta (2, MCZC); Wasatch Co.: Lost Lake Camp, Uinta Mountains, 29 Aug 1940, 7800 9 , HP Chandler (2, CASC). Washington: Easton (1, CASC); Baring, Jul (1, CASC); Olympia (1, MCZC); King Co.: 14 km S North Bend, 4 Mar 1981, moss on rocks, RE Nelson (1, CNCI); Skagit Co.: 2 mi NW Rainy Pass, 31 Jul 1982, M Sorensson (2, LUND); 2 mi NW Rainy Pass, 31 Jul 1982, R Baranowski (1, LUND); Birdsview, 27 Jul 1982, R Baranowski (1, LUND). D ISCUSSION : Based on our examination of the type specimens of A. concinnum and A. effluens, these two names refer to the same species following Horn (1880). This species has been collected from every month during the year. Specimens have been collected in a variety of habitats including pine, juniper, and aspen forests and various sources of litter. The most intriguing habitat, 24 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY which has yielded numerous specimens, is the nests of Neotoma sp. and Neotoma fuscipes. Elevation records are from 2850 ft (California) to 10,400 ft (California). AGATHIDIUM PULCHRUM SPECIES GROUP D IAGNOSIS : Members of this species group are characterized by having the mesosternum strongly concave (declivitous), the clypeus strongly excavate to extending moderately beyond the sides of front of cranium (e.g., figs. 6, 22), the metasternum not strongly shortened, the humeral angles of the elytra relatively distinct, the postocular temporum relatively short (e.g., figs. 6, 22), and males of several species with a prominent tusk or horn on the left mandible (e.g., figs. 21–29, 44, 45). Several of the species are distinctly and dramatically maculate dorsally (figs. 44– 46, 48–51), which is unusual for New World members of Agathidium. The tarsal formula is ? 5-5-4 / / 5-4-4 or 4-4-4. D ESCRIPTION : This group includes species belonging to the subgenus A. (Neoceble) Gozis auctorum, a very diverse Holarctic group. The presence of a horn on the male left mandible (also occurring in our A. brevisternum and some of our A. concinnum group species) is a particularly unique feature of several of these species in the Leiodidae and in beetles in general. Asymmetry in similar secondary sexual features is only rarely reported. The horn comes in a variety of shapes and sizes when fully developed, including those that are short, spinous, and adpressed (figs. 29, 30) to very long and variously curved (figs. 21–23, 44, 45). One species, A. aristerium, has the apex of the mandible strongly upturned and truncate apically (fig. 25, 26). In several species the mandibular horn bears a prominent brush of setae arising apically from an oblique, curved impressed line, but not all species have a setose horn. In all species where sufficient material is available for examination, the horn is variable from entirely absent to well developed. In some species, very many males are strongly modified, whereas in some very few males are modified. The development of this dimorphism is particularly dramatic in one species, A. marae, in which not only is there a left mandibular horn, but the right surface of the frons is also developed into an opposing spine (figs. 23, 24). No evidence has yet been presented for a function for the mandibular horn, but other beetles with horns use them for male-male competition, a possible scenario for these species where large numbers of males and females are often concentrated on a small food and oviposition resource where competition for mates might figure strongly. Historically, many species were recognized based on the presence or absence of a male mandibular horn, but the variability of this feature within all species that possess it makes this an unreliable character for keying or diagnosing species. In the past, species identifications have also relied heavily on the number of female protarsomeres (5 or 4). Although this is a good diagnostic feature of some species, we have avoided using this character as an exclusive diagnostic character since it is not always possible to positively associate males with females. It seems apparent from previous work on the group that often female and males were misassociated, leading to erroneous conclusions about the tarsal formula for females of certain species. Most of these species occur in forests of western and northern North America. There is an additional assemblage of species in northeastern North America and a few that occur only in the central portion of the continent. KEY TO A. PULCHRUM SPECIES GROUP 1. Elytra maculate, either irregularly brown and yellow or red with a broad, black triangle at bases of elytra (figs. 48–51); female with five protarsomeres .................. 2 — Elytra concolorous brown to piceous; female with four or five protarsomeres ....... 5 2(1). Elytra red with a broad, black triangle medially at base of elytra and black around elytral margins (fig. 48); mesosternum with distinctive lobe extending ventrad between mesocoxae; lateral lobes apically prominently expanded (fig. 53); median lobe of aedeagus without medial excavation on ventral surface visible in lateral aspect (fig. 53) ................... .............. A. difforme (LeConte) — Elytra yellow with irregular black maculae (figs. 49–51); mesosternum without dis- 2005 25MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Fig. 44. Agathidium pulchrum, dorsal habitus. tinctive posteroventral lobe; lateral lobes apically not prominently expanded; median lobe of aedeagus with distinctive medial excavation on ventral surface visible in lateral aspect .............. 3 3(2). Elytron without sutural stria (fig. 49); anterior clypeal margin near level of anterolateral margins of frons (fig. 49); male left mandibular horn, when fully formed, relatively straight and flattened; operculum apically narrowed to two spinous apices (fig. 54) ........A. maculosum Brown — Elytron with distinct sutural stria; anterior clypeal margin strongly excavate, not extending to level of anterolateral margins of frons (figs. 22, 51); male left mandibular horn, when fully formed, relatively curved and rounded (e.g., figs. 21, 22); operculum reduced to inconspicuous, flattened hyaline lobe (figs. 56, 58) ..... 4 4(3). Elytral maculation usually more extensive, covering much of elytra (fig. 51); median lobe of aedeagus in lateral aspect with apical portion slender, apically narrowly rounded or pointed (fig. 57), in ventral aspect with apical portion broader, lateral 32 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 73); northern California north to Alaska (fig. 91) .................... ........A. rotundulum Mannerheim — Size larger (TBL . 2.2 mm) ; anterior clypeal margin distinctly excavated, though in some cases not strongly so; antennomere II distinctly longer than antennomere III, if II approaching length of III, then TBL clearly greater than 2.2 mm and anteriorly clypeal margin prominently excavated; male with metasternal fovea; male left mandibular horn present or absent; female tarsi 4-4-4 or 5-4-4; median lobe of aedeagus variable; distribution variable ............................ 12 12(11). Male metasternal fovea located at center of metasternum ............... 13 — Male metasternal fovea located distinctly anterad of center of metasternum .... ............................ 14 13(12). Male mandibular horn, when present, formed by extension of apex of mandible in long, evenly curved, flattened structure (figs. 25, 26); median lobe of aedeagus in lateral aspect slender, with small basal portion, evenly and shallowly curved (fig. 75), in ventral aspect apically expanded and very broadly pointed (fig. 74) ................ ............ A. aristerium Wheeler — Male mandibular horn, when present, straight, extending from middle of mandible in oblique angle to right over right mandible (figs. 27, 28); median lobe of aedeagus in lateral aspect robust, with large basal portion, strongly constricted and curved distad of expanded basal portion (fig. 77), in ventral aspect apically not or only slightly expanded and more strongly pointed (fig. 76) ......... A. atronitens Fall 14(12). Male lateral lobe apically abruptly expanded and obliquely truncate (fig. 79); median lobe of aedeagus in lateral aspect robust, apex slender, straight and sharply pointed (fig. 79) .......... ........A. oregonense, new species —Male lateral lobe not expanded apically, of even width to near apex; median lobe of aedeagus in lateral aspect variable, but without long, slender, straight apex ........................ 15 15(14). Median lobe of aedeagus in lateral aspect slender, strongly bent medially, nearly in right angle (fig. 81); male mandibular horn, when present represented by a short, flattened spine extending posterad over clypeus (figs. 29, 30) ..... .................. A. picipes Fall — Median lobe of aedeagus in lateral aspect moderately curved, but not strongly bent medially; male mandibular horn, when present, represented by a long, rounded, setose horn ........... 16 16(15). Median lobe of aedeagus in lateral aspect apically with small, abrupt hook (fig. 83) ...... A. hamulum, new species — Median lobe of aedeagus in lateral aspect apically without hook (fig. 85) .... .............. A. politum LeConte Agathidium difforme (LeConte) Figures 48, 52, 53, 86 Phalacrus difformis LeConte, 1850: 222. Agathidium difformis, LeConte, 1853: 286; Horn, 1880; Fall, 1934b. Agathidium difforme, Leng, 1920. Agathidium canadensis Brown, 1930: 89; Fall, 1934b (synonymized with A. difforme). T YPE M ATERIAL :Agathidium difforme: lectotype (designated here to clarify assignment of this name to this species), ? in MCZC labeled ‘‘[light green circular disk with two sides cut away]/Type 3175 [number handwritten, 2/3 of label red]/A. difforme Lec [handwritten]’’. Since LeConte did not indicate how many specimens he had at the time of description, we have selected a male specimen in the MCZC to be the lectotype. Agathidium canadense: holotype ? ,in CNCI labeled ‘‘Brittania Ont. VI-8-1928 W.J.Brown/ ? /HOLOTYPE Agathidium canadensis 2878 No. Brown [red label]’’. There are two female paratypes with the same collection information as the holotype. T YPE L OCALITY :Agathidium difforme: North side of Lake Superior (LeConte, 1853). Agathidium canadense: Canada, Ontario, Britannia. D IAGNOSIS : This species is diagnosed by the unique dorsal color pattern. The elytra are red with a large, black basal triangle (fig. 48). The anterior clypeal margin is strongly excavate (fig. 48). The male mandibular horn, when fully formed, is long, stout, and curved, extending from the medial surface of the left mandible up and over the clypeus. The median lobe of the aedeagus is slender, relatively straight, and apically pointed (fig. 53). The lateral lobes are apically broadly ex- 2005 33MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Fig. 86. Geographic distribution of Agathidium difforme. 34 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 87. Geographic distribution of Agathidium maculosum. panded and broadly rounded (fig. 53). Females have 5-4-4 tarsi. D ESCRIPTION : Body moderately large (TBL 5 2.26–3.88 mm), somewhat elongate (PNW/TBL 5 0.46–0.47), moderately contractile. Head piceous with medial red macula, clypeus red; pronotum yellow laterally with large region of red to piceous medially; elytron red medially with black broadly around lateral margins and with large, black triangular macula anteriorly along suture; venter piceous; antennae dark red, club dark brown; palpi and legs dark red. Head broad (fig. 48) (MDL/OHW 5 0.52– 0.53), flattened, often with moderately prominent transverse, medial crease; temporum short, inconspicuous; head finely and sparsely punctate, smooth and shiny on surface between punctures; eyes large, rounded, finely faceted; frontoclypeal suture moderately well defined medially; clypeal margin strongly excavate (fig. 48); anterior margin of labrum entire; antennomere ratios: length I:II:III 5 2.3:1.0:1.5, width VII:VIII:IX 5 0.9:1.0:1.7. Pronotum broad (PNL/PNW 5 0.55), laterally not strongly produced, anterolateral angle subquadrate, posterolateral angle relatively strongly angulate; punctation and surface microsculpture slightly coarser than that of head. Elytra somewhat elongate (SEL/ELW 5 1.08–1.11); punctation slightly coarser and more dense than on pronotum, some punctures in indistinct series, surface between punctures shiny and smooth; sutural stria about three-fifths length of elytron on most specimens. Mesosternum strongly concave posteriorly, with prominent lobe extending ventrally between mesocoxae, anterior portion about one-half length of posterior portion; medial longitudinal carina reduced on anterior portion. Metasternum relatively broad medially (MTL/MTW 5 0.33–0.35), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres slightly expanded and with small ventral field of spatulate setae; mandibular horn, when fully developed, long, curving posteriorly over front of head, subcircular throughout in cross-section, punctate and setose along its length, apical fovea prominent with large cluster of setae; metafemur moderately slender, unmodified; metasternal fovea small, anterior, with small brush of fine, dense setae. Median lobe in lateral aspect moderately slender, evenly curved, apical portion slender, gently curved dorsad, apex with small triangular, dorsal expansion (fig. 53); in ventral aspect moderately slender, lateral margins subparallel for most of length, slightly sinuate at base of apical portion which is subtriangular, and apically pointed (fig. 52); operculum broad, flat, apically divided with each ramus pointed and convergent (fig. 52); lateral lobes slender, evenly curved, apically expanded, prominently flattened, rounded, with two long, subapical setae (fig. 53). Female tarsi 5-4-4. D ISTRIBUTION : This species is known from Alaska south to California and Nevada and across Canada to New Hampshire (fig. 86). S PECIMENS E XAMINED : CANADA: Alberta: Jumpinground Creek, 6 Sep 1964, fungus under rotten log, BF and JL Carr (2, CARR); George Lake 50 km NW Edmonton, 11 Jun 1984, spruceaspen forest, FIT, S and J Peck (1, PECK); Ghost Dam, 12 Jun 1980, BF and JL Carr (4, CARR). 2005 35MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Fig. 88. Geographic distribution of Agathidium pulchrum. 36 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 89. Geographic distribution of Agathidium pulchrum-group species: A. athabascanum 5m ; columbianum 5. ;A. laetum 5v ;A. rusticum 5m . 2005 37MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Fig. 90. Geographic distribution of Agathidium pulchrum-group species: A. amae 5m ;A. repentinum 5v . 38 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 91. Geographic distribution of Agathidium pulchrum-group species: A. aristerium 5m ;A. marae 5. ;A. rotundulum 5v . 2005 39MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II British Columbia: Mt Robson P Park, Berg Lake Trailhead, 11 Jul 1984, FIT, RS Anderson (7, CNCI); Alaska Hwy, 37 km W Ft Nelson, 12 Jun 1984, aspen-spruce forest, FIT, S and J Peck (3, PECK); Mt Robson P Park, Berg Lake Trailhead, 11 Jul 1984, FIT, RS Anderson (2, PECK); Lorna, 20 Jul 1924, R Hopping (1, CASC). Ontario: Smooth Rock Falls, 56 km NW Cochrane, 3 Jun 1984, mixed spruce-birch, evening car netting, S and J Peck (2, CNCI); 25 km W Ignace 75 km E Dryden, 5 Jun 1984, fir-maple forest, FIT, S and J Peck (1, PECK); Smooth Rock Falls 56 km NW Cochrane, 3 Jun 1984, mixed spruce birch, evening car netting, S and J Peck (3, PECK). Quebec: Duparquet, 18 Jun 1939, lake shore, GS Smith (1, CASC); Duparquet, 9 Jun 1939, GS Smith (1, CASC); Duparquet, 3 Jun 1948, GS Smith (1, CASC). Saskatchewan: Jay Jay Lake, Rt 102, 130 km NE Candle Lake, 8 Jun 1984, pine-fir forest, FIT, S and J Peck (2, CNCI); Turtle Lake Jct, 60 km S Meadow Lake, spruce-moss forest, FIT, S and J Peck (1, PECK). Yukon: Simpson Lake, 81 Km N Watson Lake, 15 Jun 1984, pine-willow, FIT, S and J Peck (1, PECK). UNITED STATES: Alaska: Chena River Rec. Area, 30 mi E Fairbanks, 22 Jun 1984, spruce forest, malaise FIT, S and J Peck (4, PECK); Kenai Pen, Cooper Landing, Rt 1, 25 Jun 1984, poplar spruce forest, malaise FIT, S and J Peck (1, PECK); Kenai Pen, E Skilak Rd Jct Rt 1, 25 Jun 1984, aspen-spruce forest, malaise FIT, S and J Peck (15, PECK); Chena Ridge Rd, 5 mi W Fairbanks, 22 Jun 1984, poplar forest, malaise FIT, S and J Peck (5, PECK); Bonanza Creek Exp. Forest nr Fairbanks, 12 Jul 1989, on Stemonitis,SL Stephenson (1, CUIC); Houston, 24 Jun 1984, spruce-birch forest with moss, malaise FIT, S and J Peck (3, PECK); Seward, Salmon Creek, 27 Jun 1984, rotted log pile, S and J Peck (1, PECK); 35 mi N Willow Trapper Creek, mi 106, 23 Jun 1984, spruce-birch forest, malaise FIT, S and J Peck (2, PECK). California: Sequoia Natl Park, 11 May 1979, R Baranowski (2, LUND); El Dorado Co.: 0.5 mi N Stumpy Meadows Lake, 20 Apr 1989, 4200 9 , F Andrews, Te Eichlin (1, FGAC); 7.7 mi SE Virner Blodgett Forest, 29 Mar 1984, under bark of Pinus ponderosa, FG Andrews (2, FGAC); Blodgett Forest 13 mi E Georgetown, 14 Jun 1967, PF Warner (1, WSUC); Lassen Co.: Ine Creek, 2 May 1950, HP Chandler (1, CASC); Pine Creek, 2 May 1950, 5200 9 , HP Chandler (1, CASC); Pine Creek, 2 May 1950, 5000 9 ,HP Chandler (2, CASC); Mono Co.: Convict Creek, 24 May 1963 (1, FGAC); Nevada Co.: Sagehen Creek, 8 Jul 1970, CS Glaser (2, CASC); Tulare Co.: Sequoia Natl Park, Halstead Creek, 23 May 1984, 7000 9 , R Baranowski (1, LUND); Tuolumne Co.: Tiog Rd 9 mi E Crane Flat, 18 May 1976, squirrel middens and litter, fir forest, Berlese, A Newton, M Thayer (1, FMNH). Montana: Gallatin Co.: Bozeman Creek, 8 Jun 1987, 5300 9 ,DL Gustafson (1, MTEC); Judith Basin Co.: Little Belt Mts, 5 mi S Deadhorse Ck, 27 May 1988, 6600 9 , FIT, CE Seibert (1, MTEC). Nevada: Washoe Co.: 1.5 mi W top of pass, Mt Rose, 27 Aug 1969, 8550 9 , HB Leech (2, CASC). New Hampshire: White Mts (1, MCZC); Coos Co.: Norton Pool, 3 mi NE East Inlet Dam, 25 Jun 1986, FIT, DS Chandler (2, CNCI); Jefferson Notch, 20 Nov 1974, on ‘‘red tubular slime mold’’, A Newton (1, MCZC); 1 mi NE East Inlet Dam, 27 May 1986, FIT, DS Chandler (1, CNCI); 0.3 mi S Jefferson Notch, 23 Jul 1980, 895 m, Picea-Abies for., window trap, A Newton, M Thayer (2, MCZC); 1.0 mi S Jefferson Notch, 7 Nov 1975, 2700 9 , litter fir-spruce-birch forest, Berlese, A Newton, M Thayer (1, MCZC); Mt Washington auto road, 1 Jul 1982, 3000 9 , rotten birch, DS Chandler (1, DENH). New York: Adiron. Mt., Mt Seward [Adirondacks?], 22 Jun 1945 (1, MCZC); Mt. Whiteface, 25 Jul 1936 (1, CASC). Oregon: Clakamas Co.: 1.5 mi S Jct US 26 Oregon 35, 11 Jul 1975, mixed conifer forest litter, A Newton, M Thayer (3, FMNH); Klamath Co.: 2 mi S 2 mi W Lake of the Woods, 14 Nov 1972, 5700 9 , rotten wood, E Benedict (6, CNCI). D ISCUSSION : Based on our examination of the holotype and two paratypes of A. canadense, we follow Fall’s (1934b) synonymy of that species with A. difforme. This species has been collected throughout the year except during the winter months. It has been collected from flight intercept traps and sifting in a variety of forest types including fir, pine, spruce, birch, maple, and poplar. Elevation records are from 2700 to 8550 ft. Host records from label data include Stemonitis sp. and a ‘‘red, tubular slime mold’’. Agathidium maculosum Brown Figures 46, 49, 54, 55, 87 Agathidium maulosum Brown, 1928: 45 lapsus calami. Agathidium maculosum, Fall, 1934b: 25; Hatch, 1957. Agathidium maculosum var. franciscanum Fall, 1934b: 26. T YPE M ATERIAL :Agathidium maculosum: holotype ? in CNCI labeled ‘‘Victoria, B.C. 11.IV.1921 W. Downes/HOLOTYPE Agathidium maculosum 2784 No. Brown [red label]’’. Agathidium franciscanum: holotype ? in 40 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY MCZC labeled ‘‘Berkeley Cal I-8 1920 H.Dietrich [date handwritten]/ ? /T YPE franciscanum [handwritten, ‘‘type’’ with red underline]/M.C.Z. Type 24037 [red label, number handwritten]/H. C. FALL COLLECTION.’’ T YPE L OCALITY :Agathidium maculosum: Canada, British Columbia, Victoria. Agathidium maculosum var. franciscanum: United States, California, Berkeley. D IAGNOSIS : This species can be distinguished by the prominent, irregular maculations on the elytra (figs. 46, 49), which distinguish the species from all others except A. pulchrum and A. amae, which it greatly resembles. From these species it differs by lacking a sutural elytral stria (fig. 49) and having the anterior clypeal margin less strongly excavated (fig. 49). The male operculum is very prominent, thick, and apically formed into two sharply pointed rami (fig. 54). Also, the macula at the base of the elytra is characteristically broad and extends in lateral, posteriorly directed lobes (fig. 49). D ESCRIPTION : Body moderately large (TBL 5 2.72–3.17 mm), rotund (PNW/TBL 5 0.47–0.53), laterally broadly rounded, moderately strongly contractile. Head yellow-brown; pronotum yellow with large central area of brown; elytra yellow with brown along suture, anteromedially and in two elongate, often confluent medial spots, lateral one located slightly more anterad than other; venter yellow; antennae yellow with basal two club antennomeres brown and apical antennomere yellow; palpi yellow; legs yellow-brown to yellow. Head broad (figs. 46, 49) (MDL/OHW 5 0.55–0.56), dorsally flattened, without medial crease; temporum short, inconspicuous, forming a distinct rim at posterior margin of eye; head finely and sparsely punctate, surfaces shiny and smooth between punctures; eyes large, rounded, finely faceted; frontoclypeal suture moderately prominent medially; clypeal margin slightly excavate, extending about to level of anterolateral margins of frons (figs. 46, 49); labrum entire; antennomere ratios: length I:II:III 5 2.0:1.0:1.9, width VII:VIII:IX 5 0.9:1.0:2.0. Pronotum broad (PNL/PNW 5 0.56–0.58), laterally not strongly produced, anterolateral angle subquadrate, posterolateral angle distinctive, somewhat rounded; punctation similar to that of head. Elytra moderately robust (SEL/ELW 5 0.89–1.17); lateral margins rounded; punctation moderately coarse but sparse, some in indistinct series, surface between punctures shiny and smooth; sutural stria absent. Mesosternum strongly concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion about one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum moderately broad medially (MTL/MTW 5 0.26–0.31), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres slightly expanded and with moderately large ventral field of spatulate setae; mandibular horn, when fully developed, very long, extending posteriorly over front of head in long, slightly curved, flattened, parallel-sided horn, apex flattened and rounded, setose along its length, with prominent subapical fovea and marginal series of fine setae; metafemur slender, unmodified; metasternal fovea transverse, linear, curved, slightly anterior of middle, with line of fine setae. Median lobe in lateral aspect with long, prominent basal portion and submedial constriction, straight dorsad of constriction, apical portion slightly curved, robust, apically truncate (fig. 55); in ventral aspect slender, parallel-sided, apical portion moderately slender, evenly tapered to rounded apex (fig. 54); operculum broad, apically acuminate, pointed apex narrowly emarginate (fig. 54); lateral lobes slender, curved basally, apically pointed with 2 long, stout setae (figs. 54, 55). Female tarsi 5-4-4. D ISTRIBUTION : This species occurs along the west coast of North America from central California north to British Columbia (fig. 87). S PECIMENS E XAMINED : CANADA: British Columbia: Hotel Lake, Pender Harbor, 7 Jul 1920, GR Hopping (1, CASC); Victoria, Vancouver Island (1, OSUC). UNITED STATES: California: state only (1, CASC); Berkeley, 8 Jan 1920 (9, MCZC); San Francisco Bay area, D Giuliani (1, CASC); Gln. Aplne., Jul (1, CASC); Alameda Co.: Redwood Canyon, 1 Jun 1920 (1, CASC); Redwood Canyon, 1 Jun 1920 (6, CASC); Redwood Canyon, 1 Jun 1920 (1, CASC); Contra Costa Co.: Moraga 2005 41MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Valley, 18 Apr 1932 (2, CASC); El Dorado Co.: Blodgett Forest, 27 Aug 1975, Pinus ponderosa log, Berlese, FG Andrews (3, FGAC); Camino, 7 Mar 1964, S Semonoff (1, FGAC); Humboldt Co.: Trinidad, 16 Oct 1984, malaise, F Adams (2, FGAC); Marin Co.: 14 Jan 1920, Deitrich (1, MCZC); Lagunitas, 9 Apr 1911, van Dyke (6, CASC); Taylorville, 28 Dec 1919 (1, CASC); Taylorville, 28 Dec 1919 (4, CASC); Mendocino Co.: Naverro River, 22 Nov 1932, RP Allen (2, CASC); 3 mi S Leggett, 20 Oct 1962, JF Lawrence (2, MCZC); Santa Clara Co.: Stevens Creek Area, 16 Apr 1956 (6, Utah St); Sonoma Co.: Sugar Loaf Ridge St Park, 4 Feb 1980, on Polyporus sp., A Hardy (1, FGAC). Oregon: Newport, 1 Nov 1961, K Goeden (1, RLWE); Astoria (1, OSUC); Croisan Gulch, S Salem, 24 Nov 1974, RL Westcott (4, RLWE); Clackamus Co.: Salmon River near Zigzag, 27 Aug 1982, R Baranowski (1, LUND); Lane Co.: Linslaw Co. Park 24 mi E Florence on Suislaw River, 6 May 1972, tree hollow, rotten wood (1, PECK); Marion Co.: Lake Selmac, 2 May 1969, RL Westcott (2, RLWE); Silver Falls SP, Hwy 214 at S entrance, 8 Nov 1974, RL Westcott (1, RLWE); Polk Co.: Homan Wayside, 4 May 1969, RL Westcott (1, RLWE); Yamhill? Co.: Champage St. Park W Wilsonville, 1 Jan 1980, under bark (4, EMEC). Washington: Beutralia?? (1, MCZC); Olympic NP Dosewallips Ranger Station, 18 Jul 1966, W Gagne J Haddock (1, EMEC). D ISCUSSION : Brown (1928) erected this species under the name ‘‘A. maulosum’’. Since Brown apparently intended to name this species after its most distinguishing feature, the dorsal maculae, it seems likely that he meant to call it A. maculosum, and this is the name others have consistently used to refer to the species. Also, the name on the type label in CNCI is A. maculosum. Based on this evidence we regard A. maulosum as a lapsus calami. As further evidence that the original spelling was unintended, the description following Brown’s description of A. maculosum is apparently also misspelled as ‘‘Callops olahomensis’’ rather than ‘‘Callops oklahomensis’’ for a species from Oklahoma. Agathidium maculosum var. franciscanum was erected based on putative differences in punctation and coloration. However, A. m. franciscanum is clearly the same species as typical A. maculosum, and we synonymize the names. Although Fall (1934b) claimed that the female tarsal formula is 4-4-4, we have found it to be 5-4-4 in female specimens examined. Agathidium maculosum has been collected during every month of the year. Label data indicate it has been found in rotting wood in conifer forests. A single elevation record is from 3600 ft. Host records include Polyporus sp. and a ‘‘slime mold’’. A. pulchrum LeConte Figures 21, 22, 44, 45, 51, 56, 57, 88 Agathidium pulchrum LeConte, 1853: 286; Horn, 1880; Leng, 1920; Fall, 1934b; Hatch, 1957; Silfverberg, 1979 (synonymized with A. pulchellum Wankowicz, 1869); Daffner, 1985 (distinguished from A. pulchellum Wankowicz, name resurrected); Wheeler, 1990 (description of larva). Agathidium mandibulatum Mannerheim, 1853: 203; Horn, 1880 (synonymized with A. pulchrum); Leng, 1920. T YPE M ATERIAL :Agathidium pulchrum: lectotype (designated here to clarify assignment of this name to this species), ? in MCZC labeled ‘‘[circular gold disc]/TYPE 3176 2 [numbers handwritten, 2/3 of label red]/pulchrum 2 [handwritten]’’. LeConte did not indicate the number of specimens on which he based his description. We have selected a specimen from the MCZC that is labeled as the type to be the lectotype. Agathidium mandibulatum: Type not examined. T YPE L OCALITY :Agathidium pulchrum: United States, California, San Jose. Agathidium mandibulatum: United States, Alaska. D IAGNOSIS : This species can be distinguished by the prominent, irregular maculations on the elytra (figs. 44, 45, 51), which distinguish the species from all others except A. maculosum and A. amae, which it greatly resembles. From A. maculosum it differs by having a distinctive sutural elytral stria (fig. 51) and by having the anterior clypeal margin very strongly excavated (figs. 22, 51). Also, the male left mandibular horn, when fully formed, is more strongly curved (figs. 21, 22). The median lobe of the aedeagus in ventral aspect has the apical portion relatively broad and robust and the operculum reduced to a rounded, hyaline lobe (fig. 56), which distinguishes this species from both A. 48 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY lateral angle subquadrate, posterolateral angle distinctive, but rounded; punctation and surface microsculpture similar to that of head, some specimens nearly impunctate. Elytra robust, broad (SEL/ELW 5 0.83–0.99); coarsely punctate, more dense apically, not in series, with fine irregular microreticulation on surfaces between punctures; sutural stria about one-half length of elytron on most specimens. Mesosternum strongly concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion less about one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum relatively broad medially (MTL/MTW 5 0.36), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres slightly expanded and with small ventral field of spatulate setae; left mandible unmodified in all specimens examined; metafemur very slender, otherwise unmodified; metasternal fovea small, slightly anterad of middle, slightly transverse with small brush of fine, dense setae. Median lobe in lateral aspect robust, with large basal region and sub-basal constriction, broad and slightly curved distad of constriction, apical portion long, slender and strongly and evenly curved dorsad (fig. 63); in ventral aspect broad, lateral margins subparallel, apical portion broad, broadly acuminate to sharp apex, orifice large (fig. 62); operculum fused to proximal margin of orifice, long, flat, slightly expanded, apex broad and medially emarginate (fig. 62); lateral lobes long, slender, curved basally, apex narrowly rounded with 2 long setae (figs. 62, 63). Female tarsi 5-4-4. D ISTRIBUTION : This far northern species is known from Alaska and the Yukon south through British Columbia and Alberta east to high elevation areas of New Hampshire (fig. 89). S PECIMENS E XAMINED : CANADA: Alberta: Jumpinground Creek, 20 Oct 1962, forest litter, BF and JL Carr (4, CARR); Elke Island NP White Spruce Trail, 9 May 1984, moose dung and Sphagnum, RS Anderson (2, CNCI); Waiparous, 4 Sep 1960, on ‘‘toadstools’’, BF and JL Carr (3, CARR); 3.8 m N Cadomin, 4 Aug 1985, lodgepole pine forest, FIT, RS Anderson (1, CNCI); Demmitt, 3 km W 90 km NW Grand Prairie, 12 Jun 1984, poplar forest, FIT, S and J Peck (1, CNCI). British Columbia: Mt Robson P Park, Berg Lake Trailhead, 11 Jul 1984, FIT, RS Anderson (2, PECK); Barkeyville, 15 Jul 1969, on forest floor, BF and JL Carr (1, CARR). Manitoba: Devils Lake 100 km S Grand Rapids, 7 Jun 1984, pine-aspen forest, FIT, S and J Peck (1, CNCI); Churchill, 3 Aug 1950, WJ Brown (1, CNCI). Yukon: Whitehorse, 4 Sep 1936, SM Williams (1, MCZC); Dempster Hwy km 141 Blackstone River, 6 Jul 1985, dung traps, SA Marshall (21, PECK); Carmacks, 2 Sep 1936, SM Williams (2, MCZC); km 147 Dempster Hwy, Fen Willows-spruce, 1 Jul 1985, dung pan trap, SA Marshall (3, PECK). UNITED STATES: Alaska: Tolovana Rd 1 Elliot Hwy mi 57, 18 Jul 1985, mushroom trap, SA Marshall (1, PECK); Delta River mi 227 Rich. Hwy., 5 Jul 1958, Lindroth (1, MCZC); Kenai Pen. E Skilak Rd Jct Rt 1, 25 Jun 1984, aspenspruce forest, malaise FIT, S and J Peck (1, PECK); Circle Hot Springs, 6 Aug 1984, alderpoplar-willow forest litter, S and J Peck (1, PECK); Kenai Pen. Summit Lake, Rt 1, 26 Jun 1984, treeline dwarf willow litter, Berlese, S and J Peck (2, PECK); Denali NP Reily Creek Camp, 10 Aug 1984, 1700 9 , taiga forest, on ‘‘mushrooms’’, Berlese, S and J Peck (4, PECK); Chena Ridge 5 mi W Fairbanks, 27 Jul 1984, alder litter, Berlese, S and J Peck (2, PECK); Tenana, 21 Aug 1936, SM Williams (1, MCZC). New Hampshire: Mt Washington, 7 Jul 1899 (1, MCZC); White Mts, subalpine (8, MCZC); White Mts, 4 Jul 1896, subalpine (15, MCZC); Mossiland, 24 Aug 1896 (1, MCZC). D ISCUSSION : The names synonymized here, A. athabascanum and A. alticola, were erected based in part on purported differences in the shape of the lateral margin of the pronotum. However, we find that the types of each of these fall well within the range of variation of a single species diagnosed from the shape of the median lobe of the aedeagus. Agathidium athabascanum comes first in Fall’s (1934b) treatment, so we selected it as the valid name. This is one of the most northerly occurring species in North America. Specimens have been collected from typical northern habitats such as alder, aspen, conifer, and mixed forests, including areas near or north of the tree line. The only host records from label data are ‘‘toadstools’’ and ‘‘mushrooms’’. 2005 49MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Agathidium columbianum Fall Figures 8, 64, 65, 89 Agathidium columbianum Fall, 1934b: 121; Hatch, 1957. T YPE M ATERIAL : Holotype, ? in MCZC labeled ‘‘Terrace, B.C. Mrs. M. E. Hippisley 1932 [number handwritten]/ ? /TYPE columbianum [name handwritten, red line under ‘‘TYPE’’]/M.C.Z. Type 24032 [number handwritten, red label]/H.C. FALL COLLECTION/Agathidium columbianum Fall. [handwritten, red line around border]’’. Only the single holotype specimen of this species is known. T YPE L OCALITY : Canada, British Columbia, Terrace. D IAGNOSIS : This species differs from other members of this group by the concolorous, light brown dorsal surfaces, the anterior clypeal margin convex and extending distinctly, but not greatly, beyond the anterolateral margins of the frons (fig. 8), the mesosternum without a ventrally extending, flattened lobe between the mesocoxae, males without a left mandibular horn, the median lobe of the aedeagus in lateral aspect with the apex pointed and straight (fig. 65), and the operculum rounded without apical emargination (fig. 64). Agathidium columbianum is most similar to A. athabascanum, but that species has the apex of the median lobe of the aedeagus sharply pointed and strongly curved dorsad (fig. 65) and the operculum is apically distinctly emarginate (fig. 64). D ESCRIPTION : Body large (TBL 5 2.74 mm), robust (PNW/TBL 5 0.49), broadly rounded, moderately contractile. Head, pronotum, and elytra yellow-brown to yellow-red; venter yellow to yellowbrown; antennae and palpi yellow; legs yellow to yellow brown. Head broad (fig. 8) (MDL/OHW 5 0.61), somewhat convex, without transverse, medial crease; temporum short, often forming distinct rim at posterior margin of eye (fig. 8); head finely punctate, punctures sparse, only one size present; eyes large, rounded, finely faceted; frontoclypeal suture moderately well defined medially; clypeal margin curved, extending distinctly beyond anterolateral margins of frons (fig. 8); anterior margin of labrum entire; antennomere ratio: length I:II:III 5 1.8:1.0:1.6. Pronotum broad (PNL/PNW 5 0.56), laterally not strongly produced, anterior margin not strongly concave, anterolateral angle subquadrate, posterolateral angle distinctive, but rounded; punctation and surface microsculpture similar to those of head. Elytra robust, broad (SEL/ELW 5 1.13); coarsely punctate, more dense apically, not in series, with fine irregular microreticulation on surfaces between punctures; sutural stria about one-half length of elytron on most specimens. Mesosternum strongly concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion less than one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum relatively broad medially (MTL/ MTW 5 0.34), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres slightly expanded and with small ventral field of spatulate setae; left mandible unmodified in all specimens examined; metafemur very slender, otherwise unmodified; metasternal fovea small, slightly anterad of middle, slightly transverse with small brush of fine, dense setae. Median lobe in lateral aspect robust, with large basal region and subbasal constriction, broad and slightly curved distad of constriction, apical portion long, slender, directed dorsad and nearly straight (fig. 65); in ventral aspect broad, lateral margins subparallel, apical portion broad, slightly acuminate to sharp apex, orifice large (fig. 64); operculum fused to proximal margin of orifice, long, flat, slightly expanded, apex broadly truncate, not emarginate (fig. 64); lateral lobes long, slender, curved basally, apex narrowly rounded with 2 long setae (figs. 64, 65). Female unknown. D ISTRIBUTION : This species is known only from the type locality in British Columbia (fig. 89). Agathidium rusticum Fall Figures 9, 16, 66, 67, 89 Agathidium rusticum Fall, 1934b: 117. T YPE M ATERIAL : Holotype, ? in MCZC labeled ‘‘N.H./Crawford Notch 9/25/08/TYPE rusticum [‘‘rusticum’’ handwritten, red line 50 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY under ‘‘TYPE’’]/M.C.Z. Type 24044 [number handwritten, red label]/H.C. FALL COLLECTION’’. T YPE L OCALITY : United States, New Hampshire, White Mountains, Crawford Notch. D IAGNOSIS : This species differs from others in the group by having concolorous dorsal surfaces, the head and pronotum with prominent and distinct microreticulation consisting of fine, impressed lines forming small, isodiametric cells, antennomere VII large, 2 times length of VIII and distinctly wider (fig. 16), the male metasternal fovea large, transverse, located slightly anterad of middle, the median lobe relatively straight in lateral aspect, with small basal portion, the apex sharply pointed and strongly curved dorsad (fig. 67). The female has 4-4-4 tarsi. D ESCRIPTION : Body relatively small (TBL 5 2.48–2.82 mm), robust (PNW/TBL 5 0.45–0.52), laterally broadly rounded, moderately contractile. Head red to dark red with large paler area medially, clypeus yellow; pronotum light red; elytra red to dark red; venter yellow to dark red; basal antennomeres red-yellow, club antennomeres brown; palpi yellow; legs yellow to yellow-red. Head broad (fig. 9) (MDL/OHW 5 0.55– 0.60), flattened, often with prominent transverse, medial crease; temporum very short, often forming distinct rim at posterior margin of eye; head finely punctate, with microsculpture of fine cells on surfaces between punctures; eyes large, rounded, finely faceted; frontoclypeal suture moderately well defined medially; clypeal margin moderately excavate (fig. 9); anterior margin of labrum entire; antennomere ratios: length I:II:III 5 1.6:1.0:1.3, width VII:VIII:IX 5 1.3:1.0:1.5. Pronotum broad (PNL/PNW 5 0.61–0.65) (fig. 16), laterally not strongly produced, anterolateral angle subquadrate, posterolateral angle distinctive, but rounded; punctation and surface microsculpture similar to those of head. Elytra robust (SEL/ELW 5 0.87– 1.12); punctation coarser than pronotum, but sparse, surface between punctures shiny and smooth; sutural stria about one-half length of elytron on most specimens. Mesosternum strongly concave posteriorly, with prominent lobe extending ventrally between mesocoxae, anterior portion less than one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum relatively narrow medially (MTL/MTW 5 0.19–0.24), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres moderately broad and with small ventral field of spatulate setae; mandibular horn absent on most specimens, one specimen with left mandible thickened and extending up and slightly over right mandible; metafemur moderately slender, unmodified; metasternal fovea large, transversely linear and slightly curved, located slightly anterad of middle, smaller and rounded on some specimens. Median lobe in lateral aspect slender, evenly curved from base, basal portion not large, apical portion narrowed, apex slender and curved dorsad, pointed, in some specimens more distinctly hooked (fig. 67); in ventral aspect slender, lateral margins subparallel, apical portion with lateral margins slightly sinuate, convergent to narrowly rounded apex (fig. 66); operculum divided into two short, parallel, flat, truncate rami (fig. 66); lateral lobes long, slender, evenly curved, apices narrowly rounded with 2 long, stout setae (figs. 66, 67). Female tarsi 4-4-4. D ISTRIBUTION : This species is known from northeastern North America (fig. 89). S PECIMENS E XAMINED : CANADA: Ontario: Chaffeys Locks QUBS, 7 Sep 1980, forest, malaise FIT, S Peck (2, CNCI). UNITED STATES: New Hampshire: White Mts SUBALPINE (1, MCZC). New York: Slide Mt, 3 Jun 1941, W Spector (1, CASC). D ISCUSSION : The antennae in this species approach the state found in many other nonAgathidium Agathidiini where antennomere VII is distinctly longer and wider than VIII. However, in other respects, such as a distinctive temporum and postocular carina and excavate anterior clypeal margin, the species resembles other A. pulchrum group members. Agathidium repentinum Horn Figures 10, 47, 68, 69, 90 Agathidium repentinum Horn, 1880: 304; Leng, 1920; Fall, 1934b. T YPE M ATERIAL : Holotype, / in MCZC la- 2005 51MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II beled ‘‘Wt Mts N. H. Woods/HOLOTYPE 3024 [number handwritten, red label]/A. repentinum Horn [handwritten]’’. Horn had a single specimen when he described this species and it is a holotype by monotypy. T YPE L OCALITY : United States, New Hampshire, White Mountains. D IAGNOSIS : This species differs from others in the group by having concolorous dorsal surfaces, the head and pronotum prominently and distinctly microreticulate, antennomere VII similar in size to VIII, the male metasternal fovea round, located slightly posterad of middle, the median lobe relatively strongly curved basally, with a very large basal portion, the apex sharply pointed, long, and straight (fig. 69), the male left mandibular horn, when present, consisting of a flattened, spine-ike process extending posterad over the clypeus. The female has 4-4-4 tarsi. D ESCRIPTION : Body moderately large (TBL 5 2.67–2.85 mm), rotund (PNW/TBL 5 0.44), laterally broadly rounded, moderately contractile. Head, pronotum, and elytra dark red; venter dark red-brown to yellow-red; antennae yellow basally, apical 5 antennomeres brown; palpi yellow-brown; legs yellow-red. Head broad (fig. 10) (MDL/OHW 5 0.59– 0.66), dorsally flattened, without transverse crease; temporum short, forming distinct rim at posterior margin of eye (fig. 10); head finely and moderately densely punctate, surfaces between punctures with conspicuous microreticulation of fine, isodiametric cells; eyes large, rounded, finely faceted; frontoclypeal suture obsolete medially; clypeal margin slightly excavate, extending nearly to anterolateral margins of frons (fig. 10); labrum entire; antennomere ratios: length I:II:III 5 2.1: 1.0:1.2, width VII:VIII:IX 5 1.0:1.0:1.8. Pronotum broad (PNL/PNW 5 0.63–0.65), laterally not strongly produced, anterolateral angle subquadrate, posterolateral angle distinctive, similar in form to anterolateral angle; punctation similar to that of head, with similar microreticulation. Elytra very broad and robust, lateral margins rounded (SEL/ ELW 5 1.21–1.32); punctation slightly more coarse than on pronotum, but sparse, surface between punctures shiny and smooth; sutural stria about one-half length of elytron. Mesosternum strongly concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion much shorter than posterior portion; medial longitudinal carina absent on anterior portion. Metasternum moderately broad medially (MTL/ MTW 5 0.29–0.31), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres slightly expanded and with moderately large ventral field of spatulate setae; left mandibular horn when fully developed short, flattened, spinous, extending posteriorly over clypeus in adpressed fashion, without apical fovea, many specimens unmodified; metafemur moderately slender, unmodified; metasternal fovea submedial, moderately large, transversely slightly oval, with large brush of fine, dense setae. Median lobe in lateral aspect with large, robust basal portion and submedial constriction, slightly curved distal to constriction, moderately robust, apical portion long, straight, slender, tapering to long slender point (fig. 69); in ventral aspect moderately slender, lateral margins tapering gradually to broadly pointed apex (fig. 68); operculum a broad, rounded, hyaline lobe (fig. 68); lateral lobes slender, curved basally, apically narrowly rounded and each with 2 long, stout setae (figs. 68, 69). Female tarsi 4-4-4. D ISTRIBUTION : This is a northern species collected from Alaska east to New Hampshire (fig. 90). S PECIMENS E XAMINED : CANADA: Alberta: TP25Rg3 W5 mer, 14 Nov 1973, B and J Carr (1, CARR). Manitoba: Churchill, 15 Jun 1937, WJ Brown (2, CNCI). Yukon: Dempster Hwy, km 141 Blackstone River, 6 Jul 1985, mushroom traps in Sphagnum, SA Marshall (1, PECK). UNITED STATES: Alaska: Denali NP, Reily Creek Camp, 10 Aug 1984, 1700 9 , winter moose dung, taiga-willow forest, Berlese, S and J Peck (1, PECK); Nenana, 11 Aug 1984, riverside poplar forest litter, Berlese, S and J Peck (15, PECK). New Hampshire: Hermit Lake, White Mts, 6 Aug 1906, S Martin (1, MCZC); Subalpine, White Mts, 30 Jun 1896 (2, MCZC); Coos Co.: 1 mi NE East Inlet Dam, 26 Nov 1986, conifer leaf litter, DS Chandler (3, CNCI). D ISCUSSION :Agathidium repentinum was described from a single female specimen. However, we are confident assigning male specimens to the species based on the pres- 52 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY ence of prominent microreticulation on especially the head and pronotum. The species is relatively rarely collected but is widespread across the northern latitudes. It has been collected from conifer and poplar forests and ‘‘taiga-willow forest’’, A single elevation record is from 1700 ft. Agathidium marae Miller and Wheeler, new species Figures 23, 24, 70, 71, 91 T YPE M ATERIAL : Holotype, ? in WSUC labeled ‘‘WASHINGTON: Mt. Spokane SP, nr Bald Knob Cmpgr. Spokane Co., 5200 ft. W. J. Turner 28-VI-1977 Malaise trap/HOLOTYPE Agathidium marae Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : United States, Washington, Spokane Co., Mt Spokane State Park, near Bald Knob Campground, 5200 9 . D IAGNOSIS : This species differs from others in the group by having concolorous dark red-brown elytra, no microreticulation on the head and pronotum, the anterior clypeal margin very strongly excavated (fig. 24), and the temporum forming a very abrupt, prominent rim at the posterior margin of the eye (fig. 24). Also, many males have a prominent left mandibular horn and a variously developed right frontal horn (figs. 23, 24). These horns are highly variable in development, but the right frontal horn is only well developed when the mandibular horn is also well developed. The median lobe of the aedeagus in lateral aspect is slender and elongate with a small basal portion and relatively simple apex (fig. 71). The lateral lobes are broadly fused dorsally for about one-third length of the median lobe of the aedeagus (figs. 70, 71). Females have 5-4-4 tarsomeres. D ESCRIPTION : Body moderately large (TBL 5 2.52–2.67 mm), rotund (PNW/TBL 5 0.43–0.45), laterally broadly rounded, strongly contractile. Head dark red-brown with indistinct red medial macula; pronotum, elytra, and venter dark red-brown; antennae, palpi, and legs yellow-red. Head broad (fig. 24) (MDL/OHW 5 0.64– 0.66), dorsally flat, often with indistinct, transverse, medial crease; temporum short, forming rim at posterior margin of eye (fig. 24); head finely punctate, surfaces shiny and smooth between punctures; eyes large, rounded, finely faceted; frontoclypeal suture obsolete medially; clypeal margin strongly excavate (fig. 24); labrum anteromedially entire; antennomere ratios: length I:II:III 5 2.2: 1.0:1.6, width VII:VIII:IX 5 0.9:1.0:1.9. Pronotum broad (PNL/PNW 5 0.59–0.61), laterally not strongly produced, lateral margins moderately broad, anterolateral angle subquadrate, posterolateral angle distinctive, but more rounded than anterolateral angle; punctation similar to that of head. Elytra moderately elongate (SEL/ELW 5 0.99– 1.00); lateral margins broadly rounded; punctation coarser than pronotum, surface between punctures shiny and smooth, some punctures in indistinct series; sutural stria about one-half length of elytron. Mesosternum strongly concave posteriorly, with prominent lobe extending ventrally between mesocoxae, anterior portion much less than one-half length of posterior portion; medial longitudinal carina obsolete on anterior portion. Metasternum broad medially (MTL/ MTW 5 0.30–0.31), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres moderately broad and with large ventral field of spatulate setae; mandibular horn, when present, very long, curving posteriorly over front of head, slightly flattened in cross-section, punctate and setose along its length, with very prominent apical fovea (figs. 23, 24) and very large series of fine, dense setae; frons with long, straight, pointed horn extending upward from surface at right of clypeus, apex reaching to near apex of mandibular horn (figs. 23, 24), some specimens with only a small prominence in this position, some specimens with no frontal horn or prominence but with mandibular horn, some specimens with no horns whatsoever; metafemur slender, unmodified; metasternal fovea anterior, large, transverse, curved, with large series of fine, dense setae. Median lobe in lateral aspect elongate, slender, with small basal portion, curved basally, apical portion short, slender, slightly curved dorsad, apex narrowly rounded (fig. 71); in ventral aspect slender, lateral margins subparallel, slightly convergent in apical half, 2005 53MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II apical portion broadly triangular, apex narrowly rounded (fig. 70); operculum reduced to an inconspicuous, short, flattened structure that is apically bilobed (fig. 70); lateral lobes long, broadly fused along dorsal margin for about one-third length of median lobe, apices narrowly rounded, each with 2 long, stout setae (figs. 70, 71). Female tarsi 5-4-4. E TYMOLOGY : This species is named in honor of Darlene Marie Platt, significant other of the junior author, for her encouragement and support. D ISTRIBUTION : This species is known from the Pacific Northwest east to Montana (fig. 91). P ARATYPES : CANADA: British Columbia: 5.6 mi E E Border Glacier NP on #1, 11 May 1984, FIT, RS Anderson (10, CNCI); Prospect Creek W Merritt, 27 Jun 1984 (1, QDWC). UNITED STATES: Oregon: Clackamas Co.: 1.5 mi S Jct US 26 Ore 35, 11 Jul 1975, 3500 9 , conifer forest, Berlese, A Newton, M Thayer (8, MCZC); Mt Hood NF, Still Creek Cpgd, 10 Jul 1975, 3700 9 , litter, mixed conifer forest, Berlese, A Newton (2, MCZC). D ISCUSSION : The unusual horn arising from the right side of the frons of some male specimens (figs. 23, 24) is particularly unique to this species. Behavioral correlates of this particular morphology have not been observed. This species has been collected in conifer forests. Altitude records from label data are from 3500 to 5200 ft. Agathidium rotundulum Mannerheim Figures 11, 72, 73, 91 Agathidium rotundulum Mannerheim, 1852: 370; Horn, 1880; Leng, 1920; Fall, 1934b. Agathidium kincaidi Hatch, 1936: 39; 1957 (synonymized with A. rotundulum). T YPE M ATERIAL :Agathidium rotundulum: Lectotype (designated here to fix this name with the species), ? in MCZC labeled ‘‘95/ Type 7945 [red label]/A rotundulum Sitkha Mannh. [handwritten]’’. Agathidium kincaidi: holotype, / in USNM labeled ‘‘Loveland, Wash. April 8, 1913/TYPE ? Agathidium (Neoceble) kincaidi 1935—M.H. Hatch [handwritten, red label]’’. T YPE L OCALITY :Agathidium rotundulum: United States, Alaska. Agathidium kincaidi: United States, Washington, Loveland. D IAGNOSIS : This species differs from other members of the group by its small size (TBL , 2.2 mm), concolorous elytra, the anterior clypeal margin extending approximately to level of anterolateral margins of frons, antennomere II subequal in length to III, the male metasternal fovea absent, the male left mandibular horn absent, the median lobe of the aedeagus in lateral aspect with a very large basal portion and the apical portion moderately long, straight, and apically slightly expanded and rounded (fig. 73). The female tarsi are 4-4-4. D ESCRIPTION : Body small (TBL 5 1.78– 2.12 mm), rotund (PNW/TBL 5 0.50–0.52), laterally broadly rounded, moderately strongly contractile. Head piceous, clypeus red; pronotum redbrown, yellow along margins; elytra redbrown; venter dark red-brown to yellow-red; antennae and palpi yellow-brown; legs yellow-red. Head broad (MDL/OHW 5 0.65), dorsally flattened, with prominent medial crease; temporum very short, inconspicuous; head finely and sparsely punctate, surfaces shiny and smooth between punctures; eyes large, rounded, finely faceted; frontoclypeal suture obsolete medially; clypeal margin very slightly excavate, extending about to level of anterolateral margins of frons (fig. 11); labrum entire; antennomere ratios: length I:II: III 5 1.3:1.0:0.8, width VII:VIII:IX 5 1.0: 1.0:1.6. Pronotum broad (PNL/PNW 5 0.57–0.62), laterally not strongly produced, anterolateral angle subquadrate, posterolateral angle distinctive, somewhat rounded; punctation similar to that of head, very fine and sparse. Elytra broad and robust, lateral margins rounded (SEL/ELW 5 0.88–0.98); punctation slightly more coarse than on pronotum, but sparse, surface between punctures shiny and smooth; sutural stria absent. Mesosternum moderately concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion shorter than posterior portion; medial longitudinal carina absent on anterior portion. Metasternum moderately narrow medially (MTL/ MTW 5 0.22), slightly convex, setose; oblique carinae absent. 54 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Male tarsi 5-5-4; proand mesobasotarsomeres slightly expanded and with moderately large ventral field of spatulate setae; left mandible not modified; metafemur moderately slender, unmodified; metasternal fovea absent. Median lobe in lateral aspect with large, robust basal portion and submedial constriction, relatively straight distal to constriction, apical portion moderately long, straight, apically slightly expanded and rounded (fig. 73); in ventral aspect slender lateral margins subparallel, apical portion elongate-triangular, apex narrowly rounded (fig. 72); operculum broad, short, flat, apically broadly rounded (fig. 72); lateral lobes moderately broad basally, slightly curved, apically narrowly rounded and with 2 long, stout setae (figs. 72, 73). Female tarsi 4-4-4. D ISTRIBUTION : This species occurs along the west coast of North America from Alaska south to northern California (fig. 91). S PECIMENS E XAMINED : CANADA: Alberta: 11 mi NE Robb, 4 Aug 1985, lodgepole pine forest, FIT, RS Anderson (1, CNCI). British Columbia: Queen Charlotte Island (1, CASC); Monashee Mtn nr Cherryville, 10 Aug 1982, 1400–1600 m, M Sorensson (1, LUND); Toco, 28 Apr 1984, R and J Camenisch (2, CNCI); Charlotte Islands, Grahm Island, 1 mi NW Tiell, 27 Jun 1984, Sitka spruce, hemlock forest, RS Anderson (15, CNCI); Vancouver (5, MCZC); Queen Charlotte Island (1, MCZC); Massett, Graham Island, moss, Clark (1, MCZC); Vancouver Island (2, MCZC). UNITED STATES: Alaska: Seward, 26 Jun 1984, mature poplar-spruce, malaise-FIT, S and J Peck (1, PECK). California: Santa Cruz Mts (1, MCZC); Amador Co.: 1 mi W Pine Grove, 24 Jun 1975, leaf litter, mixed hardwood conifer forest, A Newton, M Thayer (1, MCZC); Del Norte Co.: Crescent City, 1 Mar 1978, pine duff, Berlese, TR Haig (12, FGAC); Humboldt Co.: Kneeland, 9 Jan 1980, redwood duff, Berlese, TR Haig (3, FGAC); Blue Lake, 21 Oct 1977, redwood duff, Berlese, TR Haig (16, FGAC); Orick, 29 Oct 1976, redwood duff, Berlese, TR Haig (25, FGAC). Oregon: Linn Co.: 32.5 mi S Sweet Home, 20 Oct 1967, Abies, spruce, pine, hemlock duff, Malcolm (1, CUIC). Washington: Jefferson Co.: 20 km NE Kalaloch, 1 Sep 1982, M Sorensson (4, LUND). D ISCUSSION :Agathidium rotundulum and A. kincaidi are synonyms based on our examination of the type specimens and following Hatch (1957). This species has been collected from various forest habitats including redwood, lodgepole pine, poplar-spruce, Sitka sprucehemlock, etc. Altitude records are from 1400 to 1600 m. Agathidium aristerium Wheeler Figures 25, 26, 74, 75, 91 Agathidium aristerium Wheeler, 1987: 399; 1990 (description of larva). T YPE M ATERIAL : Holotype, ? in CUIC (see Wheeler, 1987). T YPE L OCALITY : United States, New York, Tompkins County, Smith Woods, Trumansburg. D IAGNOSIS : This species is characterized by the dorsally red, shiny coloration without maculae or microreticulation, the anterior clypeal margin strongly excavate (fig. 26), the head narrowed immediately posterad of the compound eye without a posterior ocular rim (fig. 26), the male with a metasternal fovea located medially along the midline of the metasternum, the male mandibular horn, when present, formed by extension of the apex of the mandible in a long, dorsally curved, flattened structure (figs. 25, 26), and the median lobe in lateral aspect slender, evenly and shallowly curved (fig. 75) and in ventral aspect with apex expanded and broadly pointed (fig. 74). Females have 4-44 tarsi. D ESCRIPTION : Body moderately large (TBL 5 2.33–2.68 mm), robust, broadly rounded (PNW/TBL 5 0.44–0.46), moderately contractile. Head dark red; pronotum and elytra red to dark red; venter red-yellow; basal antennomeres red-brown, club antennomeres red-yellow; palpi brown; legs red-yellow. Head broad (fig. 26) (MDL/OHW 5 0.61– 0.64), broadly convex, without transverse, medial crease; temporum short, inconspicuous (fig. 26); head finely and sparsely punctate, surface between punctures shiny; eyes large, rounded, finely faceted; frontoclypeal suture obsolete medially; clypeal margin strongly excavate (fig. 26); anterior margin of labrum slightly concave; antennomere ratios: length I:II:III 5 1.4:1.0:1.1, width VII: VIII:IX 5 1.1:1.0:2.6. Pronotum broad (PNL/PNW 5 0.63–0.72), lateral margin rel- 2005 55MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II atively short, not strongly produced, posterolateral angle more angulate than anterolateral angle; punctation similar to that of head. Elytra broad, robust (SEL/ELW 5 1.06–1.21); punctation fine and sparse; sutural stria about one-half length of elytron on most specimens. Mesosternum strongly concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion less than one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum broad medially (MTL/MTW 5 0.20–0.22), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres moderately broad and with moderately large ventral field of spatulate setae; mandibular horn, when fully developed, formed by apex of mandible extending in broad, flat, sweeping curve dorsad and to the right over the right mandible (figs. 25, 26), finely setose but without prominent setose fovea, some specimens with mandible variously thickened, others without modification; metafemur moderately broad, unmodified; metasternal fovea small, round, medial, with small brush of fine, dense setae. Median lobe in lateral aspect slender, slightly curved basally, apical portion short, directed slightly dorsad, narrowly rounded (fig. 75); in ventral aspect slender, lateral margins subparallel, apical portion broad, apex abruptly, but evenly, narrowed to broadly pointed apex (fig. 74); operculum an inconspicuous, broad flat lobe (fig. 74); lateral lobes slender, curved basally, apically narrowly rounded with 2 long setae (figs. 74, 75). Female tarsi 4-4-4. D ISTRIBUTION : This species is known from northeastern North America (fig. 91). S PECIMENS E XAMINED : CANADA: Manitoba: Rennie Lily Pond Whiteshell Prov. Park, 6 Jun 1984, maple poplar forest, FIT, S and J Peck (2, CNCI); Rennie, Lily Pond Whiteshell Prov. Park, 6 Jun 1984, maple, poplar forest, FIT, S and J Peck (13, PECK). Ontario: Chaffeys Locks QUBS, 21 Sep 1980, malaise-FIT, S Peck (1, CUIC); Markham, 24 May 1980, deciduous forest, RS Anderson (1, CNCI); 25 km W Ignace, 75 km E Dryden, 5 Jun 1984, fir-maple forest, FIT, S and J Peck (1, CNCI); Hackston, 20 km SE Kemptville, 24 Jun 1984, FIT, M Kaulbars (5, PECK); Shirleys Bay 15 km W Ottawa, 15 May 1984, M Kaulbars (3, PECK); Ottawa, Stittsville, 10 Jul 1977, M Sanbourne (1, CNCI). Quebec: Hull, Gatineau Park nr Pinks Lake, 8 Aug 1979, S Peck, A Davies (1, PECK); Hull, Gatineau Park nr Pinks Lake, 13 Apr 1980, malaise (2, CUIC). UNITED STATES: Illinois: Union Co.: Pine Hills Field Station, 15 May 1967, rotten log, Berlese, JM Campbell (3, CNCI). Michigan: Kalamazoo Co.: Hickory Corners, Gull Lake, 11 Jun 1981, FIT, RS Anderson (10, CNCI); Hickory Corners, Gull Lake, 11 Jun 1981, FIT, R Anderson (3, CNCI). New Hampshire: Carroll Co.: The Bowl, 2.5 mi NW Wonalancet, 5 Jun 1985, FIT, DS Chandler (5, CNCI); Coos Co.: Norton Pool, 3 mi NE East Inlet Dam, 25 Jul 1986, FIT, DS Chandler (3, CNCI). New York: Tompkins Co.: Trumansburg, Smith Woods, 6 Aug 1986, on Physarum polycephalum plasmodium on Pleurotus fruiting body, QD Wheeler (20, MCZC). Ohio: Fairfield Co.: 13 Sep, DJ and JL Knull (22, OSUC). D ISCUSSION :Agathidium aristerium has been collected from May to September. Specimens have been found in deciduous and fir forests. A single host record is from a Physarum polycephalum plasmodium on Pleurotus fruiting body (see Wheeler, 1987). All instars of the larva were described by Wheeler (1990). Agathidium atronitens Fall Figures 27, 28, 76, 77, 92 Agathidium atronitens Fall, 1934b: 122. T YPE M ATERIAL : Holotype, / in MCZC labeled ‘‘Urbana ILL 4/20/25/ / /TYPE atronitens [name handwritten, red line under ‘‘TYPE’’]/M.C.Z. Type 24029 [number handwritten, red label]/H.C. FALL COLLECTION’’. T YPE L OCALITY : United States, Illinois, Urbana. D IAGNOSIS : This species is distinguished from others in the group by the concolorous, dark red dorsal surfaces which lack microreticulation, the head narrowed immediately behind eye and without postocular rim (figs. 27, 28), anterior clypeal margin strongly excavated (fig. 28), the male metasternal fovea located medially along the midline of the metasternum, the male mandibular horn straight, extending from middle of mandible in oblique angle to right over the right mandible (figs. 27, 28), and the median lobe in lateral aspect robust, with large basal portion and 56 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 92. Geographic distribution of Agathidium pulchrum-group species: A. atronitens 5v ;A. oregonense 5m ;A. politum 5. . 2005 57MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II submedial constriction and with small apical spine (fig. 77). Females have 4-4-4 tarsi. D ESCRIPTION : Body moderately large (TBL 5 3.15–3.21 mm), robust (PNW/TBL 5 0.48–0.50), broadly rounded, moderately contractile. Head dark red; pronotum and elytra red to dark red; venter red to red-yellow; basal antennomeres red-brown, club antennomeres brown; palpi red; legs red to red-yellow. Head broad (fig. 28) (MDL/OHW 5 0.51– 0.60), flattened, often with prominent transverse, medial crease; temporum very short, without rim at posterior margin of eye (fig. 28); head finely punctate, with microsculpture of fine cells on surfaces between punctures; eyes large, rounded, finely faceted; frontoclypeal suture moderately well defined medially; clypeal margin strongly excavate (fig. 28); anterior margin of labrum entire; antennomere ratios: length I:II:III 5 1.6:1.0: 1.0, width VII:VIII:IX 5 1.0:1.0:1.6. Pronotum broad (PNL/PNW 5 0.55–0.61), laterally not strongly produced, anterolateral angle subquadrate, posterolateral angle distinctive, but rounded; punctation and surface microsculpture similar to those of head. Elytra robust (SEL/ELW 5 1.00); nearly impunctate to finely and sparsely punctate, surface between punctures shiny and smooth; sutural stria about one-half length of elytron on most specimens. Mesosternum strongly concave posteriorly, without prominent lobe extending ventrally between mesocoxae, anterior portion less than one-half length of posterior portion; medial longitudinal carina absent on anterior portion. Metasternum relatively short medially (MTL/MTW 5 0.21– 0.24), slightly convex, setose; oblique carinae absent. Male tarsi 5-5-4; proand mesobasotarsomeres slightly expanded and with small ventral field of spatulate setae; mandibular horn highly variable in length, straight, extending from middle of mandible in oblique angle to right over right mandible (figs. 27, 28), setose, apically pointed, with small, indistinct setose fovea near pointed apex; metafemur moderately broad, unmodified; metasternal fovea medial, transversely slightly ovoid, with large brush of fine, dense setae. Median lobe in lateral aspect robust, basal portion extremely large and robust, with distinct submedial constriction, straight and short distad of constriction, apical portion short, stout, directed slightly dorsad, apically narrowly rounded (fig. 77); in ventral aspect robust, lateral margins subparallel, apical portion broad, apex broadly pointed (fig. 76); operculum broad, flat, apically narrowly emarginate, hyaline and indistinct (fig. 76); lateral lobes slender, curved basally, apically narrowly round with 2 long, stout setae (figs. 76, 77). Female tarsi 4-4-4. D ISTRIBUTION : This species is found in eastern North America from Saskatchewan and New Hampshire south through Oklahoma to Louisiana and Alabama and east to Washington, DC (fig. 92). S PECIMENS E XAMINED : CANADA: Ontario: Sudbury, Laurentian Univ., 26 May 1982, FIT, R DeClerck (1, CNCI); Alfred Bog, Alfred, 13 Aug 1984, M Sanborne (1, CNCI). Saskatchewan: Prince Albert NP 16 km W Christopher, 9 Jun 1984, fir-aspen forest, FIT, S and J Peck (2, CNCI). UNITED STATES: Alabama: Monroe Co.: Big Flat Creek 31 8 36 9 30 0 N, 87 8 24 9 53 0 W, 27 May 1995, upland decid, Berlese, CE Carlton (1, LSAM). District of Columbia: paratype (1, MCZC). Kentucky: 5 mi W Hopkinsville, 22 Sep 1967, deciduous duff, Berlese, JM Campbell (2, CNCI). Louisiana: W Feliciana Co.: 6 mi ESE St Francisville, 17 May 1995, forest litter, D Colby, D Landau (1, LSAM); Cabin Area 30 8 47 9 N, 91 8 15 9 W, 1 Dec 2000, fallen logs, FIT, AR Cline (5, LSAM); Tunica WMA 30 8 55 9 N, 91 8 30 9 W, 28 Nov 1998, FIT, C Carlton, J Johnson, A Tischechkin (3, LSAM). New Hampshire: Grafton Co.: Hubbard Brook Exp. Sta., Bear Brook, 6 Jul 1983, rotten wood, DS Chandler (5, DENH); Straf. Co.: 1 mi SW Durham, 27 Jun 1987, FIT, DS Chandler (1, CNCI). New York: Westchester Co.: Calder Ecology Center, Armonk, 24 Apr 1979, Berlese, K Schmidt (16, AMNH); Calder Ecology Center, Armonk, 24 Apr 1979, Berlese, K Schmidt (4, AMNH). Ohio: Franklin Co.: Columbus, 13 Sep 1975, QD Wheeler (4, QDWC); Columbus, 13 Sep 1975, QD Wheeler (2, QDWC). Oklahoma: Latimer Co.: 5 mi W Red Oak, Dec 1980, K Stephan (2, QDWC); K Stephan (2, QDWC). Pennsylvania: Lebanon Co.: 1 mi E Indian Town Gap, 13 Jul 1969, Berlese, W Michmore (1, CNCI). Tennessee: Sevier Co.: Appalachian Trail at Beech Gap on Clingmans Dome Rd 83 8 26 9 50 0 W, 35 8 36 9 36 0 N, 28 Jun 2001, 1750 9 , forest litter, Berlese, C Carlton, A Tsichechkin, V Mosely (5, 64 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY not clearly fit into any. Therefore, we have elected to place them in their own group. Two of the three species, one eastern and one western, possess a prominent medial lobe on the first visible abdominal sternite, but the third species, the western A. fenderi, does not. The abdominal lobe is a seemingly clear synapomorphy, but the similarities between A. fenderi and the lobed species are not strong, and this grouping may be artificial. KEY TO A. COMPRESSIDENS SPECIES GROUP 1. First visible abdominal sternite with a prominent, medial, lobelike prominence near posterior margin ...................... 2 — First visible abdominal sternite without a prominent, medial lobe . . A. fenderi Hatch 2(1). Size smaller (TBL , 2.6 mm) ; male metasternal fovea very large, circular, located medially; median lobe of aedeagus in ventral aspect with apical portion very abruptly narrowed and sharply acuminate (fig. 96), with operculum large and deeply and broadly bifid with each ramus long, parallel-sided and straight (fig. 96); California and Oregon (fig. 104) ........ ...... A. vesperpressidens new species — Size larger (TBL . 2.6 mm) ; male metasternal fovea moderately large, transverse, located distinctly posterad of middle; median lobe of aedeagus with apex only slightly acuminate, more gradually narrowed (fig 99), with operculum moderately large and rounded, platelike with narrow medial emargination (fig. 99); eastern North America (fig. 104) ........... .............. A. compressidens Fall Agathidium fenderi Hatch Figures 94, 95, 104 Agathidium fenderi Hatch, 1957: 36. T YPE M ATERIAL : Holotype, / in USNM labeled ‘‘Boyer, Ore. May 15, 37 [handwritten]/soil samp [handwritten]/H-314/1-1189/ Agathidium 3 [handwritten, label folded]/ TYPE ? Agathidium (s. str.) fenderi 1953— M.H. Hatch [handwritten, red label]’’. T YPE L OCALITY : United States, Oregon, Lincoln Co., Boyer. D IAGNOSIS : This species differs from other members of this group by lacking a medial lobe on the first visible abdominal sternite. Also, the male metasternal fovea is relatively large, rounded, and located slightly posterad of the middle. The median lobe of the aedeagus is relatively simple (figs. 94, 95) with the operculum reduced to a small, truncate lobe (fig. 94). D ESCRIPTION : Body moderately large (TBL 5 2.16–2.55 mm), broad (PNW/TBL 5 0.48–0.49), robust, rounded, strongly contractile. Head and pronotum piceous to dark red; elytra dark red; venter red-brown; antennae, palpi, and legs red-brown. Head broad (MDL/OHW 5 0.47–0.70), dorsal surface very broadly convex, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes protruding and prominent, large, finely faceted; gula slightly concave, unmodified; clypeus strongly excavate; antennomere ratios: length I:II:III 5 2.1:1.0:1.8, width VII:VIII:IX 5 1.0:1.0:2.2. Pronotum very large, broad (PNL/PNW 5 0.67–0.70), strongly convex, anterolateral lobes strongly produced, lateral margin with rounded, but distinct anteroand posterolateral angles; surface with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.00–1.25); punctation similar to pronotum; sutural stria absent. Flight wings fully developed. Mesosternum broad, anterior portion longer than posterior portion; distinctly concave posteriorly, but not strongly declivitous; medial longitudinal carina well developed on anterior portion. Metasternum narrow medially (MTL/MTW 5 0.12), subhorizontal to slightly convex medially; oblique femoral carinae faint or absent, not meeting medially. Abdominal sternite III (first visible sternite) without prominent medial lobe. Male tarsi 5-5-4; proand mesobasotarsomeres relatively broadly laterally expanded, with large field of ventral spatulate setae; mandibles not modified; metafemur relatively slender, unmodified; metasternal fovea large, prominent, round, with pencil of long, fine, dense setae, located posterad of middle of metasternum. Median lobe in lateral aspect very robust, curved medially, apical portion broad, curved dorsad, apex sharply 2005 65MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Figs. 94–103. Agathidium compressidensand A. iota-group species, aedeagus: 94, 95, A. fenderi: 94, ventral; 95, lateral. 96–98, A. vesperpressidens:96, ventral; 97, median lobe, lateral; 98, left lateral lobe, lateral. 99–101, A. compressidens:99, ventral; 100, median lobe, lateral; 101, left lateral lobe, lateral. 102, 103, A. iota:102, ventral; 103, lateral. pointed (fig. 95); in ventral aspect broad, lateral margins subparallel, apical portion broad, with lateral margins rounded and converging to broadly rounded apex (fig. 94); operculum reduced to small truncate lobe which is apically slightly emarginate (fig. 94); lateral lobes moderately broad, apically somewhat expanded with 2 long, stout subapical setae that are unequal in length (figs. 94, 95). Female tarsi 5-4-4. D ISTRIBUTION . This species is known only from Oregon and northern California (fig. 104). S PECIMENS E XAMINED : UNITED STATES: California: Del Norte Co.: Crescent City, 25 Mar 66 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 104. Geographic distribution of Agathidium compressidens-group species: A. compressidens 5 v ;A. fenderi 5m ;A. vesperpressidens Fall 5m . 2005 67MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II 1978, pine duff, Berlese, TR Haig (1, AMNH). Oregon: Benton Co.: Mary’s Peak, 17 May 1983, 3500 9 ,Ribes litter, DS Chandler (1, DENH); Josephine Co.: 6 mi N Galice, Rogue River, 8 Apr 1972, 500 9 , madrone duff, E Benedict (1, CNCI). D ISCUSSION : The few specimens examined during this study were collected during April and May from soil samples and pine or madrone duff. Elevation records are from 500 to 3500 ft. Agathidium vesperpressidens Miller and Wheeler, new species Figures 96–98, 104 T YPE M ATERIAL : Holotype, / in CASC labeled ‘‘CALIF:Butte Co. 5 mi SE Butte Mds Cherry Hill Cpgd. V-7 thru 9-1976 Fred G. Andrews/Collected flying at dusk/HOLOTYPE Agathidium vesperpressidens Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : USA, California, Butte Co., 5 mi NE Butte Meadows, Cherry Hill Campground. D IAGNOSIS : This species is distinguishable by the presence of a prominent lobe medially on the ventral surface of the first visible abdominal sternite and by the shape of the male genitalia. The median lobe of the aedeagus is very robust; the apical portion consists of a broad basal portion abruptly narrowed to a long, slender spine in ventral aspect (fig. 96), and the operculum is deeply emarginate, consisting of long, parallel-sided, broadly separated rami (fig. 96). The species is most similar to A. compressidens from which it may be distinguished by the shape of the aedeagus (figs. 96–98). Also, A. compressidens is larger (TBL . 2.7 mm) and occurs in eastern North American (fig. 104), whereas A. vesperpressidens is smaller (TBL , 2.6 mm) and occurs in western North America (fig. 104). D ESCRIPTION : Body moderately large (TBL 5 2.43–2.52 mm), broad (PNW/TBL 5 0.48–0.49), robust, rounded, strongly contractile. Head and pronotum red; elytra red, not iridescent; venter yellow to yellow-red; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.55–0.62), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes protruding and prominent, large, finely faceted; gula slightly concave, unmodified; clypeus strong excavate; antennomere ratios: length I:II:III 5 1.6:1.0: 1.2, width VII:VIII:IX 5 1.1:1.0:2.0. Pronotum very large, broad (PNL/PNW 5 0.62– 0.63), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.97–1.05); punctation similar to pronotum; sutural stria absent to faintly present only apically. Flight wings fully developed. Mesosternum broad, anterior portion longer than posterior portion; distinctly concave posteriorly, but not strongly declivitous; medial carina well developed on anterior portion. Metasternum narrow medially (MTL/MTW 5 0.12–0.15), subhorizontal, slightly convex medially; oblique femoral carinae faint or absent, not meeting medially. Abdominal sternite III (first visible sternite) with prominent medial lobe. Male tarsi 5-5-4; proand mesobasotarsomeres somewhat laterally expanded, with small field of ventral spatulate setae; mandibles not modified; metafemur relatively slender, without subapical tooth on posterior margin, but with small subbasal tooth; metasternal fovea very large, prominent, round, with pencil of long, fine, dense setae, located posterad of middle of metasternum. Median lobe in lateral aspect very robust, strongly curved medially, apical portion slender, straight, apex sharply hooked dorsad, sharply pointed (fig. 97); in ventral aspect very broad, apical portion broad basally, strongly narrowed, apically a long, slender process, slightly tapering to sharply pointed apex (fig. 96); operculum large, flat, medially very deeply emarginate, emargination elongaterectangular, each ramus long, parallel-sided, apically truncate (fig. 96); lateral lobes small, slender, curved, shorter than median lobe, apically slightly expanded, each with 2 long, stout subapical setae (figs. 96, 98). Female tarsi 5-4-4. E TYMOLOGY : This species is named from 68 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY the Latin words vesper, meaning ‘‘west’’, and pressidens, a portion of the root for compressidens, the specific name of a similar member of this species group, to signify the similarity of this species with A. compressidens and its western distribution. D ISTRIBUTION : This species has been collected from northern California (fig. 104). P ARATYPES : UNITED STATES: California: Tallac, July (1, CASC); Butte Co.: 5 mi NE Butte Meadows, Cherry Hill Campground, 7 May 1976, collected flying at dusk, F Andrews (1, FGAC); El Dorado Co.: Blodgett Forest, 27 Aug 1975, Pinus ponderosa log, Berlese, F Andrews, M Wasbauer (1, FGAC); Tulare Co.: Sequoia Natl Park, Huckleberry Meadow, 28 May 1984, 8000 9 , R Baranowski (2, LUND); Sequoia Natl Park, Dorst Creek, 31 May 1984, 7000 9 , R Baranowski (6, LUND); Sequoia Natl Park, Lost Grove, 4 Jun 1984, 7000 9 , R Baranowski (1, LUND). D ISCUSSION : This species has been collected from a ponderosa pine log and ‘‘while flying’’. Elevation records are from 7000 to 8000 ft. Agathidium compressidens Fall Figures 99, 100, 104 Agathidium compressidens Fall, 1934b: 106. T YPE M ATERIAL : Holotype, / in MCZC labeled ‘‘Round Knob N.C. 6124 [handwritten]/ / /TYPE compressi-dens [name handwritten, red line under ‘‘TYPE’’]/M.C.Z. Type 24033 [number handwritten, red label]/ H.C. FALL COLLECTION’’. T YPE L OCALITY : United States, North Carolina, Round Knob. D IAGNOSIS : This species is distinguishable by the presence of a prominent lobe medially on the ventral surface of the first visible abdominal sternite and by the shape of the male genitalia. The median lobe of the aedeagus is very robust, the apical portion is broad, flat, and apically pointed (fig. 99), and the operculum is broad, flat, and broadly rounded apically with a deep, narrow medial emargination (fig. 99). The species is most similar to A. vesperpressidens from which it may be distinguished by the shape of the aedeagus. Also, A. vesperpressidens is generally smaller (TBL , 2.7 mm). D ESCRIPTION : Body moderately large, broad (TBL 5 2.84–3.38 mm), robust (PNW/TBL 5 0.48–0.49), rounded, strongly contractile. Head and pronotum red; elytra red, not iridescent; venter yellow to yellow-red; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.58–0.61), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes protruding and prominent, large, finely faceted; gula slightly concave, unmodified; clypeus strongly excavate; antennomere ratios: length I:II:III 5 1.6:1.0: 1.1, width VII:VIII:IX 5 1.1:1.0:2.3. Pronotum very large, broad (PNL/PNW 5 0.58– 0.61), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.94–1.01); punctation similar to pronotum; sutural stria absent to faintly present only apically. Flight wings fully developed. Mesosternum broad, anterior portion longer than posterior portion; distinctly concave posteriorly, but not strongly declivitous; medial longitudinal carina well developed on anterior portion. Metasternum narrow medially (MTL/MTW 5 0.14–0.16), subhorizontal, slightly convex medially; oblique femoral carinae faint, not meeting medially. Abdominal sternite III (first visible sternite) with prominent medial lobe. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small field of ventral spatulate setae; mandibles not modified; metafemur relatively slender, with very small subapical tooth on posterior margin; metasternal fovea moderately large, prominent, transversely ovoid, with pencil of long, fine, dense setae, located posterad of middle of metasternum. Median lobe in lateral aspect very robust, strongly curved medially, apical portion slender, curved dorsad, apex pointed (fig. 100); in ventral aspect very broad, apical portion broad, short, narrowed to pointed apex (fig. 99); operculum large, flat, rounded (fig. 99); lateral lobes small, slender, curved, much 2005 69MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II shorter than median lobe, each with 2 long, stout subapical setae (figs. 99, 101). Female tarsi 5-4-4. D ISTRIBUTION : This species is known from eastern North America from Ontario and Manitoba to New Hampshire and south to South Carolina and Tennessee (fig. 104). S PECIMENS E XAMINED : CANADA: Manitoba: Rennie, Lily Pond, Whiteshell Prov. Park, 6 Jun 1984, maple-poplar forest, FIT, S and J Peck (1, PECK). Ontario: Chaffeys Locks QUBS, 2 Sep 1980, litter, on ‘‘fleshy fungi’’, S Peck (1, CNCI). UNITED STATES: New Hampshire: White Mts, woods (1, MCZC); Coos Co.: Jefferson Notch, 19 Sep 1974, 3000 9 , under conifer bark, A Newton, JF Lawrence (1, MCZC); Grafton Co.: Hubbard Brook Exp. Forest, Bear Brook, 15 Oct 1982, leaf litter, DS Chandler (1, DENH). North Carolina: Round Knob, 24 June (1, MCZC); Appalachian Trail to Clingman’s Dome, 19 Aug 1981, on Comatrichia-like slime mold, QD Wheeler (1, QDWC). South Carolina: Oconee Co.: 12 mi NW Walhalla Oconee St Park, 5 Jun 1981, 1700 9 , log-bark litter, 1, S Peck (1, CNCI). Tennessee: Cocke Co.: GSMNP, Albright Grove Tr. 83 8 16 9 45 0 W, 35 8 44 9 10 0 N, 19 Oct 2001, 1000 m, decid stump, Carlton (2, LSAM). Virginia: Giles Co.: Mtn Lake Biol. Sta., 15 Sep 1990, 3700 9 , on Arcyria denudata, SL Stephenson (1, QDWC). D ISCUSSION : This species was described from female specimens only. However, we are confident in the assignment of male specimens to this species since we were able to examine a male and female in the same series. The species is relatively rare in collections, and the longest series of specimens examined was two. Specimens have been collected from June to October. They have been found from under conifer bark and from various litter sources including maple-poplar forest, leaves, logs and bark, and a deciduous stump. Elevation records are from 1700 to 3000 ft. Host records include a ‘‘fleshy fungus’’, Arcyria denudata, and a Comatrichalike slime mold. AGATHIDIUM IOTA SPECIES GROUP D IAGNOSIS : The single species in this group has the anterior portion of the mesosternum very broad (broader than the posterior portion), the mesosternum in the same plane as the metasternum, strong contractility, obsolete humeral angles of the elytra, minute size (TBL , 1.6 mm), and 4-3-3 tarsal formulae in both males and females. Males lack a metasternal fovea, and the posterior margin of the metafemur is unmodified. D ISCUSSION : This species further confounds the already dubious definition of the genus Agathidium. The species possesses a distinctive 3-segmented antennal club, moderately, but distinctly, excavated anterior clypeal margin, and distinctive temporum and supraocular carina extending posterad of the eye. However, it also has the unusual character state of 4-3-3 tarsal formulae in both males and females. The wings, when present, are long and fringed with fine setae, a feature characteristic of many small insects that balloon with the wind. The single species has the anterior portion of the mesosternum very broad, much broader than the posterior portion, which is very short and not at all concave or declivitous. The mesosternum is in the same plane as the metasternum. Also, specimens are very stout and strongly contractile and the metasternum is short. Thus, the species is very similar to members of the A. oniscoides group. However, the species lacks oblique femoral lines on the metasternum and has a particularly unusual tarsal formula, prompting us to place it in its own group. Agathidium iota Miller and Wheeler, new species Figures 102, 103, 359 T YPE M ATERIAL : Holotype, ? in AMNH labeled ‘‘MEXICO: S.L.P. 14 mi. W Xilitla, 4800 ft., Liq. for. VI.29.73 A.Newton/leaf & log litter/HOLOTYPE Agathidium iota Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, San Luis Potosi, 14 mi W Xilitla, 4800 9 . D IAGNOSIS : This is the only species in the group and has the diagnostic characters of the group (see above). D ESCRIPTION : Body minute (TBL 5 1.15– 1.51 mm), rounded (PNW/TBL 5 0.43– 0.53), strongly contractile. Head, pronotum, and elytra red-brown, not iridescent; venter, antennae, palpi, and legs yellow-brown. Head broad (MDL/OHW 5 0.40–0.72), dorsal surface flattened, dorsoventrally com- 70 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY pressed; temporum reduced, not prominent, supraocular carina extending posteriorly to margin of head; surface of head with extremely fine punctures, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes somewhat reduced, dorsoventrally compressed, relatively large-faceted; gula unmodified; clypeus moderately excavate; antennae relatively short (ratios: length I:II:III 5 1.8:1.0:1.0, width VII:VIII:IX 5 1.0:1.0:1.9). Pronotum very large, broad (PNL/PNW 5 0.69–0.80), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with minute, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, slightly elongate, but not apically acuminate (SEL/ELW 5 0.73– 1.47); punctation similar to pronotum; sutural stria indistinctly present apically. Flight wings absent or present, long, narrow, fringed with fine setae. Anterior portion of mesosternum very broad, convex, medial carina prominent anteriorly, obscured posteriorly; posterior portion very short, not concave. Metasternum very short (MTL/MTW 5 0.09–0.14), flattened, medially slightly sloped dorsad anteriorly; oblique femoral carinae absent. Male tarsi 4-4-3; proand mesobasotarsomeres not laterally expanded, with few or no ventral spatulate setae; mandibles not modified; metafemur relatively slender, unmodified; metasternal fovea absent. Median lobe in lateral aspect very broad and robust basally, with subbasal constriction, curved, short, and moderately slender distad of constriction, apical portion narrowed, tapering to long, sharp point (fig. 103); in ventral aspect very broad, lateral margins broadly curved, apical portion broadly triangular, apex broadly pointed (fig. 102); operculum absent (fig. 102); lateral lobes short, slender, glabrous (figs. 102, 103). Female tarsi 4-4-3. E TYMOLOGY : This species is named after the smallest Greek letter iota, for the extremely small size of members of this species. D ISTRIBUTION : This species is known from southern Mexico and Guatemala (fig. 359). P ARATYPES : MEXICO: Guerrero: 10.3 km SW Filo de Caballo, 17 Jul 1992, 2700 m, oak-pinefir forest, leaf log litter, Berlese, R Anderson (1, CNCI); 10.3 km SW Filo de Caballo, 17 Jul 1992, 2700 m, oak-pine-fir forest (wet), leaf-log litter, Berlese, RS Anderson (5, MCZC). Michoacan: 18 Sep 1973, 9400 9 , oak-pine litter, A Newton (2, CNCI). Oaxaca: S. Suchixtepec, 24 Jul 1992, wet riparian alder forest, leaf litter, Berlese, RS Anderson (12, MCZC). Verapaz: 14.5 km S Purulha, 26 May 1991, 1600 m, riparian bottomland, oak forest litter, RS Anderson (2, MCZC). D ISCUSSION :Agathidium iota has been collected from leaf litter in a variety of habitats including pine, oak, and fir mixed forests and wet alder riparian forests. All known specimens were collected from sifting or Berlese samples. Elevation records are from 1600 to 2700 m. This is the smallest species of Agathidiini known. AGATHIDIUM ONISCOIDES SPECIES GROUP D IAGNOSIS : This group is characterized by the relatively horizontal ventral surface of the mesosternum, which is in approximately the same plane as the metasternum (with the posterior portion not concave), the broad anterior portion of the mesosternum which generally has a prominent medial carina, highly contractile body form, very broad pronotum, and broadly rounded posterolateral angles of the pronotum (fig. 105), broadly rounded humeral angles of the elytra, relatively impunctate dorsal surface and presence of distinct, oblique femoral lines on the metasternum which often meet medially in a prominent, posteriorly directed flange. The species range from minute to large, are usually dark redbrown to piceous dorsally, and males often have a distinctive tooth on the posterior margin of the metafemur. D ISCUSSION : This species group corresponds to the subgenus Agathidium sensu stricto auctorum. The group includes a fantastic diversity of relatively similar species. Although a large number of species are described here, it is certainly likely that the true species diversity remains yet to be described. The species are particularly numerous in the eastern United States, particularly the southern Appalachians, and in the mountains of Mexico, where we especially expect the 2005 71MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Fig. 105. Agathidium aztec, dorsal habitus. number of species to increase with additional collection of specimens. Many of the species in this group are apparently cryptic in habitat and are often collected from forest litter samples and Berlese extraction. Often these species have the eyes reduced and are apterous. The strongly contractile nature and ballshaped habitus make these species distinctive. To facilitate management of this large group, the species were arranged into subgroups. Unfortunately, a great many of the species and groups of species are identifiable only from males, and many of these only by 72 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 106–112. Agathidium oniscoides-group species, heads, left lateral aspect: 106, A. vaderi. 107, A. kimberlae.108, A. oculeum.109, A. stenomma.110, A. microphthalmum.111, A. nimbosilva.112, A. chauliodoum. Bars 5 0.5 mm. dissection of genitalia. The key relies on dissection and critical examination of male genitalia, an unavoidable problem because of the state of our knowledge of the group at this time. Other characters of importance are the shape of the male metafemoral tooth, the presence of modifications such as lobes or tubercles on the surface of the gula, the development of the eyes and the nature of the oblique metasternal carinae and male metasternal fovea. KEY TO A. ONISCOIDES SPECIES GROUP 1. Median lobe of aedeagus apically deeply emarginate, with two long apical rami (figs. 174, 178); operculum long, slender, and with conspicuous lateral teeth (figs. 173, 177); eyes reduced to elongate triangles (figs. 106, 107) (A. kimberlae subgroup) ............. 5 — Median lobe of aedeagus apically entire, operculum various, in some cases apically expanded into lateral teeth, but generally without multiple, conspicuous lateral teeth; eyes various .......................... 2 2(1). Median lobe of aedeagus with moderate to very prominent longitudinal, lateral sulcus at base of apical portion which is subtended by two lateral carinae or a single dorsal carina and a lateral carina (e.g., figs. 184, 186); lateral lobes apically variously sinuate and often expanded (e.g., figs. 184, 187), sinuate portion of lobe fitting into lateral sulcus of median lobe of aedeagus; Mexico and Central America (A. aztec subgroup) ............... 6 —Median lobe of aedeagus without prominent lateral sulcus at base of apical portion; lateral lobes straight or evenly curved apically; distribution various ........ 3 3(2). Operculum of median lobe of aedeagus completely divided into two long rami (e.g., fig. 309); median lobe of aedeagus in ventral aspect with lateral margins distinctly sinuate near base (e.g., fig. 309); metasternum very short medially (MTL/ MTW 5 0.08–0.17); male metafemoral tooth large and prominent, subapical, and sharply pointed; eastern United States (A. dentigerum subgroup) ............ 40 —Operculum of median lobe of aedeagus sometimes apically emarginate, in some species relatively deeply so, but never entirely divided to base; median lobe of aedeagus in ventral aspect with lateral margins not sinuate near base; metasternum various, but generally longer (MTL/ MTW 5 0.12–0.28); male metafemoral tooth various, from reduced to a small, apical tooth on the posterior margin to a relatively large subapical tooth; distribution various ..................... 4 4(3). Eyes extremely reduced to tiny ovals or minute triangles consisting of only a few facets (figs. 110–112); Mexico and Central America (A. microphthalmum subgroup) ........................ 50 —Eyes relatively large and multifaceted to somewhat dorsoventrally compressed but not strongly reduced to tiny ovals or minute triangles; United States and Canada (A. oniscoides subgroup) .......... 52 5(1) (A. kimberlae subgroup). Apical rami of median lobe of aedeagus apically expanded and truncate (fig. 173); operculum long, extending nearly to apex of median lobe of aedeagus (fig. 173); male metafemoral tooth large and metafemur very broad (fig. 120); eyes very strongly re- 2005 73MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Figs. 113–117. Agathidium oniscoides-group species, sterna and legs: 113, A. gomezae, metasternum, arrow indicating medial metasternal lobe. 114, A. skoliosternum, mesosternum, arrow indicating anterior excavation of mesosternal margin. 115, A. tribulosum, ? left proleg, arrow indicating spine on ? left protrochanter. 116, A. tribulograndum, ? left proleg, arrow indicating spine on ? left procoxa. 117, A. framea, ? left metaleg, arrow indicating broad lobe along anterior margin of ? left metacoxa. duced (fig. 106) ................... ..............A. vaderi, new species — Apical rami of median lobe of aedeagus apically slender, pointed, or narrowly rounded and slightly convergent (fig. 177); operculum shorter, ending distinctly short of apex of median lobe of aedeagus (fig. 177); male metafemoral tooth smaller and metafemur narrower (fig. 119); eyes strongly reduced (fig. 107) ......... ........... A. kimberlae, new species 6(2) (A. aztec subgroup). Gula with two prominent tubercles, one on each side of midline; male metafemur with large, curved apical tooth (fig. 121) .............. ........ A. bituberculum, new species — Gula without two tubercles or with only a single, medial tubercle; male metafemur various ......................... 7 7(6). Anteromedial margin of metasternum with a transverse line which forms a distinct posteriorly directed lobe medially on metasternal surface (fig. 113) ......... 8 — Anteromedial margin of metasternum with transverse line straight medially, not forming a medial metasternal lobe . . 10 8(7). Operculum of median lobe of aedeagus shallowly emarginate apically (fig. 184) .............A. oedema, new species — Operculum of median lobe of aedeagus very deeply emarginate apically (figs. 188, 192) ....................... 9 9(8). Lobe formed by sinuate line on metasternum swollen, forming a prominent tumidity; male metafemoral tooth broad, rounded, obsolete (fig. 123); apical portion of median lobe of aedeagus relatively narrow (fig. 189) .................... ............ A. gomezae, new species — Lobe formed by sinuate line on metasternum not swollen, flattened; male metafemoral tooth small but distinct and acutely 80 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 232–260. Agathidium oniscoides-group species, aedeagus: 232–235, A. tribulosum:232, ventral; 233, median lobe, apex, ventral; 234, median lobe, lateral; 235, left lateral lobe, lateral. 236–239, A. tribulograndum:236, ventral; 237, median lobe, apex, ventral; 238, median lobe, lateral; 239, left lateral lobe, lateral. 240–243, A. invisitatum:240, ventral; 241, median lobe, apex, ventral; 242, median lobe, lateral; 243, left lateral lobe, lateral. 244–247, A. multidentatum:244, ventral; 245, median lobe, 2005 81MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II ← apex, ventral; 246, median lobe, lateral; 247, left lateral lobe, lateral. 248–250, A. sejunctum:248, ventral; 249, median lobe, lateral; 250, left lateral lobe, lateral. 251–253, A. grandidentatum:251, ventral; 252, median lobe, lateral; 253, left lateral lobe, lateral. 254–256, A. andersoni:254, ventral; 255, median lobe, lateral; 256, left lateral lobe, lateral. 257–260, A. disgregum:257, ventral; 258, median lobe, apex, ventral; 259, median lobe, lateral; 260, left lateral lobe, lateral. 30(29). Apex of median lobe of aedeagus distinctly recurved in lateral aspect, apically pointed and abruptly hooked ventrad ........................ 31 — Apex of median lobe of aedeagus not distinctly recurved in lateral aspect, apically straight or evenly curved and in some species angled dorsad or ventrad ............................ 32 31(30). Operculum of median lobe of aedeagus short, narrow basally, apically relatively broadly expanded and emarginate with each ramus broad and apically pointed (fig. 264); eyes prominent and multifaceted (fig. 108) ............ .......... A. oculeum, new species — Operculum of median lobe of aedeagus long, relatively slender, apex emarginate with each ramus narrowly rounded apically (fig. 267); eyes dorsoventrally compressed, reduced ............. ........A. recurvatum, new species 32(31). Operculum of median lobe of aedeagus relatively short and broad, length about 2 3 greatest width ............. 33 — Operculum of male long and slender, about 2.5–4 3 greatest width ..... 35 33(32). Size large (TBL . 4.0 mm); apical portion of median lobe of aedeagus expanded medially along ventral margin in lateral aspect (fig. 272) ........ ......... A. impensum, new species — Size smaller (TBL , 4.0 mm); apical portion of median lobe of aedeagus not expanded in lateral aspect ....... 34 34(33). Apex of median lobe of aedeagus in lateral aspect slender, curved slightly dorsad, apex narrowly rounded (fig. 275) ........... A. cheneyi, new species — Apex of median lobe of aedeagus in lateral aspect broad, not curved dorsad, apex broadly rounded (fig. 278) ..... ....... A. tenangoense, new species 35(32). Apex of operculum of median lobe of aedeagus in ventral aspect with apex laterally flared into relatively prominent, pointed processes ............. 36 — Apex of operculum of median lobe of aedeagus in ventral aspect with apex with lateral margins not flared or produced ............................ 37 36(35). Apex of median lobe of aedeagus in lateral aspect distinctly expanded medially along ventral margin (fig. 282); operculum in lateral aspect apically curved ventrad, expanded apically (fig. 282) ..... A. rumsfeldi, new species — Apex of median lobe of aedeagus in lateral aspect evenly narrowed to pointed apex (fig. 286); operculum in lateral aspect straight to narrowly rounded apex (fig. 286) . . A. hirsutum, new species 37(35). Apex of operculum of median lobe of aedeagus in lateral aspect broad and hooked dorsad (fig. 290); apex of median lobe in ventral aspect very narrow, straight, apically very narrowly rounded (fig. 289) ................... ........... A. cortezi, new species — Apex of operculum of median lobe of aedeagus in lateral aspect apically not broad, not hooked dorsad; apex of median lobe in ventral aspect not as strongly narrowed, apically broadly rounded ..................... 38 38(37). Gula medially with a broad, prominent tumidity ....................... ....... A. tumidiventre, new species — Gula unmodified ................ 39 39(38). Apical portion of median lobe of aedeagus in lateral aspect expanded along ventral margin (fig. 297); operculum of median lobe shorter and slightly broader, ending distinctly short of apex of median lobe (fig. 296) ........... .............A. aztec, new species — Apical portion of median lobe of aedeagus in lateral aspect not expanded along ventral margin (fig. 301); operculum of median lobe longer and more slender, extending nearly to apex of median lobe (fig. 299) ........... ......... A. iridescens, new species 40(3) (A. dentigerum subgroup). Gula with small but prominent medial lobe on gula ........................ 41 — Gula unmodified, without medial lobe . . ............................ 43 82 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 261–287. Agathidium oniscoides-group species, aedeagus: 261–263, A. oaxacaense:261, ventral; 262, median lobe, lateral; 263, left lateral lobe, lateral. 264–266, A. oculeum:264, ventral; 265, median lobe, lateral; 266, left lateral lobe, lateral. 267–269, A. recurvatum:267, ventral; 268, median lobe, lateral; 269, left lateral lobe, lateral. 270–272, A. impensum:270, ventral; 271, median lobe, lateral; 272, left lateral lobe, lateral. 273–276, A. cheneyi:273, ventral; 274, median lobe, apex, ventral; 275, median lobe, lateral; 276, left lateral lobe, lateral. 277–279, A. tenangoense:277, ventral; 278, median lobe, lateral; 279, left lateral lobe, lateral. 280–283, A. rumsfeldi:280, ventral; 281, median lobe, apex, ventral; 282, median lobe, lateral; 283, left lateral lobe, lateral. 284–287, A. hirsutum:284, ventral; 285, median lobe, apex, ventral; 286, median lobe, lateral; 287, left lateral lobe, lateral. 2005 83MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Figs. 288–302. Agathidium oniscoides-group species, aedeagus: 288–291, A. cortezi:288, ventral; 289, median lobe, apex, ventral; 290, median lobe, lateral; 291, left lateral lobe, lateral. 292–295, A. tumidiventre:292, ventral; 293, median lobe, apex, ventral; 294, median lobe, lateral; 295, left lateral lobe, lateral. 296–298, A. aztec:296, ventral; 297, median lobe, lateral; 298, left lateral lobe, lateral. 299–302, A. iridescens:299, ventral; 300, median lobe, apex, ventral; 301, median lobe, lateral; 302, left lateral lobe, lateral. 41(40). Apical portion of median lobe of aedeagus strongly and broadly sinuate in lateral aspect (fig. 305) ............. .............. A. dentigerum Horn — Apical portion of median lobe of aedeagus straight or with only apex of median lobe abruptly sinuate in lateral aspect ........................ 42 42(41). Apical portion of median lobe of aedeagus straight to apex in lateral aspect (fig. 308) . . A. stephani, new species — Apical portion of median lobe of aedeagus with apex abruptly sinuate in lateral aspect (fig. 310) ............. ....... A. akrogeneios, new species 43(42). Posteromedial margin of male metacoxa with broad, rounded, prominent lobe (fig. 117); apical portion of median lobe of aedeagus in lateral aspect strongly curved dorsad (fig. 312), in ventral aspect with apex distinctly arrowhead-shaped (fig. 311) ........ ........... A. framea, new species — Posteromedial margin of male metacoxa not modified; apical portion of median lobe of aedeagus variously curved or straight in lateral aspect, in ventral aspect not arrowhead-shaped ...... 44 44(43). Size very small (TBL , 2.2 mm) ; median lobe of aedeagus with operculum 84 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 303–328. Agathidium oniscoides-group species, aedeagus: 303–305, A. dentigerum:303, ventral; 304, median lobe, lateral; 305, left lateral lobe, lateral. 306–308, A. stephani:306, ventral; 307, median lobe, lateral; 308, left lateral lobe, lateral. 309–310, A. akrogeneios:309, ventral; 310, lateral. 311–313, A. framea:311, ventral; 312, median lobe, lateral; 313, left lateral lobe, lateral. 314–316, A. appalachium:314, ventral; 315, median lobe, lateral; 316, left lateral lobe, lateral. 317–319, A. pocahontasae:317, ventral; 318, median lobe, lateral; 319, left lateral lobe, lateral. 320–322, A. carolinense: 320, ventral; 321, median lobe, lateral; 322, left lateral lobe, lateral. 323–325, A. gallititillo:323, ventral; 324, median lobe, lateral; 325, left lateral lobe, lateral. 326–328, A. divaricatum:326, ventral; 327, median lobe, lateral; 328, left lateral lobe, lateral. 2005 85MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II very short, with rami straight and not apically expanded (figs. 314, 317) . . . ............................ 45 — Size larger (TBL . 2.2 mm) ; median lobe of aedeagus with operculum longer, rami of some species apically expanded ...................... 46 45(44). Apex of median lobe of aedeagus curved dorsad in lateral aspect (fig. 315), in ventral aspect apically truncate (fig. 314) . . A. appalachium, new species — Apex of median lobe of aedeagus straight in lateral aspect (fig. 318), in ventral aspect apically pointed (fig. 317) .... ...... A. pocahontasae, new species 46(44). Apical portion of median lobe of aedeagus in lateral aspect strongly curved dorsad (fig. 321), in ventral aspect slender and evenly tapered to narrowly rounded apex (fig. 320) .......... ........A. carolinense, new species — Apical portion of median lobe of aedeagus in lateral aspect not strongly curved dorsad, in ventral aspect various, generally somewhat expanded medially, but not evenly tapered to narrowly rounded apex ........... 47 47(46). Median lobe of aedeagus in lateral aspect distinctly bent ventrad at base of apical portion (fig. 324) ................ ..........A. gallititillo, new species — Median lobe of aedeagus in lateral aspect only slightly or not bent ventrad at base of apical portion .............. 48 48(47). Apical portion of median lobe of aedeagus abruptly narrowed near base, lateral margins subparallel thereafter (fig. 326); rami of operculum apically expanded, rounded, and divergent, extending nearly to apex of median lobe (fig. 326) ...................... ....... A. divaricatum, new species —Apical portion of median lobe of aedeagus not abruptly narrowed near base in ventral aspect, lateral margins evenly convergent or slightly constricted basally, operculum not apically expanded, rounded, or divergent, not extending nearly to apex of median lobe . . . ............................ 49 49(48). Apical portion of median lobe of aedeagus with margins evenly convergent to apex (fig. 329), in lateral aspect straight to apex (fig. 330) ............... ............ A. bushi, new species —Apical portion of median lobe of aedeagus distinctly constricted near base (fig. 332), in lateral aspect distinctly curved dorsad at apex (fig. 333) .......... ....... A. georgiaense, new species 50(4) (A. microphthalmum subgroup). Eyes ovoid, dorsally directed (fig. 110); median lobe of aedeagus stout, apically abruptly curved and sharply pointed in ventral aspect (fig. 335) .......... ...... A. chauliodoum, new species —Eyes shaped like elongate triangles, more laterally directed (figs. 111, 112); median lobe of aedeagus various . . . 51 51(50). Median lobe of aedeagus with apex abruptly expanded and truncate (fig. 338) .......................... ... A. microphthalmum, new species —Median lobe of aedeagus with apex pointed, not expanded (fig. 341) ..... ........ A. nimbosilva, new species 52(4) (A. oniscoides subgroup). Median lobe of aedeagus with operculum long, slender, evenly tapered, with apex laterally expanded ...................... 53 —Median lobe of aedeagus with operculum shorter and broader, apex not expanded in most species ............... 54 53(52). Size generally larger (TBL 5 2.5–5.0 mm) ; dorsal surfaces black; metasternal fovea moderately large and conspicuous ....................... ... A. oniscoides Palisot de Beauvois —Size smaller (TBL 5 2.2–2.9 mm) ; dorsal surfaces red; metasternal fovea minute, inconspicuous .............. .................A. rubellum Fall 54(52). Lateral lobe with apical portion directed at angle dorsad in lateral aspect with subapical expansion on dorsal margin (fig. 352) . . . A. exiguum Melsheimer —Lateral lobe with apical portion various but without subapical dorsal expansion in lateral aspect (fig. 355) ........ ......... A. fawcettae, new species Agathidium kimberlae Subgroup D ISCUSSION : This group of two species is united by the median lobe of aedeagus apically bifid (figs. 174, 178) and with the operculum strongly toothed (figs. 173, 177). In addition, the eyes are strongly reduced (figs. 106, 107), and the two share a similar habitus since they are moderately elongate with a large, protruding pronotum and strong contractility. The male metafemora are very broad (figs. 119, 120). The two species occur sympatrically in a portion of the southern 86 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 329–356. Agathidium oniscoides-group species, aedeagus: 329–313, A. bushi:329, ventral; 330, median lobe, lateral; 331, left lateral lobe, lateral. 332–334, A. georgiaense:332, ventral; 333, median lobe, lateral; 334, left lateral lobe, lateral. 335–337, A. chauliodoum:335, ventral; 336, median lobe, lateral; 337, left lateral lobe, lateral. 338–340, A. microphthalmum:338, ventral; 339, median lobe, lateral; 340, left lateral lobe, lateral. 341–343, A. nimbosilva:341, ventral; 342, median lobe, lateral; 343, left lateral lobe, lateral. 344–347, A. oniscoides:344, ventral; 345, apex of median lobe, ventral, specimen from Macon City, NC; 346, median lobe, lateral; 347, left lateral lobe, lateral. 348–350, A. rubellum:348, ventral; 349, median lobe, lateral; 350, left lateral lobe, lateral. 351–353, A. exiguum: 351, ventral; 352, median lobe, lateral; 353, left lateral lobe, lateral. 354–356, A. fawcettae:354, ventral; 355, median lobe, lateral; 356, left lateral lobe, lateral. 2005 87MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Figs. 357, 358. Geographic distribution of Agathidium oniscoides-group species: 357, A. vaderi. 358, A. kimberlae. Appalachians where they are commonly collected from forest litter. Agathidium vaderi Miller and Wheeler, new species Frontispiece, figures 106, 120, 173–176, 357 T YPE M ATERIAL : Holotype, ? in AMNH labeled ‘‘NC/TN: Clingman’s Dome Gr. Smoky Mtn. Nat. Park 18.viii.81 sift. decaying wood Q. Wheeler #81343/HOLOTYPE Agathidium vaderi Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : United States, North Carolina, Great Smoky Mountains National Park, Clingman’s Dome. D IAGNOSIS : This species is distinguishable from most by the elongate form of the body combined with highly reduced eyes consisting of small, elongate triangles (frontispiece, fig. 106) and by the median lobe of the aedeagus being apically deeply emarginate (fig. 174). Agathidium vaderi is most similar to A. kimberlae from which it differs in having the apical rami of the median lobe broad, apically truncate, and with the apices straight (fig. 174), whereas in A. kimberlae the rami are narrow, apically narrowly rounded, and with the apices convergent (fig. 178). The operculum in A. kimberlae is apically more broadly expanded, shorter (ending far short of apex of median lobe), and bears fewer teeth in general (fig. 177), whereas in A. vaderi the operculum extends nearly to the apex of the median lobe and is heavily toothed (fig. 173). The metafemur is slightly narrower (fig. 119), and the eyes are slightly larger (fig. 107) in A. kimberlae than in A. vaderi (figs. 106, 120). D ESCRIPTION : Body moderately large (TBL 88 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 5 2.59–3.24 mm), oblong (PNW/TBL 5 0.38–0.45), strongly contractile. Head piceous to reddish-brown; pronotum piceous to reddish-brown, lighter reddish along posterior margin; elytra piceous to reddish-brown, lighter apically; venter brown to yellow-brown; antennae and palpi yellow to yellow-brown; legs yellow-brown. Head broad (MDL/OHW 5 0.53–0.78), dorsoventrally conspicuously compressed, laterally with distinct bead, tentorium laterally protuberant; with few, sparse, fine punctures, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eye strongly reduced to an elongate triangle (fig. 106); gula concave; antennomere ratios: length I:II:III 5 2.0:1.0:1.7, width VII:VIII:IX 5 1.0:1.0:1.5. Pronotum very large, broad (PNL/PNW 5 0.72–0.92), strongly convex, extending anteromedially over head, posterior margin weakly bisinuate, lateral margin broadly curved, not angulate; with few, fine, scattered punctures, surface between punctures shiny, smooth. Elytra broad, convex, apically not strongly attenuate, elongate (SEL/ELW 5 0.96–1.17); with very few fine, scattered punctures, surface between punctures shiny, smooth; without sutural stria. Flight wings absent. Mesosternum broad, medially with distinct longitudinal carina. Metasternum moderately broad, not reduced (MTL/MTW 5 0.17– 0.26); oblique femoral lines weakly present, meeting medially in low line. Male tarsi 5-5-4; proand mesobasotarsomeres moderately laterally expanded, with large ventral field of spatulate setae; mandibles not modified; metafemur very broad, posteroapical angle broadly triangular and strongly produced in large, flat tooth, apical margin strongly curved (fig. 120); metasternal fovea present, medial, circular and small with dense pencil of fine setae. Median lobe in lateral aspect moderately curved, apical portion straight and with margins approximately parallel, apex narrowly rounded (fig. 175); in ventral aspect slender, apically bifurcate, rami somewhat divergent, each ramus apically expanded and broadly truncate, medial emargination moderately broad (figs. 173, 174); operculum long, extending to or nearly to apex of median lobe, with conspicuous and strong lateral dentation along entire length, apex abruptly laterally expanded, curved toward apex of median lobe, extending nearly to apex of median lobe (fig. 173); lateral lobes slender, long, apically narrowly rounded with two long setae (fig. 176). Female tarsi 5-4-4. E TYMOLOGY : This species is named for the fictional villain Darth Vader in the movie ‘‘Star Wars’’, who shares with A. vaderi a broad, shiny, helmetlike head. D ISTRIBUTION : This species is known from mountainous areas of North Carolina, Tennessee, and Georgia (fig. 357). P ARATYPES : UNITED STATES: Georgia: White-Union Co.: Tesnatee Gap, 20 Jul 1967, 3100 9 , log litter, Berlese, S Peck, A Fiske (13, PECK); Tesnatee Gap, 20 Jul 1967, log litter, Berlese, S Peck, A Fiske (1, CNCI). North Carolina: Clingman’s Dome, Great Smoky Mts Natl Park, 18 Aug 1981, decaying wood, Q Wheeler (1, QDWC); Highlands, Jun 1888 (1, MCZC); Cherokee Co.: Joanna Bald, 26 Jul 1967, 4700 9 , log litter, Berlese, S Peck, A Fiske (2, CNCI); Macon Co.: 5 mi NW Highlands, California Gap, 3000– 3500 9 , leaf litter, Rhododendron, Q Wheeler (1, QDWC); Wayah Bald, 5 Jun 1981, hardwood litter, Q Wheeler (1, QDWC); Highlands, Jul, QD Wheeler (1, QDWC); Swain Co.: Great Smoky Mts Natl Park, Flat Creek Trail, 83 8 10 9 21 0 W, 35 8 33 9 1 0 N, 31 Jul 2001, 1500 m, leaf litter, Berlese, C Carlton (8, LSAM). Tennessee: Great Smoky Mts P.P., 19 Jun 1981, Q Wheeler (1, QDWC); Great Smoky Mts NP, 19 Jun 1981, leaf litter, Q Wheeler (1, QDWC); Blount Co.: Great Smoky Mts Natl Park, Cades Cove, 8 Jun 1960, 1700 9 , wet leaf duff, J Wagner, W Suter (4, FMNH); Cocke Co.: Great Smoky Mts Natl Park, Albright Grove Trail, 83 8 16 9 45 0 W, 35 8 44 9 10 0 N, 29 Jun 2001, 1000 m, old growth, Berlese, C Carlton (1, LSAM); Sevier Co.: Great Smoky Mts Nat Park, Newfound Gap, 16 Sep 1949, J Wagner, W Suter (3, FMNH); Great Smoky Mts Natl Park, EElkmont, 8 Jun 1960, 2250 9 , leaf duff, W Suter, J Wagner (5, FMNH); Appalachian Trail at beech Gap on Clingman’s Dome Rd, 83 8 26 9 50 0 W, 35 8 36 9 36 0 N, 28 Jun 2001, 1750 m, forest litter, Berlese, A Tischckhin, V Moseley (3, LSAM); Great Smoky Mts Natl Park, 0.5 km NE Newfound Gap, 82 8 24 9 46 0 W, 35 8 38 9 9 0 N, 26 Jun 2001, CE Carlton (7, LSAM). D ISCUSSION : This species has been collected mainly from log and leaf litter (particularly Rhododendron litter). Altitude records are from 1700 to 4700 ft. 2005 89MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Agathidium kimberlae Miller and Wheeler, new species Figures 107, 119, 177–180, 358 T YPE M ATERIAL : Holotype, ? in AMNH labeled ‘‘NC: Macon Co. Highlands, 2 mi NW Nantahala NF 7.VIII.81/ex. leaf litter Q.D.Wheeler Lot. #81231 J. Pakaluk, Q.Wheeler/HOLOTYPE Agathidium kimberlae Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : United States, North Carolina, Macon Co., Nantahala National Forest, 2 mi NW Highlands. D IAGNOSIS : This species is distinguishable from most by the elongate form of the body combined with highly reduced eyes consisting of small, elongate triangles (fig. 107) and by the median lobe of the aedeagus being apically deeply emarginate (fig. 178). Agathidium kimberlae is most similar to A. vaderi. See the diagnosis of that species for characters to separate the two species. D ESCRIPTION : Body moderately small (TBL 5 2.38–3.04 mm), broad, oblong (PNW/TBL 5 0.40–0.47), strongly contractile. Head reddish-brown; pronotum reddishbrown, lighter reddish along posterior margin; elytra reddish-brown, lighter apically; venter brown to yellow-brown; antennae and palpi yellow to yellow-brown; legs yellowbrown. Head broad (MDL/OHW 5 0.55–0.77), dorsoventrally conspicuously compressed, laterally with distinct bead, tentorium laterally protuberant; with few, sparse, fine punctures, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eye strongly reduced to an elongate triangle (fig. 107); gula strongly concave; antennomere ratios: length I:II:III 5 2.0:1.0: 1.8, width VII:VIII:IX 5 1.0:1.0:1.7. Pronotum very large, broad (PNL/PNW 5 0.71– 0.85), strongly convex, extending anteromedially over head, posterior margin weakly bisinuate, lateral margin broadly curved, not angulate; with few, fine, scattered punctures, surface between punctures shiny, smooth. Elytra broad, elongate, convex, apically not strongly attenuate (SEL/ELW 5 1.08–1.31); with very few fine, scattered punctures, surface between punctures shiny, smooth; without sutural stria. Flight wings reduced to absent. Mesosternum broad, medially with distinct longitudinal carina. Metasternum short (MTL/MTW 5 0.13–0.19); oblique femoral lines weakly present, meeting medially in low line. Male tarsi 5-5-4; proand mesobasotarsomeres distinctly laterally expanded, with small field of ventral adhesive setae; mandibles not modified; metafemur broad, expanded anteriorly, posteroapical angle triangular and posteriorly produced into large, broad tooth, apical margin strongly curved (fig. 119); metasternal fovea present, very small to moderately large, medial, circular, and with dense pencil of fine setae. Median lobe in lateral aspect moderately curved, apical portion straight and with margins approximately parallel, apex narrowly rounded (fig. 179); in ventral aspect slender, apically bifurcate, rami slender, curved, convergent apically, apices very narrowly rounded, medial emargination moderately broad (figs. 177, 178); operculum moderately long, not extending to apex of median lobe, with conspicuous and strong lateral dentation along entire length, apex abruptly laterally expanded, curved toward apex of median lobe, ending distinctly before apex of median lobe (fig. 177); lateral lobes slender, long, apically narrowly rounded with two long setae (fig. 180). Female tarsi 5-4-4. E TYMOLOGY : This species is named in honor of the junior author’s former wife, Kimberly Jo Peters Wheeler, for her understanding and support of his taxonomic ways for a quarter century of marriage. D ISTRIBUTION : This species is found in the southern Appalachian Mountains of North Carolina, Georgia, Tennessee, and Virginia (fig. 358). P ARATYPES : UNITED STATES: Georgia: Clayton, 2000–3700 9 (2, CNCI); Rabun Co.: Rabun Bald, 2 Aug 1981, 4080 9 , leaf litter, J Pakaluk (8, QDWC); Chatahoochee Natl Forest 10 mi NE Clayton, 12 Jun 1981, 2400 9 , leaf litter, J Pakaluk (1, QDWC); Rabun Bald, 11 Aug 1981, 4000 9 , hardwood leaf litter, J Pakaluk and Q Wheeler (1, QDWC); Rabun Bald, 25 Jun 1975, 4300 9 , forest litter, JF Cornell (3, CUIC); Rabun Bald, 2 Aug 1981, leaf litter, QD Wheeler (4, QDWC); Rabun Bald, 11 Aug 1981, 4000 9 , leaf litter, Berlese, QD 96 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.95–1.03); punctation and surface similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.12– 0.20), flat medially, distinctly dorsally sloped anteriorly, with distinct curved line medially on surface; oblique femoral carinae moderately well developed, meeting medially in low carina. Male tarsi 5-5-4; proand mesobasotarsomeres distinctly laterally expanded, protarsomeres more so and with large ventral field of spatulate setae; mandibles not modified; metafemur broad, with small, sharp tooth submedially on posterior margin (fig. 124); metasternal fovea minute, with small pencil of fine setae. Median lobe in lateral aspect long, slender, strongly curved basally, apicomedially with lateral, lobelike carinate expansions between which fit the lateral lobes, apical portion prominent, moderately broad, straight, subapically expanded slightly, apex a sharply curved point (fig. 194); in ventral aspect moderately slender, lateral lobelike carina prominent, apical portion slender medially, apex prominently expanded, apically rounded (figs. 192, 193); operculum in lateral aspect long, slender, prominently expanded and curved at apex, in ventral aspect broad, lateral margins curved, with deep, narrow medial emargination, each ramus somewhat expanded apically and rounded (fig. 192); lateral lobes long, slender basally and curved, medially and apically expanded, sinuate with 2 stout setae (fig. 195). Female tarsi 5-4-4. E TYMOLOGY : Named after the type locality of this species. D ISTRIBUTION : This species is known only from the type locality in Hidalgo (fig. 360). P ARATYPES : MEXICO: Hidalgo: Tlanchinol, 43 km SW Huejutla, 14 Jun 1983, 1500 m, cloud forest litter, S and J Peck (3, PECK). D ISCUSSION : The type specimens were collected from cloud forest litter at 1500 m. Agathidium skoliosternum Miller and Wheeler, new species Figures 114, 125, 196–198, 360 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEX:Ver;7 km E Huatusco 22.VI.83,1040 m Anderson & Peck cloud forest litter/HOLOTYPE Agathidium skoliosternum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Veracruz, 7 km E Huatusco, 1040 m. D IAGNOSIS : This species is immediately distinguishable from all A. oniscoides group species by the modified anterior margin of the mesosternum which is deeply excavated on each side of the midline, making the anterior margin strongly sinuate (fig. 114). This occurs in both males and females, although in females the excavations are not as deep. In other respects this species strongly resembles A. aztec. D ESCRIPTION : Body moderately large (TBL 5 2.93–3.22 mm), broad, robust (PNW/TBL 5 0.48–0.49), strongly contractile. Head and pronotum testaceous; elytra testaceous, not iridescent; venter yellow-brown, antennae, palpi, and legs yellow to yellowbrown. Head broad (MDL/OHW 5 0.59–0.60), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes prominent, not strongly compressed; gula slightly concave; antennomere ratios: length I:II:III 5 1.6:1.0:1.6, width VII:VIII:IX 5 1.0:1.0:1.6. Pronotum very large, broad (PNL/PNW 5 0.73–0.78), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.87–0.99); punctation similar to pronotum; sutural stria present only at apex of elytron. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous, anterior margin strongly excavated on each side, medially with anteriorly directed prominence that is apically truncate (fig. 114); with longitudinal carina present 2005 97MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II only posteriorly. Metasternum moderately narrow (MTL/MTW 5 0.13), flattened, sloping slightly dorsad anteriorly; oblique femoral carinae well developed, meeting medially in low, posteriorly directed lobe. Male tarsi 5-5-4; proand mesobasotarsomeres somewhat laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur relatively narrow, with small subapical tooth along posterior margin, apical margin rounded (fig. 125); metasternal fovea transversely linear, broad, with dense line of fine setae. Median lobe in lateral aspect robust, medially expanded, dorsal margin sinuate, apex slender, slightly curved dorsally, slightly expanded medially along ventral margin, apex slender and pointed (fig. 197); in ventral aspect slender, moderately elongate, medially expanded laterally, apex slender to relatively broadly expanded, arrowhead-shaped apex (fig. 196); operculum long and broad, not extending to apex of median lobe, lateral margins slightly curved, apex bifid, each ramus triangularly pointed, medial emargination narrow (fig. 196); lateral lobe in lateral aspect long, slender, medially slightly expanded and sinuate (fig. 198). Female tarsi 5-4-4; sinuation of anterior margin of mesosternum less strongly pronounced than in male. E TYMOLOGY : This species is named from the Greek words skolios, meaning ‘‘curved’’, and sternum, meaning ‘‘chest’’, for the strongly excavated and bisinuate anterior margin of the mesosternum in this species. D ISTRIBUTION : This species is known from Puebla and Vera Cruz (fig. 360). P ARATYPES : MEXICO: Puebla: 24 km N Xicotepec de Juarez, 17 Jun 1983, oak forest litter, R Anderson (4, PECK); 4.4 mi SW Huachinango, 25 Jul 1969, 1700 m, moist ravine oak forest, malt traps, S and J Peck (1, PECK). D ISCUSSION : This species has been collected from oak forest litter, cloud forest litter, and moist ravine oak forest. Elevation records are from 1040 to 1700 m. Agathidium erythromelas Miller and Wheeler, new species Figures 126, 199–202, 361 T YPE M ATERIAL : Holotype, ? in EMEC labeled ‘‘Morelia, 34 mi E., Michoacan, Mex.VII-2-63/J.Doyen Collector/HOLOTYPE Agathidium erythromelas Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Michoacan, 34 mi E Morelia. D IAGNOSIS : This species is relatively large and rather conspicuously reddish on the head and pronotum with contrasting black on the elytra. The eyes are relatively large and finely faceted. The metasternum is moderately broad and the oblique metasternal carinae are poorly developed. The male metafemoral tooth is small and anteapical (fig. 126). The male genitalia are distinct with the median lobe slender and strongly curved medially and strongly expanded along dorsal and ventral margins at the base of the apical portion in lateral aspect (fig. 201). D ESCRIPTION : Body large (TBL 5 3.47– 3.86 mm), broad, robust (PNW/TBL 5 0.47– 0.48), strongly contractile. Head and pronotum red; elytra piceous, red apically and along lateral margin, slightly blue-iridescent laterally; venter red, testaceous on metasternum; antennae, palpi, and legs yellow-red. Head broad (MDL/OHW 5 0.58–0.67), dorsal surface flattened, dorsoventrally compressed; impunctate, smooth, shiny; frontoclypeal suture obsolete medially; eyes prominent, not compressed; gula slightly concave; antennomere ratios: length I:II:III 5 2.6:1.0: 2.3, width VII:VIII:IX 5 1.0:1.0:2.2. Pronotum very large, broad (PNL/PNW 5 0.66– 0.71), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.05–1.06); punctation similar to pronotum; sutural stria present in apical one-third. Flight wings fully developed. Mesosternum broad, somewhat convex; medial carina prominent. Metasternum narrow (MTL/MTW 5 0.16–0.17), flattened, evenly sloping dorsad anteriorly; oblique femoral carinae not prominent, obsolete medially. Male tarsi 5-5-4; proand mesobasotarsomeres distinctly laterally expanded, protarsomeres more so and with large ventral field 98 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 361, 362. Geographic distribution of Agathidium oniscoides-group species: 361, A. erythromelas 5v ;A. rhamphastes 5m ;A. triangularum 5m .362, A. grumum 5m ;A. lobosternum 5m ; A. megoniscoides 5. . 2005 99MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II of spatulate setae; mandibles not modified; metafemur broad, with small tooth about one-third length from apex (fig. 126); metasternal fovea small, medial, transversely oval, with dense pencil of fine setae. Median lobe in lateral aspect slender in basal half, strongly curved, broadly expanded along dorsal and ventral margins submedially, with prominent lateral sulcus and associated carinae for reception of lateral lobe, apical portion of median lobe slender, evenly curved ventrally, apex pointed (fig. 201); in ventral aspect slender basally, broadly laterally expanded medially, apical portion evenly narrowed basally, then abruptly narrowed, apex a long slender process (figs. 199, 200); operculum in lateral aspect very slender, apically narrowly rounded, in ventral aspect long, slender, lateral margins gently curved, apex narrowly emarginate (fig. 199); lateral lobes very slender, strongly curved, expanded submedially and slightly subapically, broadly sinuate in apical half each with 2 stout subapical setae (fig. 202). Female not examined. E TYMOLOGY : This species is named from the Greek words erythro, meaning ‘‘red’’, and melas, meaning ‘‘black’’, for the red head and pronotum and contrasting black elytron in this species. D ISTRIBUTION : This species is known only from southern Mexico (fig. 361). P ARATYPES : MEXICO: Mexico: Temascaltepec, Real de Arriba, Jul 1934, 6000–7000 9 (2, BMNH); 7 air km WSW Juchitepec, 24 Aug 1987, 2750 m, JT Doyen (3, EMEC). Michoacan: 34 mi E Morelia, 2 Jul 1963, J Doyen (2, EMEC); 34 mi E Morelia, 2 Jul 1963, J Doyen (9, EMEC). Vera Cruz: 10 mi S Las Vigas, 15 Jul 1971, 10,000 9 , under bark of Pinus, A Newton (1, FMNH). D ISCUSSION :Agathidium erythromelas has been collected from under pine bark from 6000 to 10,000 ft. Agathidium rhamphastes Miller and Wheeler, new species Figures 127, 203–205, 361 T YPE M ATERIAL : Holotype, ? in AMNH labeled ‘‘14 mi SW El Salto, Dg. MEX. VI.20 ’64 E.E.Lindquist, coll./bunch grass & sod below [handwritten]/HOLOTYPE Agathidium rhamphastes Miller and Wheeler, 2003 [red label with black line border]’’. Only the holotype was examined of this species. The specimen is in several pieces glued to a point. T YPE L OCALITY : Mexico, Durango, 14 mi SW El Salto. D IAGNOSIS : This species is somewhat similar to A. recurvatum externally but with male proand mesobasotarsomeres somewhat more broadly expanded and with a large, flat, triangular process extending anteriorly from the dorsal surface of the labium. The male genitalia (figs. 203–205) are somewhat similar to A. recurvatum (figs. 267–269) as well but differ somewhat in the shape of the apical portion of the median lobe and the operculum. In addition, this species bears distinct microreticulation on the dorsal surface of the pronotum and elytron consisting of small, isodiametric cells, a condition unique among Mexican Agathidium, but similar to more northern species such as A. exiguum. D ESCRIPTION : Body moderate in size (TBL 5 3.88 mm), broad, robust (PNW/TBL 5 0.46), strongly contractile. Head and pronotum red; elytra red, not iridescent; venter yellow-red, antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.57), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, with indistinct microreticulation consisting of fine, isodiametric cells; frontoclypeal suture obsolete medially; with prominent, anteriorly directed, flat, triangular process extending from (apparently) labium, process upturned apically; eyes moderately large, not dorsoventrally compressed; gula slightly concave; antennomere ratios: length I:II:III 5 1.9:1.0:1.7, width VII:VIII:IX 5 1.0:1.0:2.1. Pronotum very large, broad (PNL/PNW 5 0.77), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, with indistinct microreticulation consisting of fine, isodiametric cells. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.06); punctation similar to pronotum but with microretic- 100 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY ulation much more distinctly evident; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, flattened; medial carina well developed. Metasternum narrow (MTL/ MTW 5 0.16), flattened, sloping dorsad anteriorly; oblique femoral carinae moderately prominent laterally, medially meeting in moderately prominent, posteriorly directed lobe. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small field of ventral spatulate setae; mandibles not modified; metafemur moderately broad with broad, moderately large tooth subapically on posterior margin (fig. 127); metasternal fovea moderately large, rounded with moderately large pencil of long, dense, fine setae. Median lobe in lateral aspect slender basally, strongly curved, expanded submedially, with prominent dorsal prominence and lateral carina between which is a sulcus in which fits the lateral lobe, apical portion elongate triangular, slightly sinuate, apex sharply pointed and directed anterad (fig. 204); in ventral aspect slender, long, lateral margins subparallel, apical portion evenly narrowed to slender, sharply pointed apex (fig. 203); operculum moderately long, apex with V-shaped emargination, each ramus triangular in shape, with points directed laterad and ventrad (fig. 203); lateral lobes slender, evenly curved through much of length, apically strongly sinuate and distinctly expanded submedially and apically, apex rounded with 2 stout setae (fig. 205). Female unknown. E TYMOLOGY : This species is named from the Greek word rhamphos, meaning ‘‘curving beak’’, for the unusual anteriorly produced structure of the labium. D ISTRIBUTION : This species is known only from the type locality in Durango (fig. 361). D ISCUSSION : The type specimen was collected from ‘‘bunch grass and sod’’. Agathidium megoniscoides Miller and Wheeler, new species Figures 128, 206–209, 362 T YPE M ATERIAL : Holotype, ? in AMNH labeled ‘‘MEX.:Mor. 7 mi S Tres Cumbres VII-7-1975/QDWheeler colr./HOLOTYPE Agathidium megoniscoides Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Morelos, 7 mi S Tres Cumbres. D IAGNOSIS : This species is among the largest of Nearctic Agathidium and is about the size of many A. oniscoides. The male metafemoral tooth is subapical and relatively small (fig. 128). The male genitalia are diagnostic with the median lobe long, the apical portion very short and stout, and with weakly developed lateral sulci for reception of the lateral lobes (fig. 208). D ESCRIPTION : Body large (TBL 5 4.17– 4.69 mm), broad, robust (PNW/TBL 5 0.44– 0.50), strongly contractile. Head and pronotum dark red; elytra dark red to testaceous, not iridescent; venter yellow, red-brown on metasternum; antennae, palpi, and legs yellow to yellow-red. Head broad (MDL/OHW 5 0.62–0.70), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes slightly compressed, but protuberant and conspicuous; gula convex; antennomere ratios: length I:II: III 5 2.2:1.0:2.3, width VII:VIII:IX 5 1.0: 1.0:2.1. Pronotum very large, broad (PNL/ PNW 5 0.76–0.79), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.81– 1.13); punctation similar to pronotum; sutural stria extending from apex to about half elytral length. Flight wings strongly reduced. Mesosternum with surface convex, broad, anterior margin broadly curved; medial carina obscured anteriorly. Metasternum moderately narrow (MTL/MTW 5 0.12–0.18), flattened, slightly sloping dorsad anteriorly; oblique femoral carinae low, not strongly developed, especially laterally, medially low and obsolete. Male tarsi 5-5-4; proand mesobasotarsomeres distinctly laterally expanded, with large field of ventral spatulate setae; mandibles not modified; metafemur broad, with relatively small curved subapical tooth on posterior margin (fig. 128); metasternal fovea 2005 101MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II large, transversely oval, with large, dense brush of fine setae. Median lobe in lateral aspect long, moderately straight throughout length, subapically broad, expanded along dorsal margin, apical portion short, robust, hooked dorsad (fig. 208); in ventral aspect long, straight and parallel-sided, slender, subapically expanded laterally, apical portion short and broadly triangular, apex pointed (figs. 206, 207); operculum in lateral aspect straight, slender, in dorsal aspect very broad, robust, apically truncate with prominent, robust lateral teeth (fig. 207); lateral lobes long, slender, evenly curved through most of length, apex expanded and sinuate, apex pointed, without stout setae (fig. 209). Female tarsi 5-4-4. E TYMOLOGY : This species is named from the Greek word mega, meaning ‘‘large’’, and the name oniscoides, a common eastern North American species for which this entire species group is named, in reference to the large size of this species and its placement in this group. D ISTRIBUTION : This species is known only from the type series from Morelos, Mexico (fig. 362). P ARATYPES : MEXICO: Morelos: 7miSTres Cumbres, 7 Jul 1975, QD Wheeler (4, QDWC). Agathidium grumum Miller and Wheeler, new species Figures 129, 210–212, 362 T YPE M ATERIAL : Holotype, ? in AMNH labeled ‘‘MEXICO: HIDALGO 5 mi NE Acatzingo 27.VI.1975 Q.D.Wheeler [handwritten]/HOLOTYPE Agathidium grumum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Hidalgo, 5 mi NE Acatzingo. D IAGNOSIS : Members of this species can be distinguished from others by the presence of a prominent medial tumidity on the gula, the large, acutely pointed male metafemoral tooth (fig. 129), and the shape of the male genitalia, which are relatively undifferentiated (figs. 210–212). There is a distinct sutural stria on the apical one-third of the elytron. This species is similar to A. popocatepetlae (including the shape of the male genitalia), but that species has a smaller male metafemoral tooth (fig. 133) and lacks the tumidity on the gula. D ESCRIPTION : Body moderately large (TBL 5 3.12 mm), broad, robust (PNW/TBL 5 0.48), rounded, strongly contractile. Head and pronotum dark red; elytra dark red-brown, not iridescent; venter yellow; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.59), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes moderately large, only slightly compressed; gula with prominent medial tumidity; antennomere ratios: length I:II:III 5 2.4:1.0:1.9, width VII:VIII:IX 5 1.0:1.0:2.0. Pronotum very large, broad (PNL/PNW 5 0.71), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.01); punctation and surface similar to pronotum; sutural stria present in apical one-third of elytron. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed, slightly obscured anteriorly. Metasternum narrow (MTL/MTW 5 0.14), slightly concave medially, distinctly dorsally sloped anteriorly; oblique femoral carinae moderately well developed, meeting medially in low, but prominent, posteriorly directed carina. Male tarsi 5-5-4; proand mesobasotarsomeres only somewhat laterally expanded, with moderate ventral field of spatulate setae; mandibles not modified; metafemur moderately broad, with very large, flat, triangular tooth subapically on posterior margin (fig. 129); metasternal fovea large, transversely oval with large, dense brush of long fine setae. Median lobe in lateral aspect moderately long, moderately stout, strongly bent basally, apical portion slender, angled dorsad, apically curved ventrad, acutely pointed (fig. 211); in ventral aspect moderately broad, parallelsided, apically evenly narrowed to narrowly rounded apex (fig. 210); operculum flat, moderately broad, evenly narrowed to rounded apex (fig. 210); lateral lobes long, slender, 102 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY slightly expanded and slightly sinuate apically, apices rounded with 2 stout setae (fig. 212). Female not examined. E TYMOLOGY : Named from the Latin word grumus, meaning ‘‘mound’’, for the prominent medial tumidity on the gula. D ISTRIBUTION : This species is known from southern Mexico (fig. 362). P ARATYPES : MEXICO: Mexico: 2miNETenancingo, 11 Sep 1973, 7100 9 , litter, pine-madrosaoak forest, Berlese, A Newton (1, CNCI); Morelia, 7 mi S Tres Cumbres, 7 Jul 1975, QD Wheeler (13, QDWC). D ISCUSSION : Habitat records for this species are from pine-oak forest litter at 7100 ft elevation. Agathidium triangularum Miller and Wheeler, new species Figures 130, 213–215, 361 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEX:Hgo;18 KmE Jacala, nr.El Alamo 10.VI.83, 1700 m S&J.Peck, moist oak for.litter/HOLOTYPE Agathidium triangularum Miller and Wheeler, 2003 [red label with black line border]’’. The only specimen examined of this species was the holotype. T YPE L OCALITY : Mexico, Hidalgo, 18 km E Jacala nr El Alamo. D IAGNOSIS : Members of this species can be distinguished from other species by the combination of a distinct ventral prominence on the gula which is foveate on the anterior surface, the prominent flat, triangular flange formed where the oblique metasternal carinae meet medially, moderately large and finely faceted eyes, the small male metafemoral tooth (fig. 130), and the male genitalia, which are relatively simple (figs. 213–215). The median lobe is relatively slender and straight and the apical portion is fairly short, apically pointed, and undifferentiated (fig. 213). The operculum is moderately slender and apically slightly expanded and rounded (fig. 213). D ESCRIPTION : Moderately large (TBL 5 3.14 mm), body broad, robust, rounded (PNW/TBL 5 0.5), strongly contractile. Head and pronotum dark red-brown; elytra dark red-brown, iridescent laterally; venter yellow-brown; antennae, palpi, and legs yellow to yellow-red. Head broad (MDL/OHW 5 0.59), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes protruding and prominent but dorsoventrally compressed; gula with prominent anteromedial tumidity which is strongly foveate on anterior surface; antennomere ratios: length I:II:III 5 1.9:1.0:2.2, width VII: VIII:IX 5 1.0:1.0:1.9. Pronotum very large, broad (PNL/PNW 5 0.73), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.74); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, not declivitous; medial carina well-developed, prominent medially. Metasternum narrowed (MTL/MTW 5 0.14), anteriorly sloping dorsad, concave medially; oblique femoral carinae well-developed, meeting medially in strongly-produced, triangular, posterior flange. Male tarsi 5-5-4; proand mesobasotarsomeres only somewhat laterally expanded, with moderate field of ventral spatulate setae; mandibles not modified; metafemur moderately slender, posteroapical angle produced into small tooth, apical margin sinuate (fig. 130); metasternal fovea prominent, large, transversely oval, with large, dense brush of fine setae. Median lobe in lateral aspect slender, curved basally, apical portion short, broad, directed slightly dorsad, apex slightly sinuate and pointed (fig. 214); in ventral aspect slender, lateral margins parallel, apical portion broad, tapered, apex formed into a narrow process (fig. 213); operculum long, flat, lateral margins convergent apically, apex a parallel-sided, apically rounded process (fig. 213); lateral lobes long, slender, apically expanded, apex pointed with 2 subapical stout setae (fig. 215). Female not examined. E TYMOLOGY : Named from the Latin word triangularus, meaning ‘‘triangle’’, for the 2005 103MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II distinctly triangular shape of the medial metasternal flange. D ISTRIBUTION : This species is known only from the type locality in Hidalgo (fig. 361). D ISCUSSION : The single type specimen was collected from moist oak forest litter at 1700 m elevation. Agathidium lobosternum Miller and Wheeler, new species Figures 131, 216–218, 362 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEX:Hgo; Tlanchinol 43 km SW Huejutla, 1500 m 14.VI.83.S&J.Peck cloud forest litter/HOLOTYPE Agathidium lobosternum Miller and Wheeler, 2003 [red label with black line border]’’. Unfortunately, the single specimen examined of this species is in several pieces glued to a point. T YPE L OCALITY : Mexico, Hidalgo, Tlanchinol 43 km SW Huejutla. D IAGNOSIS : Members of this species have a distinctive tubercle medially near the anterior margin of the gula and a very prominent lobe formed where oblique femoral carinae of the metasternum meet medially. The male genitalia are fairly simple, slender and with the lateral sulcus on the median lobe for reception of the lateral lobe only very weakly developed (fig. 217). D ESCRIPTION : Body moderately large (TBL 5 2.85 mm), broad (PNW/TBL 5 0.38), robust, rounded, strongly contractile. Head and pronotum dark red-brown; elytra dark red-brown, not iridescent; venter yellow-brown; antennae, palpi and legs yellow. Head broad (MDL/OHW 5 0.86), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes dorsoventrally distinctly compressed; gula flat, anterior margin with small, prominent tumidity, anterior surface of tumidity with slight fovea; antennomere ratios: length I:II:III 5 1.3:1.0: 1.3, width VII:VIII:IX 5 1.0:1.0:2.2. Pronotum very large, broad (PNL/PNW 5 0.98), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.02); punctation and surface similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed, obscured anteriorly. Metasternum narrow (MTL/MTW 5 0.13), slightly concave medially, distinctly dorsally sloped anteriorly; oblique femoral carinae prominent, well-developed, medially meeting in very prominent, narrowly-rounded, posteriorly-directed lobe. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur moderately broad, with small tooth subapically along posterior margin (fig. 131); metasternal fovea large, transversely oval with large brush of fine, long setae. Median lobe in lateral aspect slender, curved basally, slightly constricted medially and expanded thereafter, with weak lateral sulcus for reception of lateral lobe, apical portion of median lobe short, sinuate, slender, apex pointed and hooked ventrad (fig. 217); in ventral aspect slender, slightly narrowed medially, apical portion broad, evenly tapered to pointed apex (fig. 216); operculum flat, moderately broad, lateral margins curved to rounded apex (fig. 216); lateral lobes slender, curved medially, apically slightly sinuate, apices narrowly rounded, each with 2 stout setae (fig. 218). Female not examined. E TYMOLOGY : Named from the Greek words lobos, meaning ‘‘lobe’’, and sternum, meaning ‘‘chest’’, for the lobe formed where the oblique femoral carinae of the metasternum meet medially. D ISTRIBUTION : This species is known only from Hidalgo (fig. 362). D ISCUSSION : The single observed specimen was collected at 1500 m from cloud forest litter. Agathidium potosii Miller and Wheeler, new species Figures 132, 219–221, 363 T YPE M ATERIAL : Holotype, ? in FMNH labeled ‘‘MEX:S.L.P.14 mi.W Xilitla 4800 9 VI.29.73 A.Newton/leaf & log litter non-co- 104 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 363, 364. Geographic distribution of Agathidium oniscoides-group species: 363, A. popocatepetlae 5m ;A. potosii 5v ;A. stenomma 5. ;A. tribulosum 5m .364, A. hyle 5. ;A. grandidentatum 5V ;A. multidentatum 5v ;A. sejunctum 5m ;A. tribulograndum 5v . 2005 105MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II niferous Liquidambar for. /HOLOTYPE Agathidium potosii Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, San Luis Potosi, 14 mi W Xilitla. D IAGNOSIS : This species is similar to A. popocatepetlae,A. hyle, and other relatively undifferentiated Mexican species. The male metafemoral tooth is small (fig. 132), the male metasternal fovea is large and transverse, the metasternum is moderately narrow, and the oblique metasternal lines are not strongly prominent and meet medially in a broad, subtriangular flange. The male aedeagus is not strongly modified (figs. 219–221). The apex of the median lobe is short, subtriangular, sharply pointed and, in lateral aspect, strongly curved apically (fig. 220). The operculum is moderately long, broad, flattened, and has the apex distinctly expanded laterally with lateral teeth (fig. 219). D ESCRIPTION : Body small (TBL 5 2.22 mm), broad, robust (PNW/TBL 5 0.48), strongly contractile. Head and pronotum red; elytra red, not iridescent; venter yellow-brown; antennae and palpi yellow to yellow-brown. Head broad (MDL/OHW 5 0.61), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes relatively prominent and coarsely faceted, somewhat dorsoventrally compressed; gula strongly concave in anterior half; antennomere ratios: length I:II:III 5 2.1:1.0:1.5, width VII:VIII:IX 5 1.0:1.0: 2.3. Pronotum very large, broad (PNL/PNW 5 0.68), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.93); punctation similar to pronotum; a faint sutural stria extending from apex for about one-fifth of elytral length. Flight wings strongly reduced. Mesosternum broad, convex; medial carina prominent. Metasternum moderately broad (MTL/MTW 5 0.20), medial surface slightly concave, distinctly sloping dorsad anteriorly; oblique femoral carinae moderately well developed, meeting medially in broadly triangular, posteriorly directed flange. Male tarsi 5-5-4; proand mesobasotarsomeres moderately laterally expanded, with moderately large ventral field of spatulate setae; mandibles not modified; metafemur moderately broad, with prominent, small tooth subapically on posterior margin (fig. 132); metasternal fovea large, transverse, with large, prominent, dense brush of long fine setae. Median lobe in lateral aspect slender, strongly curved basally, relatively straight thereafter, apical portion short, slender, directed at angle dorsad, apically curved back ventrad, apex sharply pointed (fig. 220); in ventral aspect slender, slightly expanded at base of apical portion, which has the lateral margins slightly curved to broadly pointed apex (fig. 219); operculum flat, broad, moderately long, apex slightly expanded with lateral teeth (fig. 219); lateral lobes very slender, evenly curved basally, straight distally, apically slightly expanded and slightly sinuate, apices rounded with 2 stout setae (fig. 221). Female tarsi 5-4-4. E TYMOLOGY : This species is named for the type locality. D ISTRIBUTION : This species is known only from San Luis Potosi and Michoacan (fig. 363). P ARATYPES : MEXICO: Michoacan: Cerro de Garnica, Puerto Garnica, 18 Sep 1973, 9400 9 , oak-pine, Berlese, A Newton (3, CNCI). D ISCUSSION : This species has been collected from pine and Liquidambar forest from 4800 to 9400 ft elevation. Agathidium popocatepetlae Miller and Wheeler, new species Figures 133, 222–224, 363 T YPE M ATERIAL : Holotype, ? in CNCI labeled ‘‘MEX. Km. 10,000 9 V. Popocatepetl Mex., V-9-1971 J.M. Campbell /HOLOTYPE Agathidium popocatepetlae Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, State of Mexico, Popocatepetl, 10,000 9 . D IAGNOSIS : Members of this species can be distinguished from others by the combination of a small male metafemoral tooth that is lo- 112 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.93–1.07); punctation and surface similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina low but distinctly developed. Metasternum narrow (MTL/MTW 5 0.13–0.17), slightly concave medially, distinctly dorsally sloped anteriorly; oblique femoral carinae moderately well developed, forming a low carina medially. Male tarsi 5-5-4; proand mesobasotarsomeres somewhat laterally expanded, protarsomeres more so and with small field of ventral spatulate setae; mandibles not modified; metafemur moderately broad, with several prominent, small teeth medially along posterior margin (fig. 138); metasternal fovea small with small pencil of fine setae. Median lobe in lateral aspect slender basally, more robust submedially, with small, but prominent, lobelike lateral carinae between which fit the lateral lobes, apical portion relatively short, curved ventrad, apex pointed (fig. 242); in ventral aspect moderately slender, apical portion narrowed, slender to expanded, rounded apex (figs. 240, 241); operculum in lateral aspect sender, apically expanded and slightly curved ventrad (fig. 242), in ventral aspect broad, short, apically narrowly emarginate, each ramus with small lateral tooth (fig. 240); lateral lobes slender, curved basally, apical one-third broader and sinuate, apices rounded, each with 2 stout setae (fig. 243). Female not examined. E TYMOLOGY : Named for the Latin word invisitatus, meaning ‘‘strange’’, for the unusual teeth on the male metafemur. D ISTRIBUTION : This species has been collected from central Mexico (fig. 365). P ARATYPE : MEXICO: San Luis Potosi: 14 mi W Xilitla, 29 Jun 1973, 4800 9 , under bark, Liqidambar forest, A Newton (1, FMNH). D ISCUSSION : The species is known from under bark at 4800 to 11,000 ft elevation. Agathidium multidentatum Miller and Wheeler, new species Figures 139, 244–247, 364 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEX.:OAX.; 3.5 mi S Suchixtepec, 8000 9 3.vi.1971 S. Peck Ber208, leaflitter/ HOLOTYPE Agathidium multidentatum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Oaxaca, 3.5 mi S Suchixtepec. D IAGNOSIS : Males of this species are recognizable by the multidentate metafemur (fig. 139) and the very unique median lobe with the operculum long and slender and apically with lateral hooks (fig. 244). The metasternum is moderately narrow and the metasternal fovea is transverse and moderately large. D ESCRIPTION : Body relatively large (TBL 5 3.52–3.74 mm), broad, robust (PNW/TBL 5 0.46), strongly contractile. Head and pronotum testaceous; elytra testaceous, not iridescent; venter dark brown; antennae and palpi yellow-brown; legs dark brown to red brown. Head broad (MDL/OHW 5 0.51–0.53), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eye slightly compressed, but not strongly reduced; gula convex anteriorly; antennomere ratios: length I:II:III 5 1.7:1.0: 1.4, width VII:VIII:IX 5 0.9:1.0:2.1. Pronotum very large, broad (PNL/PNW 5 0.80– 0.82), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.07–1.08); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina present, but not strongly developed. Metasternum moderately narrow (MTL/MTW 5 0.12–0.13), flattened, sloping dorsad anteriorly; oblique femoral carinae well developed, meeting medially in very low carina. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur moderately broad, posteroapical margin with conspicuous, curved tooth and series of smaller teeth, apical margin irregularly excavated (fig. 2005 113MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II 139); metasternal fovea linear, transverse with line of dense, fine setae. Median lobe in lateral aspect long, slender, moderately curved, dorsal margin bisinuate, apex long, slender, straight with apical reflexed hook (fig. 246); in ventral aspect slender, with medial abrupt expansion consisting of laterally directed carina which together with the dorsal carinae form a sulcus in which fits the lateral lobe, apex slender, slightly expanded medially tapering to narrowly rounded apex (figs. 244, 245); operculum long, apex laterally produced into conspicuous laterally and dorsally directed hooks, apex distinctly emarginate (fig. 244); lateral lobes very long, slender, medially expanded and sinuate, apex slender to very narrowly rounded, with 2 long apical setae (fig. 247). Female tarsi 5-4-4. E TYMOLOGY : This species is named from the Latin words multus, meaning ‘‘much’’, and dentatus, meaning ‘‘tooth’’, for the multiple teeth present along the posterior margin of the male metafemur in this species. D ISTRIBUTION : This species is known only from Oaxaca (fig. 364). P ARATYPES : MEXICO: Oaxaca: 5.1 km S Suchixtepec, 25 Jul 1992, 2150 m, oak-alder-pine leaf litter, Berlese, RS Anderson (1, CNCI); 4.6 km S Suchixtepec, 23 Jul 1992, wet riparian alder forest, leaf litter, RS Anderson (1, CNCI); 3.5 mi S Suchixtepec, 3 Jun 1971, 8000 9 , leaf litter, S Peck (9, PECK). D ISCUSSION : The species has been collected from oak, alder, pine, and ‘‘wet riparian’’ forest litter. Elevation records are from about 2400 m. Agathidium sejunctum Miller and Wheeler, new species Figures 140, 248–250, 364 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEXICO: Chiapas: Volcan Tacana, lower slopes, ca. 4 km N Union Juarez. 18IX-1992 R.S.Anderson 92-109/cloud forest litter Elev. 1950 m./HOLOTYPE Agathidium sejunctum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Chiapas, lower slopes of Volcan Tacana, ca. 4 km N Union Juarez, 1950 m. D IAGNOSIS : This species can be distinguished from most species in this subgroup by the very large, acutely pointed male metafemoral tooth (fig. 140). The gula is unmodified. The male genitalia are relatively simple (figs. 248–250). This species is extremely similar to A. grandidentatum but specimens are larger, are generally iridescent dorsally, have the mesosternal carina medially usually with a low spot dividing the carina into anterior and posterior portions, and exhibit some differences in shape of the male genitalia. In A. sejunctum there is a prominent dorsal lobelike carinae alongside which is a large fovea in which fits the lateral lobes (fig. 249). This is much less distinct in A. grandidentatum (fig. 252). D ESCRIPTION : Body moderately large (TBL 5 2.69–3.16 mm), broad, robust, rounded (PNW/TBL 5 0.44–0.48), strongly contractile. Head and pronotum red-brown to piceous; elytra red-brown to piceous, iridescent; venter yellow to red-brown; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.58–0.66), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes compressed dorsoventrally; gula slightly concave; antennomere ratios: length I:II:III 5 2.0:1.0:1.4, width VII:VIII:IX 5 1.0:1.0: 2.2. Pronotum very large (PNL/PNW 5 0.77–0.82), broad, strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.80–1.03); punctation and surface similar to pronotum; sutural stria present in apical onefourth of elytron. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed, on most specimens clearly interrupted medially. Metasternum narrow (MTL/MTW 5 0.13– 0.15), slightly concave medially, distinctly dorsally sloped anteriorly; oblique femoral carinae moderately well developed, meeting medially in low, but prominent carina. Male tarsi 5-5-4; proand mesobasotar- 114 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY someres moderately laterally expanded, more so with protarsomeres, with moderate ventral field of spatulate setae; mandibles not modified; metafemur moderately broad, with very large, flat, acutely pointed tooth subapically on posterior margin (fig. 140); metasternal fovea moderately large, transverse with large, dense brush of long fine setae. Median lobe in lateral aspect slender, moderately long, strongly curved basally, with distinct dorsal expansion at base of apical portion along side of which is a triangular fovea for reception of lateral lobe, apical portion of median lobe slender, distinctly flexed dorsad, apex abruptly curved ventrad, sharply pointed (fig. 249); in ventral aspect moderately broad, slightly expanded medially, apical portion evenly narrowed to pointed apex (fig. 248); operculum flat, short, slender, apex rounded, not emarginate (fig. 248); lateral lobes moderately broad, long, curved basally, slightly expanded and slightly sinuate apically, apices rounded with 2 stout setae (fig. 250). Female tarsi 5-4-4. E TYMOLOGY : This species is named for the Latin word sejunctum, meaning ‘‘separated’’, for the medially interrupted mesosternal carina in most members of this species. D ISTRIBUTION : This species is known only from Chiapas (fig. 364). P ARATYPES : MEXICO: Chiapas: 7.4 km SSW Motozintla de Mendoza, 17 Sep 1992, cloud forest litter, R Anderson (12, CNCI); Volcan Tacana, lower slopes, 4 km N Union Juarez, 19 Sep 1992, 2000 m, cloud forest litter, R Anderson (43, CNCI). D ISCUSSION : The species has been collected from cloud forest litter at 2000 m. Agathidium grandidentatum Miller and Wheeler, new species Figures 141, 252, 253, 364 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEXICO: Guerrero: 5.6 km SW Filo de Caballo R.S.Anderson 92-013 17VII-1992/alder forest, Berlese litter and twigs. Elev. 2310 m/HOLOTYPE Agathidium grandidentatum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Guerrero, 5.6 km SW Filo de Caballo, 2310 m. D IAGNOSIS : This species can be distinguished from most species in this subgroup by the very large, acutely pointed male metafemoral tooth (fig. 141). The gula is unmodified. The male genitalia are relatively simple (figs. 252, 253). The lateral sulcus on the median lobe for reception of the lateral lobes is relatively indistinct compared with other members of the subgroup, though the lateral lobes are prominently sinuate (fig. 252). This species is extremely similar to A. sejunctum but specimens are smaller, are not generally iridescent dorsally, have the mesosternal carina entire, and exhibit some differences in shape of the male genitalia. In A. sejunctum there is a prominent dorsal lobelike carinae alongside which is a large fovea in which fit the lateral lobes (fig. 249). This is much less distinct in A. grandidentatum (fig. 252). D ESCRIPTION : Body small (TBL 5 2.00– 2.32 mm), broad, robust, rounded (PNW/ TBL 5 0.44–0.46), strongly contractile. Head and pronotum red-brown to piceous; elytra red-brown to piceous, not iridescent; venter yellow to red-brown; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.53–0.70), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes compressed dorsoventrally; gula flat to slightly convex; antennomere ratios: length I: II:III 5 1.4:1.0:1.3, width VII:VIII:IX 5 1.0: 1.0:2.2. Pronotum very large, broad (PNL/ PNW 5 0.73–0.84), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.87–0.98); punctation and surface similar to pronotum; sutural stria present in apical onefourth of elytron. Flight wings strongly reduced to absent. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.11–0.19), slightly concave medially, distinctly dorsally sloped anteriorly; oblique femoral carinae moderately well developed, 2005 115MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II meeting medially in low, but prominent carina. Male tarsi 5-5-4; proand mesobasotarsomeres distinctly laterally expanded, protarsomeres slightly more, with moderate field of ventral spatulate setae; mandibles not modified; metafemur moderately broad, with very large, flat, acutely pointed tooth subapically on posterior margin (fig. 141); metasternal fovea large, transverse with large, dense brush of long fine setae. Median lobe in lateral aspect slender, moderately long, strongly curved basally, apical portion slender, distinctly flexed dorsad, apex abruptly curved ventrad, sharply pointed (fig. 252); in ventral aspect moderately broad, slightly expanded medially, apical portion evenly narrowed to pointed apex (fig. 251); operculum narrow, slightly tapered apically, apex distinctly and broadly emarginate (fig. 251); lateral lobes moderately broad, long, curved basally, slightly expanded and slightly sinuate apically, apices rounded with 2 stout setae (fig. 253). Female tarsi 5-4-4. E TYMOLOGY : This species is named from the Latin words grandis, meaning ‘‘large’’, and dentatus, meaning ‘‘tooth’’, for the large tooth on the metafemur of males of this species. D ISTRIBUTION : This species has been collected only from Guerrero (fig. 364). P ARATYPES : MEXICO: Guerrero: 15.0 km SW Filo de Caballo, 16 Jul 1992, 2500 m, wet oak forest litter, Berlese, RS Anderson (13, CNCI); 10.3 km SW Filo de Caballo, 13 Jul 1992, 2700 m, wet oak-pine-fir forest, leaf log litter, Berlese, R Anderson (75, CNCI). D ISCUSSION : This species has been collected from leaf and log litter from wet oak, pine, and fir forest at 2500–2700 m elevation. Agathidium andersoni Miller and Wheeler, new species Figures 142, 254–256, 365 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘GUAT.SACATEPEQUEZ: 4.5 km SW San Miguel Duenas. Elev. 1760 m. mesic hardwd. for litter, R.Anderson 91-61, 12VI-1991/HOLOTYPE Agathidium andersoni Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Guatemala, Sacatepequez, 4.5 km SW San Miguel Duenas, 1760 m. D IAGNOSIS : Members of this species have relatively large eyes and a small, subapical male metafemoral tooth (fig. 142). The metasternum is moderately broad medially. The species is best identified by examining the male genitalia, which has the median apical portion of the median lobe and the operculum each broadly truncate apically with a shallow medial emargination in ventral aspect (fig. 254). The operculum extends to about the apex of the apical portion of the median lobe (fig. 254). The lateral sulcus and associated carinae for reception of the lateral lobe are well developed (fig. 255). D ESCRIPTION : Body size relatively small (TBL 5 2.58–2.90 mm), broad (PNW/TBL 5 0.46–0.47), robust, strongly contractile. Head and pronotum testaceous; elytra testaceous, with blue iridescence laterally; venter yellow-brown; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.55–0.62), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes prominent, not strongly compressed; gula slightly concave; antennomere ratios: length I:II:III 5 2.0:1.0:1.3, width VII:VIII:IX 5 1.0:1.0:2.2. Pronotum very large, broad (PNL/PNW 5 0.72), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.03–1.15); punctation similar to pronotum; sutural stria present in apical one-third of elytron. Flight wings fully developed. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum moderately narrow (MTL/ MTW 5 0.16–0.18), flattened, slightly sloping dorsad anteriorly; oblique femoral carinae well developed, moderately prominent medially. Male tarsi 5-5-4; proand mesobasotarsomeres moderately laterally expanded, pro- 116 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY tarsomeres more so and with large ventral field of spatulate setae; mandibles not modified; metafemur broad, posterior apical angle produced into broad tooth, apical margin broadly rounded (fig. 142); metasternal fovea large, transversely oval, with dense, fine, long setae. Median lobe in lateral aspect abruptly bent basally, approximately straight thereafter, slender, distinctly expanded submedially along dorsal margin by lateral carina which together with dorsal carinae form a sulcus in which fits the lateral lobe, apical portion of median lobe short, slender, curved, gradually narrowed to narrowly rounded apex (fig. 252); in ventral aspect moderately robust, distinctly expanded submedially, apex broad, apically truncated with slight medial emargination (fig. 251); operculum long, extending to apex of median lobe, broadly curved ventrally, broad in ventral aspect, margins parallel, apex broad, truncate with slight medial emargination (fig. 251); lateral lobes long, slender, broadly and conspicuously expanded medially, apex slightly expanded in lateral aspect, in ventral aspect sinuate medially, abruptly narrowed subapically, with a single subapical seta (fig. 253). Female tarsi 5-4-4. E TYMOLOGY : This species is named for R.S. Anderson (Canadian Museum of Nature, Ottawa), who has contributed considerably to knowledge of Coleoptera through excellent research and valuable field work. He also collected the holotype specimen and many specimens of other Agathidium. D ISTRIBUTION :Agathidium andersoni has been collected from southern Mexico and Guatemala (fig. 365). P ARATYPES : GUATEMALA: Baja Verapaz: 8 km S Purulha, 20 May 1991, 1660 m, pine cloud forest litter, R Anderson (1, PECK). Sacatepequez: 4.5 km SW San Miguel Duenas, 12 Jun 1991, 1760 m, mesic hardwood forest litter, R Anderson (4, PECK). MEXICO: Chiapas: Jitotol, 22 Sep 1986, 1500 m, leaf litter under Salix(?) shrubs, R Baranowski (1, LUND). D ISCUSSION : This species has been collected from litter unders Salix shrubs, pine cloud forest and mesic hardwood forest litter from 1500 to 1760 m. Agathidium disgregum Miller and Wheeler, new species Figures 143, 254–256, 365 T YPE M ATERIAL : Holotype, ? in CNCI labeled ‘‘MEX. Cerro Potosi, NL V-3-1971, 8500 9 J.M. Campbell/HOLOTYPE Agathidium disgregum Miller and Wheeler, 2003 [red label with black line border]’’. Only a single specimen was examined of this species. T YPE L OCALITY : Mexico, Nueva Leon, Cerro Potosi, 8500 9 . D IAGNOSIS : This species is similar to A. recurvatum in most external characters except that the eyes are a little larger. The operculum of the median lobe is approximately Tshaped with broad lateral hooks apically (fig. 257), a condition unique in this species subgroup. D ESCRIPTION : Body relatively large (TBL 5 3.81 mm), broad, robust (PNW/TBL 5 0.48), strongly contractile. Head red; pronotum dark red; elytra red, not iridescent; venter red-brown, antennae, palpi, and legs red-brown. Head broad (MDL/OHW 5 0.66), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes moderately large, not dorsoventrally compressed; gula slightly concave; antennomere ratios: length I:II:III 5 2.2:1.0: 2.2, width VII:VIII:IX 5 1.0:1.0:2.0. Pronotum very large, broad (PNL/PNW 5 0.69), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.02); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, flattened; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.18), flattened, sloping dorsad anteriorly; oblique femoral carinae moderately prominent laterally, medially meeting in low carina. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small field of ventral spatulate setae; man- 2005 117MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II dibles not modified; metafemur moderately broad with small tooth subapically on posterior margin (fig. 143); metasternal fovea moderately large, rounded with moderately large pencil of long, dense, fine setae. Median lobe in lateral aspect slender basally, strongly curved, slightly expanded submedially, with prominent dorsal prominence and lateral carina with lateral sulcus in which fits the lateral lobe, apical portion elongate triangular, slightly sinuate apex abruptly hooked and sharply pointed (fig. 259); in ventral aspect slender, long, lateral margins broadly sinuate, apical portion evenly narrowed to slender, narrowly rounded apex (figs. 257, 258); operculum slender basally, very broadly expanded apically (fig. 257); lateral lobes slender, evenly curved through much of length, apically strongly sinuate and expanded submedially and apically, apex rounded with 2 stout setae (fig. 260). Female not examined. E TYMOLOGY : Name from the Latin word disgregus, meaning ‘‘different’’, for the unusual shape of the male operculum. D ISTRIBUTION : This species is known only from the type locality (fig. 365). D ISCUSSION : A single altitude record is from 8500 ft. Agathidium oaxacaense Miller and Wheeler, new species Figures 144, 261–263, 365 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEX: Oax.; 37 mi Valle Nacional 8500 9 24.V.71 oak litter Ber 206 S. Peck/HOLOTYPE Agathidium oaxacaense Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Oaxaca, 37 mi S Valle Nacional. D IAGNOSIS : This species is externally similar to several species in the A. aztec subgroup. The eyes are moderately dorsoventrally compressed with facets of moderate size. The male metafemoral tooth is moderately large, anteapical, and there is a series of smaller serrations along the posterior margin of the metafemur (fig. 144). The male genitalia are very distinctive with the median lobe very robust and strongly curved (fig. 262). There is a very prominent dorsomedial carina with a prominent lateral fovea that receives the lateral lobe (fig. 262). D ESCRIPTION : Body moderately small (TBL 5 2.54–3.16 mm), broad, robust (PNW/TBL 5 0.46–0.53), strongly contractile. Head and pronotum testaceous; elytra testaceous, slightly iridescent in some specimens; venter, antennae, palpi, and legs dark brown. Head broad (MDL/OHW 5 0.53–0.60), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes slightly compressed, but not strongly reduced; gula slightly concave; antennae moderately long (ratios: length I:II:III 5 1.9:1.0:1.7, width VII:VIII: IX 5 1.0:1.0:1.8). Pronotum very large, broad (PNL/PNW 5 0.64–0.75), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.87–1.23); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, not declivitous, flattened; medial carina well developed. Metasternum moderately narrow (MTL/MTW 5 0.13–0.17), sloping dorsad anteriorly and concave medially; oblique femoral carinae prominent, meeting medially in large, semicircular flange. Male tarsi 5-5-4; proand mesobasotarsomeres only slightly laterally expanded, with small field of ventral spatulate setae; mandibles not modified; metafemur moderately broad, with prominent subapical tooth on posterior margin and numerous smaller serrations along posterior margin (fig. 145); metasternal fovea linear, transverse with line of dense, fine setae. Median lobe in lateral aspect short, very robust, abruptly curved basally, broadly expanded submedially due to dorsal production of dorsomedial carina, with large fovea laterad of carina for reception of lateral lobe, apical portion of median lobe slender, apically pointed, slightly hooked ventrally (fig. 262); in ventral aspect robust, lateral margins subparallel for most 118 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY of length, abruptly narrowed subapically, apex a slender, narrowly rounded process (fig. 261); operculum in lateral aspect slender basally, abruptly expanded apically, in ventral aspect short, broad, broadly rounded, apex slightly emarginate (fig. 261); lateral lobes slender basally, strongly expanded submedially, expanded portion fitting into cavity on each side of median lobe, apical portion strongly sinuate, apex broadly expanded, apically rounded, with 2 stout apical setae (fig. 263). Female tarsi 5-4-4. E TYMOLOGY : Named for the state in Mexico in which this species is found. D ISTRIBUTION : This species is known only from the state of Oaxaca (fig. 365). P ARATYPE : MEXICO: Oaxaca: 37 mi S Valle Nacional, 24 May 1971, 8500 9 , oak litter, S Peck (5, PECK). D ISCUSSION : The type series was collected from oak litter at 8500 ft. Agathidium oculeum Miller and Wheeler, new species Figures 108, 145, 264–266, 366 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEXICO: Chiapas: Volcan Tacana, lower slopes, ca. 4 km N Union Juarez. 18IX-1992. R.S.Anderson 92-109/cloud forest litter Elev. 1950 m./HOLOTYPE Agathidium oculeum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Chiapas, lower slopes of Volcan Tacana, ca. 4 km N Union Juarez, 1950 m. D IAGNOSIS : This species is very similar to A. recurvatum but the eyes are much more prominent, protruding, and finely faceted (fig. 108). The male genitalia are also similar to that species but differ in that the operculum is broader and more rounded in A. oculeum (fig. 264) than in A. recurvatum (fig. 267). D ESCRIPTION : Body moderately large (TBL 5 2.82–3.97 mm), broad, robust (PNW/TBL 5 0.44–0.48), rounded, strongly contractile. Head and pronotum red-brown; elytra redbrown, not iridescent; venter, antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.64), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes large with many facets, not reduced (fig. 108); gula slightly convex medially; antennomere ratios: length I:II:III 5 2.1:1.0:1.8, width VII: VIII:IX 5 1.0:1.0:2.1. Pronotum very large, broad (PNL/PNW 5 0.73–0.76), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.03–1.20); punctation and surface similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.12–0.16), flat medially, distinctly dorsally sloped anteriorly; oblique femoral carinae moderately developed, not prominent, low where meeting medially. Male tarsi 5-5-4; proand mesobasotarsomeres only slightly laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur moderately broad, with small tooth subapically along posterior margin (fig. 145); metasternal fovea moderately large, transverse with line of fine, dense, long setae. Median lobe in lateral aspect slender, moderately short, strongly curved medially, apical portion long, slender, sinuate, apex sharply pointed, recurved ventrad (fig. 265); in ventral aspect slender, apical portion with margins sinuate, converging, apex slightly expanded and pointed apically (fig. 264); operculum flat, short, narrow basally, expanded apically, apex emarginate, each ramus broad and apically pointed (fig. 264); lateral lobes long, slender, not expanded, sinuate, apex narrowly rounded with 2 stout setae (fig. 266). Female not examined. E TYMOLOGY : Named for the Latin word oculeus, meaning ‘‘full of eyes’’, for the large, many faceted eyes of members of this species. D ISTRIBUTION : This species has been collected from San Luis Potosi and Chiapas (fig. 366). 2005 119MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II P ARATYPES : MEXICO: Chiapas: 10 mi SE San Cristobal de las Casas, 1 Sep 1993, 8000 9 , pinemadron-oak forest litter, Berlese, A Newton (4, FMNH); Pico Cerro Tzontehuitz, 10 km NE San Cristobal, 16 Sep 1991, 2910 m, cloud forest litter, RS Anderson (1, CNCI). D ISCUSSION : This species has been collected from cloud forest litter and leaf and log litter from Liquidambar, pine, oak, and madrone forests. Elevation records are from 4800 to 8000 ft. Agathidium recurvatum Miller and Wheeler, new species Figures 146, 267–269, 366 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEX.: OAX.; 3 mi N Suchixtepec, 9500 9 4.vi.1971 S.Peck Ber209, oak litter/ HOLOTYPE Agathidium recurvatum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Oaxaca, 3 mi N Suchixtepec, 9500 9 . D IAGNOSIS : This species is similar to A. aztec and related species in having a small, subapical male metafemoral tooth (fig. 146), but differs in having relatively smaller eyes with somewhat larger facets, more well-developed oblique metasternal carinae which meet medially in a moderately developed triangular lobe and the male genitalia. The median lobe in lateral aspect is apically slender, long, and sharply recurved at the apex (fig. 268). This species is similar to A. oculeum but has smaller eyes and that species has a very broad, rounded operculum (fig. 264) whereas that of A. recurvatum is smaller and more slender (fig. 267). This species is also similar to A. stenomma (including the shape of the male genitalia); however, that species has eyes that are very strongly dorsoventrally compressed (fig. 109). D ESCRIPTION : Body moderate in size (TBL 5 2.68–3.82 mm), broad, robust (PNW/TBL 5 0.44–0.48), strongly contractile. Head dark red-brown to piceous; pronotum dark red-brown to piceous, red around margins; elytra red-brown to piceous, red around margins, not iridescent; venter, antennae, palpi, and legs red-brown. Head broad (MDL/OHW 5 0.52–0.63), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes moderately large, but distinctly dorsoventrally compressed; gula slightly concave; antennomere ratios: length I:II:III 5 1.8:1.0:1.9, width VII:VIII:IX 5 1.0:1.0:1.5. Pronotum very large, broad (PNL/PNW 5 0.73–0.83), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.93–1.11); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, flattened; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.14–0.22), flattened, sloping dorsad anteriorly; oblique femoral carinae moderately prominent laterally, medially meeting in moderately prominent, posteriorly directed lobe. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small field of ventral spatulate setae; mandibles not modified; metafemur moderately broad with broad, moderately large tooth subapically on posterior margin (fig. 146); metasternal fovea moderately large, rounded with moderately large pencil of long, dense, fine setae. Median lobe in lateral aspect slender basally, strongly curved, expanded submedially, with prominent dorsal prominence, apical portion elongate triangular, apex abruptly recurved and sharply pointed (fig. 268); in ventral aspect slender, long, lateral margins broadly sinuate, apical portion evenly narrowed to slender, sharply pointed apex (fig. 267); operculum flat, long, relatively slender, apex emarginate, each ramus narrowly rounded (fig. 267); lateral lobes slender, evenly curved through much of length, apically strongly sinuate and slightly broadened, apex rounded with 2 stout setae (fig. 269). Female tarsi 5-4-4. E TYMOLOGY : This species is named recurvatum after the prominently recurved apex of the median lobe in lateral aspect. D ISTRIBUTION : This species is known only 120 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY from the states of Mexico and Oaxaca (fig. 366). P ARATYPES : MEXICO: Oaxaca: 23 km N Oaxaca City, 12 Sep 1994, 2650 m, under bushes at roadside, mixed pine forest, R Baranowski (13, JRAC); 26 km S Yolomecatl, 10 Aug 1988, Doyen and Stockwell (1, EMEC); 17 km N Villa Diaz Ordaz, 5 Sep 1994, 2750 m, pine oak forest litter, R Baranowski (5, JRAC); 40.5 km S Suchixtepec, 25 Jul 1992, 1300 m, cloud forest, leaf litter, Berlese, RS Anderson (2, CNCI); 64.5 km SW Valle Nacional km 117.5, 28 Jul 1992, 2600 m, oak forest leaf litter, Berlese, RS Anderson (3, CNCI); 5.1 km S Suchixtepec, 25 Jul 1992, oak-alder-pine forest leaf litter, Berlese, RS Anderson (7, CNCI); 4.6 km S Suchixtepec, 23 Jul 1992, 2150 m, wet riparian alder forest leaf litter, Berlese, RS Anderson (7, CNCI); 3.5 mi S Suchixtepec, 3 Jun 1971, 8000 9 , leaf litter, S Peck (4, PECK). D ISCUSSION : This species has been collected from litter from various forest types including pine, oak, alder, and wet riparian forest. Agathidium impensum Miller and Wheeler, new species Figures 147, 270–272, 366 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEXICO: Guerrero: 15.0 km SW Filo de Caballo, R.S.Anderson 92-011 16VII-1992/oak forest (wet) litter Berlese. Elev. 2600 m/HOLOTYPE Agathidium impensum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Guerrero, 15.0 km SW Filo de Caballo. D IAGNOSIS : This species is characterized by its very large size (approaching the size of A. megoniscoides and A. oniscoides), moderately large, curved apical male metafemoral tooth (fig. 147), and very distinctive male genitalia (figs. 270–272). The median lobe is very long with the apical portion very slender and straight and with very prominent dorsal and lateral carinae for reception of the lateral lobe (fig. 271). The operculum is broad (fig. 270) and apically hooked ventrad (fig. 271). D ESCRIPTION : Size large (TBL 5 4.21– 4.73 mm), body broad (PNW/TBL 5 0.46– 0.47), robust, rounded, strongly contractile. Head and pronotum yellow-red; elytra yellow-red, distinctly iridescent; venter yellowbrown to red-brown; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.52–0.61), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes prominent, not strongly compressed; gula shallowly concave; antennae moderately long; antennomere ratios: length I:II:III 5 2.4:1.0:2.1, width VII:VIII:IX 5 0.9:1.0:2.2. Pronotum very large, broad (PNL/PNW 5 0.78–0.79), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.04–1.19); punctation and surface similar to pronotum; sutural stria present in apical onefourth of elytron. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.14– 0.17), flat medially, distinctly dorsally sloped anteriorly; oblique femoral carinae moderately well developed, meeting medially in low carina. Male tarsi 5-5-4; proand mesobasotarsomeres only moderately laterally expanded, with moderate field of ventral spatulate setae; mandibles not modified; metafemur moderately slender, with curved, robust, large tooth apically (fig. 147); metasternal fovea large, transverse, with dense brush of long fine setae. Median lobe in lateral aspect robust, evenly curved, at base of apical portion with lateral lobelike carinae, apical portion narrowed, straight, slightly expanded along ventral margin (fig. 271); in ventral aspect slender, apical portion with lateral margins irregular, narrowed, apex a slender, apically rounded process (fig. 270); operculum in lateral aspect flat, apically hooked ventrad (fig. 271), in ventral aspect broad, apex broad with narrow medial emargination, each ramus of operculum rounded apically (fig. 270); lateral lobes long, curved basally, apical one-fifth expanded and sinuate, apices rounded, each with 2 stout setae (fig. 172). Female tarsi 5-4-4. 2005 121MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II E TYMOLOGY : This species is named from the Latin word impensus, meaning ‘‘large’’, for the relatively large size of specimens. D ISTRIBUTION : This species is known only from the type locality in Guerrero (fig. 366). P ARATYPE : MEXICO: Guerrero: 15 km SW Filo de Caballo, 16 Jul 1992, wet forest litter, RS Anderson (1, CNCI). D ISCUSSION : The type specimens were extracted from wet oak forest litter using a Berlese device. They were collected from 2600 m elevation. Agathidium cheneyi Miller and Wheeler, new species Figures 148, 273–276, 366 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEXICO: Chiapas: Yerbabuena Preserve 2.1 km NW Pueblo Nuevo, Solistahuacan. R.S. Anderson 92-114 23-IX1992/HOLOTYPE Agathidium cheneyi Miller and Wheeler, 2003 [red label with black line border]’’. The holotype is the only specimen examined of this species. T YPE L OCALITY : Mexico, Chiapas, Yerbabuena Preserve, 2.1 km NW Pueblo Nuevo, Solistahuacan. D IAGNOSIS : This species is similar to A. aztec and related species. This species differs in the greater development of the oblique metasternal carinae which meet medially in a very broad, truncate flange. The median lobe differs particularly in having the operculum very short and laterally rounded with a distinct apical emargination (fig. 273). D ESCRIPTION : Body moderately small (TBL 5 3.26 mm), broad, robust, rounded (PNW/TBL 5 0.47), strongly contractile. Head and pronotum red; elytra red, iridescent; venter yellow-red; antennae, palpi, and legs yellow-brown. Head broad (MDL/OHW 5 0.58), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes moderately large, slightly compressed dorsoventrally; gula flat; antennomere ratios: length I: II:III 5 1.6:1.0:1.5, width VII:VIII:IX 5 1.0: 1.0:2.1. Pronotum very large, broad (PNL/ PNW 5 0.71), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.07); punctation and surface similar to pronotum; sutural stria present in apical one-fifth of elytron. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.16), flat medially, distinctly dorsally sloped anteriorly; oblique femoral carinae moderately well developed, meeting medially in broadly truncate carina. Male tarsi 5-5-4; proand mesobasotarsomeres somewhat laterally expanded, with moderately large ventral field of spatulate setae; mandibles not modified; metafemur moderately broad, with small tooth subapically on posterior margin (fig. 148); metasternal fovea large, transversely oval with large region of long fine setae. Median lobe in lateral aspect slender, strongly curved basally, with moderate lateral and dorsal carinae at base of apical portion for reception of lateral lobe, apical portion slender, sinuate, apex very narrowly truncate (fig. 275); in ventral aspect slender, expanded at base of apical portion that has the lateral margins slightly sinuate and convergent to near apex, which is slightly expanded and rounded (figs. 273, 274); operculum broad, short, flat, with shallow V-shaped emargination apically (fig. 273); lateral lobes slender, apically sinuate, apex narrowly rounded with 2 stout setae (fig. 276). Female not examined. E TYMOLOGY : This species is named in honor of R.B. Cheney (Casper, WY), Vice-President of the United States. D ISTRIBUTION : This species is known only from Chiapas (fig. 366). Agathidium tenangoense Miller and Wheeler, new species Figures 149, 277–279, 367 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘GUAT: HUEHUETENANAGO 10 kmW La Capellania 3.viii.78. Helava./HOLOTYPE Agathidium tenangoense Miller 128 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY rowed to a slightly expanded apex in ventral aspect (fig. 296). The operculum is fairly long and slender and has the apex rounded with a prominent, but shallow, apical emargination (fig. 296). The species is nearly identical to A. tumidiventre, including genitalic shape, except for the presence in that species of a distinct tumidity medially on the gula. D ESCRIPTION : Body moderately large (TBL 5 3.29–3.64 mm), broad, robust (PNW/TBL 5 0.47–0.48), strongly contractile. Head and pronotum testaceous; elytra testaceous, not or only faintly iridescent; venter yellow-brown, antennae and palpi yellow; legs yellow to yellow-brown. Head broad (MDL/OHW 5 0.50–0.53), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes prominent, not compressed; gula flat to slightly concave; antennomere ratios: length I:II:III 5 2.1:1.0:2.2, width VII:VIII:IX 5 1.0:1.0:1.7. Pronotum very large, broad (PNL/PNW 5 0.80–0.81), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad (SEL/ ELW 5 0.97–1.12); lateral margins strongly rounded, apically rounded; punctation similar to pronotum; sutural stria present in apical one-fifth to one-half of elytron. Flight wings strongly reduced to absent. Mesosternum broad, flattened; medial carina present, but low and not strongly developed. Metasternum relatively narrow (MTL/MTW 5 0.14– 0.16), flattened, sloping dorsad anteriorly; oblique femoral carinae moderately prominent, low and convex medially. Male tarsi 5-5-4; proand mesobasotarsomeres somewhat laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur moderately broad with posteroapical tooth and series of smaller teeth, apical margin rounded to truncate (fig. 154); metasternal fovea large, transversely oval, medial, with cluster of long, dense, fine setae. Median lobe in lateral aspect moderately slender, moderately long, strongly curved basally, moderately expanded submedially on dorsal margin, apical portion directed slightly dorsally, straight, slightly expanded medially, apex pointed (fig. 297); in ventral aspect slender, slightly constricted medially, distinctly expanded submedially, apical portion narrowed to constricted neck, thereafter expanded and apex rounded (fig. 296); operculum long and slender in lateral aspect, apically pointed (fig. 297), in ventral aspect long with lateral margins subparallel, apex emarginate (fig. 296); lateral lobes slender, strongly curved basally, sinuate, expanded medially, apex curved and slender to narrowly rounded apex (fig. 298). Female tarsi 5-4-4. E TYMOLOGY : This species is named for the Aztecs, a group of people who engaged in human sacrifice, dominated central Mexico through military subjugation of their enemies, and were slaughtered and forced to serve Spain by the conquistador Cortez. D ISTRIBUTION : This species is known from central and southern Mexico (fig. 368). P ARATYPES : MEXICO: Chiapas: 8kmSESan Cristobal, 28 Sep 1986, 2400 m, litter at logs, fungus, pine-oak forest, on fungus, R Baranowski (3, LUND). Oaxaca: 40.5 km S Suchixtepec, 25 Jul 1992, 1300 m, cloud forest leaf litter, Berlese, RS Anderson (3, CNCI); 6 mi S Valle Nacional, 19 May 1971, 2000 9 , leaf litter, S Peck (1, PECK); 40 km SW Valle Nacional km 93, 26 Jul 1992, 1900 m, oak forest leaf litter, RS Anderson (2, CNCI); 5 mi S Valle Nacional, 19 May 1971, 2000 9 , leaf litter, S Peck (1, PECK). Puebla: 24 km N Xicotepec de Juarez, 17 Jun 1983, oak forest litter, R Anderson (1, PECK). Vera Cruz: 4.4 mi N Huatusco, 29 Jul 1973, 4200 9 , cloud forest, Newton (1, FMNH); 5 km NE Coscomatepec, 22 Jun 1983, 1130 m, cloud forest litter, Anderson and Peck (9, PECK); 5 km NE Coscomatepec, 22 Jun 1983, 1130 m, cloud frest litter, Anderson and Peck (1, PECK); 6 mi NE Catemaco, 7 Jul 1976, 1500 9 , rain forest leaf litter, Newton (1, FMNH); 7 km E Huatusco, 22 Jun 1983, cloud forest litter, Anderson and Peck (2, PECK); Lake Catemaco, 1 May 1969, JM Campbell (1, CNCI). D ISCUSSION : This species has been collected from cloud forest litter and litter in pine and oak forests. Elevation records are from 1500 to 4200 ft. Agathidium iridescens Miller and Wheeler, new species Figures 155, 299–302, 367 T YPE M ATERIAL : Holotype, ? in FMNH labeled ‘‘MEX: S.L.P.14 mi.W Xilitla 4800 9 2005 129MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II VI.29.73 A.Newton/leaf & log litter non-coniferous Liquidambar for/HOLOTYPE Agathidium iridescens Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, San Luis Potosi, 14 mi W Xilitla, 4800 9 . D IAGNOSIS : This species is extremely similar externally to A. aztec and related species. It is dorsally iridescent in all specimens examined. The median lobe is diagnostic and has the operculum long and narrow with the apex slightly emarginate, with each ramus apically rounded (fig. 299). Other differences are mainly slight shape differences in the apical portion of the median lobe (fig. 300). D ESCRIPTION : Body relatively large (TBL 5 2.79–3.94 mm), broad, robust (PNW/TBL 5 0.46–0.50), strongly contractile. Head and pronotum dark red-brown; elytra dark red -brown, conspicuously iridescent, especially laterally; venter red-brown; antennae, palpi, and legs red-brown. Head broad (MDL/OHW 5 0.53–0.58), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes prominent, not compressed; gula slightly convex; antennomere ratios: length I:II:III 5 1.6:1.0:2.0, width VII:VIII:IX 5 1.0:1.0:2.1. Pronotum very large, broad (PNL/PNW 5 0.79–0.80), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.86–1.04); punctation similar to pronotum; sutural stria present in apical one-fifth of elytron. Flight wings present, fully developed. Mesosternum broad, flattened; medial carina reduced anteriorly. Metasternum moderately broad (MTL/MTW 5 0.16), flattened, not or only gently sloping dorsad anteriorly; oblique femoral carinae not prominent, obsolete medially. Male tarsi 5-5-4; proand mesobasotarsomeres moderately expanded laterally, with moderately large field of ventral spatulate setae; mandibles not modified; metafemur moderately broad with small subapical tooth along ventral margin (fig. 155); metasternal fovea transversely linear, with linear series of fine setae. Median lobe in lateral aspect slender, strongly curved basally, abruptly expanded along dorsal margin submedially, apical portion directed at angle dorsad, slender slightly expanded subapically, apex slightly curved ventrad, pointed (fig. 301); in ventral aspect slender, laterally expanded submedially, apical portion strongly and evenly narrowed, very slender, apex slightly expanded, apically rounded (figs. 299, 300); operculum in lateral aspect long, slender, straight, apex somewhat expanded (fig. 301), in ventral aspect long, nearly extending to apex of median lobe, lateral margins convergent to near apex, apex slightly expanded laterally, emarginate medially, each ramus slightly flared laterally (fig. 299); lateral lobes long, slender, strongly curved basally, sinuate apically, expanded medially and apex spatulate with 2 prominent setae (fig. 302). Female not examined. E TYMOLOGY : This species is named from the Latin word iridis, meaning ‘‘rainbow’’ or ‘‘iridescent’’, for the conspicuous iridescent coloration of the elytron in this species. D ISTRIBUTION : This species is moderately widespread in central and southern Mexico (fig. 367). P ARATYPES : MEXICO: Durango: 35 mi W El Salto, 23 Jul 1964, HF Howden (3, CNCI). Hidalgo: 4 mi SW Chapalhuacan, 3 Jul 1973, 3500 9 , on ‘‘fungus logs’’, A Newton (1, FMNH); Tianchinol, 43 km SW Huejutla, 14 Jun 1983, 1500 9 , cloud forest litter, S and J Peck (1, PECK). Oaxaca: 52 mi N Oaxaca, 17 May 1971, 9500 9 , leaf litter, S Peck (1, PECK); 33 km S Valle Nacional, 10 Sep 1986, 1600 m, litter, cloud forest, R Baranowski (1, LUND). Puebla: 5 mi NE Teziutlan, 16 Jul 1973, 5000 9 , cloud forest leaf litter, Berlese, A Newton (1, FMNH). Queretaro: 18 mi E Landa Matamoros, 11 Jun 1971, 5500 9 , leaf litter, S Peck (2, PECK); 17–18 mi E Landa de Matamoros, 30 Jun 1973, oak leaf and log litter, A Newton (1, FMNH); 17–18 mi E Landa de Matamoros, 28 Jun 1973, 5300 9 , under bark, A Newton (10, CNCI). San Luis Potosi: 14 mi W Xilitla, 29 Jun 1973, 4800 9 , leaf and log litter nonconiferous Liquidambar forest, A Newton (2, FMNH); 40 km W Xilitla, 6 Aug 1983, 1700 m, pine -oak forest litter, S and J Peck (6, PECK). D ISCUSSION : Specimens have been collected from litter in a variety of forest types including oak, pine, and Liquidambar cloud 130 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY forest and leaf and log litter. Elevation records are from 1500 to 9500 ft. Agathidium dentigerum Subgroup D ISCUSSION : This group of species is characterized by the operculum of the median lobe completely divided into two long, independent rami (e.g., fig. 309), extensive endophallic armature consisting of an elongate medial, triramous process with a medial, apically truncate lobe and lateral, flattened hyaline lobes, the base of the lateral lobes distinctly sinuate in ventral aspect (e.g., fig. 309), relatively narrow metasternum medially with moderately large male metasternal fovea, somewhat reduced eyes that are dorsoventrally compressed and ovoid with large facets, and a large, acutely pointed, occasionally somewhat falcate male metafemoral tooth. The group may be further divided into two groups based on the presence or absence of a prominent lobelike tubercle medially on the gula. Other than this, the species are very similar externally with the only other significant differences being the relative development of the medial mesosternal and oblique metasternal carinae and the development of the lobelike process posteromedially on the metasternum. Male genitalia are generally diagnostic and male specimens should be dissected for most positive identification of specimens. The species are eastern Nearctic in distribution. They have been most commonly collected by sifting various litter types and Berlese extraction. The reduced eyes, strong contractility, and winglessness of many specimens suggest they are cryptic in habitats living in litter and in the substrate. The only previously described species in this group is A. dentigerum Horn. Since the species in this subgroup are externally very similar, it is not surprising that Horn (1880) apparently included specimens in A. dentigerum that belong in what we describe here as new species. As additional collecting in the eastern United States ensues, we expect more species of this subgroup to come to light since the species appear to be numerous, cryptic in habitat, and relatively poorly collected. Agathidium dentigerum Horn Figures 156, 303–305, 371 Agathidium dentigerum Horn, 1880: 303; Leng, 1920; Fall, 1934b. T YPE M ATERIAL : Lectotype (designated to fix this name with this species), ? in MCZC labeled ‘‘Stone Cr. LeeCo Va/269. [handwritten]/Type 3187 [number handwritten, two-thirds of label red]/A. dentigerum Horn [handwritten]’’. There is a single / paratype (not examined) with same label data as the holotype (Horn, 1880). T YPE L OCALITY : United States, Virginia, Lee Co., Stone Creek. D IAGNOSIS : This species is similar to other A. dentigerum group members that have a large, acute, somewhat falcate male metafemoral tooth (fig. 156), very narrow metasternum with the oblique metasternal carinae relatively prominent and meeting medially in a large, posteriorly directed triangular lobe, large male metasternal fovea, and somewhat reduced eyes. However, this species has a very prominent lobelike tubercle medially on the gula, making it most similar to A. akrogeneios and A. stephani. From these species it may be distinguished by the shape of the median lobe which has the apical portion strongly recurved dorsally in lateral aspect (fig. 304), much more broadly so than in A. akrogeneios (fig. 310). D ESCRIPTION : Body small (TBL 5 2.17– 2.23 mm), rounded, robust (PNW/TBL 5 0.47–0.48), strongly contractile. Head, pronotum, and elytra red; venter yellow-red; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.52–0.59), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very finely microreticulate; frontoclypeal suture obsolete medially; eyes not strongly reduced, protruding, large-faceted; gula broad, flattened, with prominent median lobelike, tubercle; antennomere ratios: length I:II:III 5 1.3:1.0:0.9, width VII:VIII:IX 5 1.0:1.0:2.0. Pronotum very large, broad (PNL/PNW 5 0.71–0.74), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures 2005 131MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II smooth, shiny. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ ELW 5 0.94–1.07); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, broadly convex; medial carina prominent. Metasternum narrow (MTL/MTW 5 0.14– 0.15), medially flat, distinctly sloping dorsad anteriorly; oblique femoral carinae not prominent laterally, medially meeting in very prominent, posteriorly directed triangular flange. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur slender, with large acute, subfalcate tooth subapically on posterior margin (fig. 156); metasternal fovea posterior, large, round with large brush of long, fine setae. Median lobe in lateral aspect moderately broad, basally bent, straight thereafter, apical portion tapered, strongly sinuate, apex finely pointed (fig. 304); in ventral aspect moderately broad, lateral margins slightly curved, apical portion narrowed to broadly pointed apex (fig. 303); operculum divided into two long, slender rami, apices slightly divergent and each narrowly rounded (fig. 303); lateral lobes broad, curved basally, apices rounded with 2 long subapical setae (fig. 305). Female not examined. D ISTRIBUTION :Agathidium dentigerum is found in Tennessee and Virginia (fig. 371). S PECIMENS E XAMINED : UNITED STATES: Tennessee: Campbell Co.: Morely, 17 Mar 1976, rotten wood debris, Berlese, QD Wheeler (2, QDWC). D ISCUSSION : A single habitat record is from ‘‘rotten wood debris’’. Agathidium stephani Miller and Wheeler, new species Figures 157, 306–308, 369 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘ALA:Walker Co. Jasper, 12.X.80 T. King, litter [handwritten]/HOLOTYPE Agathidium stephani Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : United States, Alabama, Walker Co., Jasper. D IAGNOSIS : This species is similar to other A. dentigerum group members that have a large, acute, male metafemoral tooth (fig. 157), very narrow metasternum with the oblique metasternal carinae relatively prominent and meeting medially in a large, posteriorly directed triangular lobe, large male metasternal fovea and somewhat reduced eyes. However, this species has a very prominent lobelike tubercle medially on the gula, making it most similar to A. akrogeneios and A. dentigerum. From these species it may be distinguished by the shape of the median lobe, which is nearly straight in lateral aspect from near the base to the sharply pointed apex (fig. 307). D ESCRIPTION : Body small (TBL 5 1.90– 2.22 mm), rounded, robust (PNW/TBL 5 0.43–0.48), strongly contractile. Head, pronotum and elytra yellow to yellow-red; venter yellow to yellow-red; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.62–0.63), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very finely microreticulate; frontoclypeal suture obsolete medially; eyes moderately reduced, protruding, large-faceted, dorsoventrally somewhat compressed; gula broad, flattened, with small but prominent median lobelike tubercle; antennomere ratios: length I:II:III 5 1.2:1.0:1.0, width VII:VIII:IX 5 1.0:1.0:2.1. Pronotum very large, broad (PNL/PNW 5 0.77–0.82), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth, shiny. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.87–1.09); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, broadly convex; medial carina very prominent, anteriorly obscured on some specimens. Metasternum very narrow medially (MTL/MTW 5 0.08–0.11), medially slightly concave, distinctly sloping dorsad anteriorly; oblique femoral carinae not prominent laterally, medially meeting in very prominent, posteriorly directed triangular flange. Male tarsi 5-5-4; proand mesobasotar- 132 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 369. Geographic distribution of Agathidium oniscoides-group species: A. akrogeneios 5m ;A. stephani 5v . someres slightly laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur slender, with large acute tooth subapically on posterior margin (fig. 157); metasternal fovea posterior, prominent, round with large brush of long, fine setae. Median lobe in lateral aspect slender, basally bent, straight thereafter, apical portion tapered, straight to slightly dorsally bent, pointed apex (fig. 307); in ventral aspect moderately broad, lateral margins broadly curved, apex rounded (fig. 306); operculum divided into two long, slender rami, apices slightly divergent and each narrowly rounded (fig. 306); lateral lobes moderately broad, evenly curved in lateral aspect, apex pointed or narrowly rounded with 2 long subapical setae (fig. 308). Female tarsi 5-4-4. E TYMOLOGY : This species is named in honor of K. Stephan, the collector of some of the type specimens. D ISTRIBUTION : This species is found in south-central North America from Oklahoma to Kentucky and south to Louisiana and Alabama (fig. 369). P ARATYPES : UNITED STATES: Alabama: Jackson Co.: McAllister Sink Cave, 10 Aug 1980, cave litter, T King (1, CNCI); 6 mi N Princeton outside Horseshoe Cave, 30 Jun 1967, Berlese, S Peck, A Fiske (1, CNCI); Walker Co.: Jasper, 12 Oct 1980, litter, T King (1, CNCI); Jasper, 12 Oct 1980, litter, T King (2, PECK). Arkansas: Garland Co.: 3 mi W Crystal Springs, 1 Jun 1984, 800 9 , hardwood litter, J Pakaluk (1, AMNH); Logan Co.: Cove Lake RA, 16 Jun 1990, Berlese, CE Carlton (3, LSAM); 1 km E Lookout, Mt Magazine, 23 May 1986, JM Campbell (11, CNCI); Mt 2005 133MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Magazine Lookout, 26 May 1986, deciduous litter, JM Campbell (9, CNCI); Brown Springs, Mt Magazine, 23 May 1986, leaf litter along stream, JM Campbell (4, CNCI); Mt Magazine, Cameron Bluffs, 27 May 1986, leaf litter, JM Campbell (3, CNCI); Madison Co.: Withrow Spr. St Park, 27 Oct 1974, 1400 9 , mixed harwood forest ground litter, Berlese, AF Newton (5, CNCI); Montgomery Co.: 1.5 mi E Crystal Rec. Area, slopes along FS 177K, 6 Jul 1991, deciduous forest, Berlese, CE Carlton (5, LSAM); 2 mi NW Mount Ida, 1 Jun 1984, 800 9 , hardwood litter, J Pakaluk (4, AMNH); 1 mi SE Joplin, 1 Jun 1984, 700 9 , litter, J Pakaluk (1, AMNH); E Crystal Rec. Area at FS 177K, 24 Mar 1992, deciduous forest, Berlese, CE Carlton (1, LSAM); 1 mi SE Joplin, 1 Jun 1984, 700 9 , litter near stream, J Pakaluk (2, QDWC); Newton Co.: Ozark NF, FSR 92096B, 6 km NE Pelsor, 13 May 1986, oak hickory leaf litter, JM Campbell (2, CNCI); Ozark NF, Alum Cove Natl Br. 4 km NW jct Hwy 7 and 16, 13 May 1986, JM Campbell (1, CNCI); Ozark NF, Buck Br. FSR 1209, 3 mi N Hwy 123, 14 May 1986, flood debris on bank, JM Campbell (1, CNCI); Polk Co.: Rich Mt Pioneer Cemetary, 17 Feb 1988, Berlese, RT Allen (1, LSAM); Shady Lake Rec. Area, 13 Oct 1974, 1200 9 , mixed hardwood-pine forest ground litter, Berlese, AF Newton (2, CNCI); Rich Mt Eagleton Overlook, 17 Feb 1988, Berlese, RT Allen (1, LSAM); Pope Co.: Ozark NF jct Hwy 7 and FSR 1828, 12 May 1986, JM Campbell (1, CNCI); Pulaski Co.: Pinnacle Mt St Park NE Face near top, 11 Nov 1987, decid., Berlese, CE Carlton (9, LSAM); Scott Co.: 14 mi SE Waldron, 31 May 1984, forest litter, J Pakaluk (2, AMNH); Washington Co.: 3 mi S Devils Den St Park, 28 May 1979, oak hickory, S and J Peck (1, CNCI); Devils Den St Park, 8 Jun 1969, litter, Berlese, S and J Peck (8, QDWC). Kentucky: Edmonson Co.: Mammoth Cave Natl Park., Cabin Woods, 11 Apr 1979, W Suter (5, CNCI); Mammoth Cave Natl Park, Doyle Valley, 30 Aug 1967, log litter, Berlese, S Peck, A Fiske (1, CNCI). Louisiana: Rapides Par. Magnolia Rec. Area, 15 mi SW Alexandria, 4 Oct 1973, pine/ hardwood leaf litter forest floor, A Newton (1, CNCI). Oklahoma: Clayton, 21 Jun 1983, leaves and pine needles, BF and JL Carr (3, CARR); Laflore Co.: 10 mi SW Heavener winding stair watch tower, 10 Jun 1969, 1600 9 , log litter, S Peck (1, CNCI); Latimer Co.: 5 mi W Red Oak, Dec 1980, K Stephan (1, KSIC); Apr 1983, K Stephan (20, QDWC); Robbers Cave St Park, Lost Lake, 3 Jul 1987, rotten pine, DS Chandler (1, CNCI); Nov 1983, K Stephan (4, KSIC); 25 Feb 1985, litter, grass, FIT, K Stephan (48, KSIC); McCurtain Co.: Beaver Bend St Park, 27 Jul 1968, dry rivulet, W Suter (1, PECK). D ISCUSSION :Agathidium stephani has been collected in a wide variety of habitats, including leaves and pine needles, oak, hickory, pine, grass and various other litter types, cave litter, and debris on a river bank. Elevation records are from 800 to 1200 ft. Agathidium akrogeneios Miller and Wheeler, new species Figures 158, 309, 310, 369 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘GA: Dade Co., Cloud land Canyon St. Prk. 16.v.1972, S.&J. Peck Rhodo. Litter,B236/HOLOTYPE Agathidium akrogeneios Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : United States, Georgia, Dade Co., Cloudland Canyon State Park. D IAGNOSIS : This species is similar to other A. dentigerum subgroup members that have a large, acute, somewhat falcate male metafemoral tooth (fig. 158), very narrow metasternum with the oblique metasternal carinae relatively prominent and meeting medially in a large, posteriorly directed triangular lobe, large male metasternal fovea, and somewhat reduced eyes. However, this species has a very prominent lobelike tubercle medially on the gula, making it most similar to A. stephani and A. dentigerum. From these species it may be distinguished by the shape of the median lobe, which is straight in lateral aspect from near the base to near the apex where it is abruptly recurved dorsally (fig. 310). D ESCRIPTION : Body small (TBL 5 2.36– 2.45 mm), rounded, robust (PNW/TBL 5 0.45–0.51), strongly contractile. Head, pronotum, and elytra red; venter red to yellow-brown; antennae, palpi, and legs yellow to yellow-brown. Head broad (MDL/OHW 5 0.54–0.62), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes moderately prominent and protruding, but dorsoventrally compressed, large-faceted; gula broad, flattened, with prominent median lobelike tubercle; antennomere ratios: length I:II:III 5 1.6:1.0: 1.0, width VII:VIII:IX 5 1.0:1.0:2.3. Pron- 134 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY otum very large, broad (PNL/PNW 5 0.70– 0.76), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth, shiny. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.86–1.15); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, broadly convex; medial carina prominent. Metasternum narrow (MTL/ MTW 5 0.10–0.15), slightly concave medially, distinctly sloping dorsad anteriorly; oblique femoral carinae not prominent laterally, medially meeting in very prominent, posteriorly directed triangular flange. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur slender, with very sharp, acuminate, large tooth subapically on posterior margin (fig. 158); metasternal fovea posterior, prominent, round with large brush of long, fine setae. Median lobe in lateral aspect slender, basally bent, straight thereafter, apical portion tapered, straight, apex abruptly and sharply recurved dorsally (fig. 310); in ventral aspect moderately broad, lateral margins broadly curved, apex abruptly narrowed and with medial, narrowly rounded prominence (fig. 309); operculum divided into two long, slender rami, apices slightly divergent and each narrowly rounded (fig. 309); lateral lobes moderately broad, evenly curved in lateral aspect, apex pointed or narrowly rounded with 2 long subapical setae (fig. 310). Female tarsi 5-4-4. E TYMOLOGY : This species is named akrogeneios from the Greek word meaning ‘‘with a prominent chin’’, for the prominent gular tubercle in this species. D ISTRIBUTION : This species is found in southeastern North America from Texas to Georgia and north to Virginia (fig. 369). P ARATYPES : UNITED STATES: Alabama: Jackson Co.: 5 mi NW Princeton, 19 May 1972, forest litter, S Peck (1, PECK); 5 mi N Garth, 19 May 1972, S Peck (8, PECK); Madison Co.: Monte Sano St. Park, 17 May 1972, S and J Peck (3, PECK). Georgia: Dade Co.: Cloudland Canyon St. Park., 16 May 1972, Rhododendron litter, S and J Peck (2, PECK). Kentucky: Edmonson Co.: Mammoth Cave Natl Park, Doyle Valley, 30 Aug 1969, log litter, S Peck, A Fiske (1, PECK). Louisiana: W Feliciana Co.: Magnolia Glen TNC property near Tunica WMA, 16 Nov 1996, mixed mesophytic, Berlese, CE Carlton (24, LSAM); Tunica WMA Magnolia Glen TNC Property, 10 mi NW St Francisville, 25 Mar 1998, mesophytic forest, Berlese, CE Carlton (19, LSAM); Cabin area 30 8 47 9 N, 91 8 15 0 W, 1 Feb 2001, pitfall, AR Cline (3, LSAM). Tennessee: Hardin Co.: 0.5 mi W Olive Hille, 28 May 1991, old growth, Berlese, C Carlton (3, LSAM). Texas: Sabine Co.: 9 mi E Hemphill, 11 May 1988, beech, magnolia litter, Berlese, R Anderson (2, LSAM). Virginia: Lee Co.: Cumberland Gap Natl Park. nr Skyline Cave, 10 Jul 1971, dry litter at log, W Shear (1, AMNH). D ISCUSSION : This species has been collected from February to November. It has been found in a variety of habitats, including ‘‘mesophytic forest’’, and various litter types, including logs, Rhododendron, beech, and magnolia. Agathidium framea Miller and Wheeler, new species Figures 117, 311–313, 373 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘N.CAR:Wilkes Co. Jeffress Park, 3500 9 BlueRidgePkwy, mi 272 17.VIII.1981 S. Peck log—leaf litter/HOLOTYPE Agathidium framea Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : United States, North Carolina, Wilkes Co., Jeffress Park, Blue Ridge Parkway mi 272, 3500 9 . D IAGNOSIS : This species is distinguishable from most species by the relatively reduced eyes (dorsoventrally compressed and large - faceted), relatively indistinct oblique metasternal carinae which meet medially in a very small, indistinct triangle, unmodified gula, the male with a distinct, broad lobe along posterior margin of coxa (fig. 117), and very distinctive male genitalia. The apical portion of the median lobe is long and apically expanded into an arrowhead shape in ventral aspect (fig. 311). D ESCRIPTION : Body small (TBL 5 1.93– 2.12 mm), rounded (PNW/TBL 5 0.4), robust, strongly contractile. Head, pronotum, and elytra red; venter 2005 135MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II Figs. 370, 371. Geographic distribution of Agathidium oniscoides-group species: 370, A. pocahontasae.371, A. appalachium 5m ;A. dentigerum 5v . yellow to yellow-brown; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.52), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very finely microreticulate; frontoclypeal suture obsolete medially; eyes strongly compressed, reduced to elongate triangle; gula broad, flattened, with prominent rim; antennomere ratios: length I:II:III 5 1.8:1.0: 1.3, width VII:VIII:IX 5 1.0:1.0:1.6. Pronotum very large, broad (PNL/PNW 5 0.73– 0.75), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures slightly microreticulate. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.02–1.06); punctation similar to pronotum; sutural stria indistinctly present only apically. Flight wings strongly reduced. Mesosternum broad, broadly convex; medial carina present, well developed. Metasternum narrow (MTL/MTW 5 0.09–0.10), medially concave, slightly sloping dorsad anteriorly; oblique femoral carinae present, low and indistinct, meeting medially in small, triangular, posteriorly directed flange. Male tarsi 5-5-4; proand mesobasotarsomeres moderately laterally expanded, with moderately large field of ventral spatulate setae; mandibles not modified; metafemur broad, with exceptionally large, flat, triangular tooth subapically on posterior margin (fig. 117); metasternal fovea large, transverse, broadly oval with large, conspicuous brush of fine, dense setae; posteromedial margin of metacoxa with broad, flat, rounded 136 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Figs. 372, 373. Geographic distribution of Agathidium oniscoides-group species: 372, A. gallititillo. 373, A. framea 5v ;A. georgiaense 5m . lobe. Median lobe in lateral aspect slender, strongly curved basally, moderately straight thereafter, apical portion slender, tapered, apex sharply and abruptly bent dorsad, apex pointed and often slightly recurved (fig. 312); in ventral aspect slender, long, lateral margins subparallel to sinuate, apical portion tapered, apically expanded into broad, arrowhead shape, apex rounded to medially pointed (fig. 311); operculum divided into two long, slender, straight rami, rami sometimes slightly apically expanded, apices pointed; endophallic armature consisting of long, flat, broad medial lobe that is apically truncate and slightly emarginate medially and lateral hyaline lobes on each side of median lobe (fig. 311); lateral lobes long, slender, apically slightly expanded, basally, along with basal piece, strongly sinuate in ventral aspect (fig. 313). Female tarsi 5-4-4. E TYMOLOGY : This species is named from the Latin word framea, meaning ‘‘spear’’, for the spear-shaped apex of the median lobe of the aedeagus in this species. D ISTRIBUTION : This species has been collected from Georgia, North Carolina, Tennessee, and Virginia (fig. 373). P ARATYPES : UNITED STATES: Georgia: Rabun Co.: Setolah, 29 May 1983, Rhododendron and mixed leaf litter, DS Chandler (1, DENH). North Carolina: Black Mts (3, AMNH); Valley of Black Mts, 29 Jul 1906, W Beutenmuller (4, AMNH); Alleghany Co.: Roaring Gap Stone Mt St Park Rd, 18 Aug 1981, 2000 9 , leaf litter, S Peck (14, CNCI); Cumberland Knob, Blue Ridge Parkway, mi 218, 19 Aug 1981, 2700 9 , forest leaf lit- 2005 137MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II ter, S Peck (7, CNCI); Avery Co.: Grandfather Mt Blue Ridge Parkway, mi 304, 17 Aug 1981, 4000 9 , leaf litter, S Peck (17, CNCI); Linville Falls, Blue Ridge Parkway, mi 317, 16 Aug 1981, 3500 9 ,Rhododendron litter, S Peck (1, CNCI); Linville Falls, Blue Ridge Parkway mi 337, 16 Aug 1981, 3500 9 ,Rhododendron litter, S Peck (6, PECK); Henderson Co.: 0.3 mi SW Bat Cave, 22 Jul 1967, 1000 9 , litter, S Peck, A Fiske (5, PECK); Macon Co.: Highlands, QD Wheeler (1, QDWC); Mitchell Co.: Roan Mtn, 25 Jul 1967, 5000 9 , log litter, S Peck, A Fiske (2, PECK); Wilkes Co.: Jeffres Park, Blue Ridge Parkway mi 272, 17 Aug 1981, 3500 9 , leaf litter, S Peck (8, CNCI); Jeffress Park, Blue Ridge Parkway mi 272, 17 Aug 1981, 2500 9 , leaf litter, S Peck (3, PECK); Yancy Co.: Black Mts, Blue Ridge Parkway mi 152, 15 Aug 1981, 4900 9 , leaf litter, S Peck (4, CNCI). Tennessee: Cocke Co.: Great Smoky Mts NP, Albright Grove Tr 83 8 16 9 45 0 W, 35 8 44 9 10 0 N, 19 Oct 2001, 1000 m, decid stump, Berlese, CE Carlton (21, LSAM); 6 mi SE Cosby, 31 May 1983, forest litter, DS Chandler (7, DENH); Sevier Co.: Great Smoky Mts NP, Porters Creek Trail 82 8 23 9 52 0 W, 35 8 40 9 13 0 N, 18 Oct 2001, 870 m, old growth, Berlese, T Tischechkin, A Cline (4, LSAM). Virginia: Tazewell Co.: E River Mtn Summit 1 mi E Rd 662, 2 May 1971, 3300 9 ,Rhododendrum, oak, gum, hemlock litter, W Shear (1, AMNH). D ISCUSSION : This species has been collected from a variety of litter types, including oak, gum, hemlock, Rhododendron, and logs. Elevation records are from 1000 to 5000 ft. Agathidium appalachium Miller and Wheeler, new species Figures 159, 314–316, 371 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘VA: Wise Co 1.5 mi S. Norton [handwritten]/rot. stump S. Peck, ber 133 1500 9 [handwritten]/HOLOTYPE Agathidium appalachium Miller and Wheeler, 2003 [red label with black line border]’’. Only a single specimen is known of this species. T YPE L OCALITY : United States, Virginia, Wise Co., 1.5 mi S Norton. D IAGNOSIS : This species can be distinguished from most species by its very small size, reduced eyes, lack of a medial lobe on the anterior margin of the gula, relatively large metafemoral tooth (fig. 159), and distinctive median lobe which has the apex truncate in ventral aspect and the operculum relatively short (fig. 314). D ESCRIPTION : Body minute (TBL 5 1.89 mm), oblong (PNW/TBL 5 0.51), strongly contractile. Head and pronotum light red; elytra red, not iridescent; venter yellow-red; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.55), dorsal surface flattened, dorsoventrally compressed; with extremely fine punctures, surface between punctures shiny, smooth; frontoclypeal suture obsolete medially; eyes reduced, dorsoventrally compressed, and elongate, large faceted; gula slightly concave, unmodified; antennomere ratios: length I:II:III 5 1.2:1.0: 0.9, width VII:VIII:IX 5 1.1:1.0:2.1. Pronotum very large (PNL/PNW 5 0.71), broad, strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with minute, sparse punctures, each with a short, very fine seta, surface between punctures smooth. Elytra broad, lateral margins strongly rounded, slightly elongate, but not apically acuminate (SEL/ELW 5 0.97); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, rounded; medial carina well developed. Metasternum very narrow medially (MTL/MTW 5 0.16), flattened, medially strongly dorsally sloped anteriorly; oblique femoral carinae present and distinct, meeting medially in rounded flange. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small field of ventral spatulate setae; mandibles not modified; metafemur moderately broad, with strongly produced, flattened, thornlike subapical tooth on posterior margin (fig. 159); metasternal fovea prominent, transversely ovoid, with pencil of long, fine, dense setae. Median lobe in lateral aspect slender, slightly curved basally, straight thereafter, apical portion slender, apically distinctly curved dorsad (fig. 315); in ventral aspect slender, slightly widened medially, apical portion long, slender, tapered to abruptly truncate apex (fig. 314); operculum moderately small, broadly V-shaped, each ramus very slender, apically narrowly rounded (fig. 314); lateral lobes moderately robust, in lateral aspect broad, apically tapered to narrowly rounded apex, without long subapical setae (fig. 316). Female not examined. E TYMOLOGY : Named after the Appalachian 144 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY 5 2.88–3.23 mm), rounded, robust (PNW/ TBL 5 0.44–0.49), strongly contractile. Head, pronotum, and elytra yellow-red; venter yellow; antennae, palpi, and legs yellow to yellow-brown. Head broad (MDL/OHW 5 0.52–0.54), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, with fine microreticulation on most specimens; frontoclypeal suture obsolete medially; eyes prominent, protruding, somewhat dorsoventrally compressed, largefaceted; gula concave; antennomere ratios: length I:II:III 5 1.6:1.0:1.3, width VII:VIII: IX 5 1.0:1.0:1.6. Pronotum very large, broad (PNL/PNW 5 0.73–0.90), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures smooth, with fine microreticulation on many specimens. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ ELW 5 0.92–0.99); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, broadly convex; medial carina completely obscured. Metasternum narrow (MTL/MTW 5 0.16–0.17), medial area slightly convex, surface with numerous fine setae, slightly sloped dorsad anteriorly; oblique femoral carinae reduced, rounded, medially with narrowly rounded, posteriorly directed flange. Male tarsi 5-5-4; proand mesobasotarsomeres slightly laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur moderately broad, with acuminate, sharp, large tooth on posterior margin (fig. 164); metasternal fovea large, transversely oval, with large, dense brush of long, fine setae. Median lobe in lateral aspect moderately robust, strongly bent basally, relatively straight thereafter, slightly expanded medially, apical portion broad, evenly tapered to pointed, slightly dorsally curved apex (fig. 330); in ventral aspect broad, medially somewhat expanded, apical portion evenly narrowed to rounded apex (fig. 329); operculum divided into two long, slender rami, apices slightly divergent and each narrowly rounded (fig. 329); lateral lobes moderately broad, bent basally, straight thereafter, apices narrowly rounded with 2 long subapical setae (fig. 331). Female not examined. E TYMOLOGY : This species is named in honor of G.W. Bush (Crawford, TX), President of the United States. D ISTRIBUTION : This species is found in Ohio, Virginia, and North Carolina (fig. 375). P ARATYPES : UNITED STATES: North Carolina: Valley of Black Mts, 18 Aug 1906, W Beutenmuller (1, AMNH); Mt Michell, 4000 9 (1, MCZC). Ohio: Adams Co.: 2 mi SE Blue Creek, 8 Aug 1976, forest litter, Berlese, LE Watrous (1, QDWC). D ISCUSSION : Habitat records include forest and leaf litter. A single altitude record is from 4000 ft. Agathidium georgiaense Miller and Wheeler, new species Figures 165, 332–334, 373 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘GA: Dade Co., Cloud land Canyon St. Prk. 16.v.1972, S.&.J.Peck Rhodo. litter,B236/HOLOTYPE Agathidium georgiaense Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : United States, Georgia, Dade Co., Cloudland Canyon State Park. D IAGNOSIS : This species is very similar to several species in this subgroup. It is nearly identical to A. gallititillo (see ‘‘Diagnosis’’ under that species) from which it differs mainly in the shape of the male genitalia. Whereas the apical portion of the median lobe is deflected ventrad in A. gallititillo (fig. 324), it continues in approximately the same line as the medial portion of the median lobe in A. georgiaense (fig. 333). The proand mesobasotarsomeres are relatively broadly expanded in males of this species. D ESCRIPTION : Body moderately large (TBL 5 2.31–2.68 mm), rounded, robust (PNW/ TBL 5 0.50–0.51), strongly contractile. Head and pronotum red to dark red; elytra dark red; venter, antennae, palpi, and legs yellow to yellow-brown. Head broad (MDL/OHW 5 0.52–0.54), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punc- 2005 145MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II tures shiny, with fine microreticulation; frontoclypeal suture obsolete medially; eyes prominent, protruding, slightly dorsoventrally compressed, large-faceted; gula concave anteriorly, slightly swollen medially; antennomere ratios: length I:II:III 5 1.3:1.0:1.0, width VII:VIII:IX 5 1.0:1.0:1.9. Pronotum very large, broad (PNL/PNW 5 0.63–0.68), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, with fine microreticulation. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.99–1.07); punctation similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum broad, broadly convex; medial carina well developed. Metasternum very narrow medially (MTL/MTW 5 0.09– 0.10), medial area slightly convex, slightly sloped dorsad anteriorly; oblique femoral carinae obscured, low, rounded, medially with broad, flat, subtriangular, posteriorly directed flange. Male tarsi 5-5-4; proand mesobasotarsomeres somewhat laterally expanded, with small ventral field of spatulate setae; mandibles not modified; metafemur slender, subapical posterior tooth moderately large, triangular, acutely pointed (fig. 165); metasternal fovea large, transversely oval, with large brush of long fine setae. Median lobe in lateral aspect long, slender, curved basally, moderately straight thereafter, apical portion long, slender, tapered, in same line of curvature as medial portion, apically bent dorsad, very finely pointed, without small dorsal prominence (fig. 333); in ventral aspect slender, somewhat expanded medially, apical portion slightly constricted, expanded apically, apex broadly rounded (fig. 332); operculum divided, rami long but ending well short of apex of median lobe, slender, relatively straight through most of length, with apices narrowly rounded and divergent (fig. 332); lateral lobes slender, long, bent basally, straight in apical portion (fig. 334). Female not examined. E TYMOLOGY : This species is named after the state of origin of the type specimens. D ISTRIBUTION : This species is known only from the type locality in Georgia (fig. 373). P ARATYPES : UNITED STATES: Georgia: Dade Co.: Cloudland Canyon St. Park, 16 May 1972, Rhododendron litter, S and J Peck (8, PECK). D ISCUSSION : The type series was collected from Rhododendron litter. Agathidium microphthalmum Subgroup D ISCUSSION : This species subgroup is united by small body size, the very strongly reduced eyes, the large male metafemoral tooth, and distribution in southern Mexico and Guatemala. The three species are externally fairly similar but may be distinguished by the small size and shape of the eyes (figs. 110–112) and by the shape of the male genitalia with the median lobe lacking lateral carinae and sulci and the lateral lobes apically straight (figs. 335–343). Agathidium chauliodoum Miller and Wheeler, new species Figures 112, 166, 335–337, 376 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘GUAT: BAJA VERAPAZ: 7.5 km S Purulha. Elev. 1630 m.cloud for. litter. R. Anderson 91-26, 26-V-1991/HOLOTYPE Agathidium chauliodoum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Guatemala, Baja Verapaz, 7.5 km S Purulha. D IAGNOSIS : Males of this species may be distinguished from most species by the strongly reduced eyes consisting of dorsally directed ovoids (fig. 112), and by the large metafemoral tooth (fig. 166) and distinctive male genitalia with the median lobe stout and apically abruptly curved and sharply pointed in lateral aspect (fig. 336). The species is most similar to A. microphthalmum and A. nimbosilva, from which it may be distinguished by slightly larger eyes and differences in the shape of the median lobe and the more broadly expanded male probasotarsomeres. D ESCRIPTION : Body small (TBL 5 2.21– 2.43 mm), broad, robust (PNW/TBL 5 0.43– 0.52), rounded, strongly contractile. Head piceous; pronotum piceous, testaceous around margins; elytra piceous, slightly iridescent laterally; venter dark red; antennae, palpi, and legs red-brown. Head broad (MDL/OHW 5 0.58–0.69), 146 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Fig. 376. Geographic distribution of Agathidium oniscoides-group species: A. chauliodoum 5m ;A. microphthalmum 5v ;A. nimbosilva 5. . dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes strongly reduced to ovoids, directed dorsally (fig. 112); gula flat or slightly convex; antennomere ratios: length I:II:III 5 1.6:1.0:1.7, width VII:VIII:IX 5 1.0:1.0:1.6. Pronotum very large, broad (PNL/PNW 5 0.70–0.74), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.79–1.14); punctation and surface similar to pronotum; sutural stria present only apically. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.15– 0.18), slightly concave medially, strongly dorsally sloped anteriorly; oblique femoral carinae well developed, meeting medially in broad, broadly truncate flange. Male tarsi 5-5-4; proand mesobasotarsomeres distinctly laterally expanded, protarsomeres more strongly so and with large ventral field of spatulate setae; mandibles not modified; metafemur broad, with very large, flat, triangular tooth subapically on posterior margin (fig. 166); metasternal fovea moderate in size, transverse, with pencil of dense, fine, long setae. Median lobe in lateral aspect broad and robust, abruptly curved basally, apical portion short, slender, strongly hooked ventrally, apex sharply pointed (fig. 336); in ventral aspect robust, widened medially, apical portion broadly triangular (fig. 335); operculum flat, broad, lateral margins slightly divergent apically, apex broadly rounded (fig. 335); endophallic armature comprised of a main medial hyaline lobe that is apically truncate, a long hyaline lobe on each side, a strongly diverging, long hyaline lateral lobe on each side and a long dorsolateral hyaline lobe on each side; lateral lobes long, slender, 2005 147MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II strongly curved, apices directed medially, apex slightly expanded, with 2 stout setae subapically (fig. 337). Female not observed. E TYMOLOGY : Named chauliodoum from the Greek word for ‘‘having prominent teeth’’ for the very large teeth on the posterior margin of the metafemora. D ISTRIBUTION : This species is known only from Baja Verapaz, Guatemala (fig. 376). P ARATYPES : GUATEMALA: Baja Verapaz: 14.5 km S Purulha, 26 May 1991, 1600 m, riparian bottomland oak forest litter, R Anderson (1, CNCI); 3 km SW Purulha, 20 Nov 1991, litter, cloud forest, R Baranowski (19, JRAC); 7 km E Purulha, 22 May 1991, 1600 m, cloud forest litter, RS Anderson (1, CNCI); 8 km S Purulha, 20 May 1991, 1600 m, pine-cloud forest litter, RS Anderson (1, CNCI). D ISCUSSION : This species has been collected from cloud forest litter and riparian bottomland oak forest litter. Altitude records are from 1600 m. Agathidium microphthalmum Miller and Wheeler, new species Figures 110, 167, 338–340, 376 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘MEXICO: Chiapas: Volcan Tacana, lower slopes, ca. 4 km N Union Juarez, 19IX-1992. R.S. Anderson 92-110/HOLOTYPE Agathidium microphthalmum Miller and Wheeler, 2003 [red label with black line border]’’. T YPE L OCALITY : Mexico, Chiapas, Volcan Tacana, lower slopes, ca. 4 km N Union Juarez. D IAGNOSIS : Males of this species may be distinguished from most species by the strongly reduced eyes consisting of minute, slightly elongate triangles (fig. 110), the large metafemoral tooth (fig. 167), and distinctive male genitalia with the median lobe moderately slender and with the apex hooked and slightly expanded in lateral aspect (fig. 339). The species is most similar to A. chauliodoum and A. nimbosilva, from which it may be distinguished by smaller eyes, less laterally expanded male probasotarsomeres (to distinguish it from A. chauliodoum), and differences in the shape of the median lobe (figs. 338, 339). D ESCRIPTION : Body small (TBL 5 2.39 mm), broad, robust (PNW/TBL 5 0.48), rounded, strongly contractile. Head dark red-brown; pronotum dark redbrown medially, red along margins; elytra dark red-brown medially, red along margins, not iridescent; venter, antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.53–0.55), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes strongly reduced to small triangles, only a few facets present (fig. 110); gula flattened, unmodified; antennomere ratios: length I:II: III 5 2.0:1.0:1.4, width VII:VIII:IX 5 1.0: 1.0:1.9. Pronotum very large (PNL/PNW 5 0.71–0.72), broad, strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 1.00–1.12); punctation and surface similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.12–0.15), flat medially, distinctly dorsally sloped anteriorly; oblique femoral carinae well developed, meeting medially in low, but prominent carina. Male tarsi 5-5-4; proand mesobasotarsomeres distinctly laterally expanded, about equally so, each with moderate field of ventral spatulate setae; mandibles not modified; metafemur moderately broad, with very large, flat, triangular tooth subapically on posterior margin (fig. 167); metasternal fovea minute with small pencil of fine setae. Median lobe in lateral aspect stout, strongly curved basally, straight in medial portion apically flexed dorsad, apical portion slender, straight, apex abruptly expanded with dorsal and ventral points (fig. 339); in ventral aspect parallel-sided for most of length, apex evenly tapered to narrowly rounded apex (fig. 338); operculum broad, flat, apically broadly rounded (fig. 338); lateral lobes long, slender, 148 NO. 291BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY strongly curved basally, apically with 2 stout setae (fig. 340). Female tarsi 5-4-4. E TYMOLOGY : This species is named from the Greek words micro, meaning ‘‘small’’, and ophthalmos, meaning ‘‘eye’’, in reference to the very reduced eyes in this species. D ISTRIBUTION : This species is known only from the type locality in Chiapas (fig. 376). P ARATYPES : MEXICO: Chiapas: Volcan Tacana, lower slopes, 4 km N Union Juarez, 19 Sep 1992, 2000 m, cloud forest litter, RS Anderson (5, CNCI). D ISCUSSION : The type specimens were collected from cloud forest litter at 2000 m. Agathidium nimbosilva Miller and Wheeler, new species Figures 111, 168, 341–343, 376 T YPE M ATERIAL : Holotype, ? in CMNC labeled ‘‘GUATEMALA: BAJA VERAPAZ: 7 km E Purulha. Elev. 1600 m. cloud for. litter. R. Anderson 91-15 23-V-1991/HOLOTYPE Agathidium nimbosilva Miller and Wheeler, 2003 [red label with black line border]’’. Only a single specimen is known of this species. T YPE L OCALITY : Guatemala, Baja Verapaz, 7 km E Purulha. D IAGNOSIS : Males of this species may be distinguished from most species by the strongly reduced eyes consisting of minute, slightly elongate triangles (fig. 111), a small but distinct and sharp metafemoral tooth (fig. 168), and distinctive male genitalia with the median lobe moderately robust and with the apex slender and approximately straight (fig. 342). The species is most similar to A. chauliodoum and A. microphthalmum, from which it may be distinguished by smaller eyes, less laterally expanded male probasotarsomeres (both to distinguish it from A. chauliodoum), a smaller metafemoral tooth and differences in the shape of the median lobe (figs. 341, 342). D ESCRIPTION : Body small (TBL 5 2.16 mm), broad, robust (PNW/TBL 5 0.49), rounded, strongly contractile. Head dark red-brown; pronotum dark redbrown medially, red along margins; elytra dark red-brown medially, red along margins, not iridescent; venter, antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.55), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, very lightly microreticulate; frontoclypeal suture obsolete medially; eyes strongly reduced to small triangles, only a few facets present (fig. 111); gula flattened, not modified; antennomere ratios: length I:II:III 5 1.2:1.0:1.7, width VII:VIII:IX 5 1.0:1.0:2.2. Pronotum very large, broad (PNL/PNW 5 0.74), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, smooth. Elytra broad, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.98); punctation and surface similar to pronotum; sutural stria absent. Flight wings strongly reduced. Mesosternum moderately broad, not declivitous; medial carina well developed. Metasternum narrow (MTL/MTW 5 0.12), flat medially, distinctly dorsally sloped anteriorly; oblique femoral carinae well developed, meeting medially in low, but prominent carina. Male tarsi 5-5-4; proand mesobasotarsomeres not strongly laterally expanded, equally so and with a moderate field of ventral spatulate setae; mandibles not modified; metafemur moderately broad, with very large, flat, triangular tooth subapically on posterior margin (fig. 168); metasternal fovea moderately large, transversely oval with dense brush of long fine setae. Median lobe in lateral aspect moderately short, stout, strongly curved basally, straight in medial portion apically flexed dorsad, apical portion slender, straight, apex slender and pointed (fig. 342); in ventral aspect parallel-sided for most of length, apex evenly tapered to narrowly rounded apex (fig. 341); operculum broad, flat, apically broadly rounded (fig. 341); lateral lobes long, slender, strongly curved basally, apically with 2 stout setae (fig. 343). Female not observed. E TYMOLOGY : Named from the Latin words nimbus, meaning ‘‘rain cloud’’, and silva, 2005 149MILLER AND WHEELER: SLIME-MOLD BEETLES, PART II meaning ‘‘forest’’, for the habitat where the type specimen was collected. D ISTRIBUTION : This species is known only from the type locality in Baja Verapaz, Guatemala (fig. 376). D ISCUSSION : The type specimen was collected from cloud forest litter at 1500 m. Agathidium oniscoides Subgroup D ISCUSSION : These species are not characterized by any particularly discrete characters, but share a general similarity in body form and aedeagal shape. There are two species pairs in this group. One of these is A. oniscoides and A. rubellum, which have nearly identical male genitalia and relatively short elytral sutures. The other two are A. fawcettae and A. exiguum, which have dorsal microreticulation, relatively long elytral sutures and somewhat similar genitalia. These are some of the most common species in North America. Agathidium oniscoides Palisot de Beauvois Figures 169, 344–347, 377 Agathidium oniscoides Palisot de Beauvois, 1817: 160; LeConte, 1853; Horn, 1880; Matthews, 1887; Leng, 1920; Fall, 1934b; Wheeler, 1990 (description of larva). Agathidium globatile LeConte, 1878: 598; Horn, 1880 (synonymized with A. oniscoides); Leng, 1920. Agathidium piceum Melsheimer, 1844: 103; LeConte, 1853 (synonymized with A. oniscoides), Matthews, 1887; Leng, 1920, nec Erichson, 1845. T YPE M ATERIAL :Agathidium oniscoides: type (presumably in Paris) not examined. Agathidium globatile: lectotype (designated here to clarify assignment of this name to this species), / in MCZC labeled ‘‘Mic./613. [handwritten]/Type 3180 [number handwritten, two-thirds of label red]/A. globatile Lec. [handwritten]/oniscoides 6 [handwritten]’’. Paralectotype, 1 / in MCZC labeled ‘‘Marquett 29-1 Mich/13’’. LeConte had multiple specimens on which he based his description, and other specimens accompany these two but are not clearly part of the type series. Agathidium piceum: holotype (sex not determined) in MCZC labeled ‘‘[circular pink disc]/Agathidium oniscoides Beau piceum Mels. [handwritten]’’. T YPE L OCALITY :Agathidium oniscoides: United States, South Carolina. Agathidium globatile: United States, Michigan, Detroit [here restricted]. Agathidium piceum: United States, Pennsylvania. D IAGNOSIS : This species is the largest in the genus in North America and is often very black in color, making it easily distinguishable from most species. The shape of the median lobe is distinctive with the apical portion long, directed at an angle dorsad, slightly expanded medially along dorsal margin in lateral aspect (fig. 346), and with the operculum narrow and long with the apex slightly to moderately laterally expanded (fig. 344). The male metafemoral tooth is small (fig. 169) and the metasternal fovea is moderately large. The sutural stria extends about onefifth to one-third the length of the elytron. This species is extremely similar to A. rubellum, from which it can be distinguished by larger size, darker coloration, and larger metasternal fovea. D ESCRIPTION : Body moderate to very large (TBL 5 2.57–4.74 mm), oblong (PNW/TBL 5 0.44–0.54), strongly contractile. Head, pronotum, and elytra dark red to piceous; venter yellow to yellow-red; antennae, palpi, and legs yellow. Head broad (MDL/OHW 5 0.61–0.64), dorsal surface flattened, dorsoventrally compressed; with very fine punctures, each with a short, very fine seta, surface between punctures shiny, with fine microreticulation; frontoclypeal suture obsolete medially; gula without tubercle; eyes rounded, not compressed, prominent, small-faceted; gula slightly convex; antennomere ratios: length I: II:III 5 2.2:1.0:2.3, width VII:VIII:IX 5 1.0: 1.0:1.6. Pronotum very large, broad (PNL/ PNW 5 0.65–0.82), strongly convex, anterolateral lobes strongly produced, lateral margin broadly curved, not angulate; with very fine, sparse punctures, each with a short, very fine seta, surface between punctures shiny, with fine microreticulation. Elytra broad, slightly elongated, lateral margins strongly rounded, apically rounded (SEL/ELW 5 0.88–1.01); punctation similar to pronotum; sutural stria extending from apex to one-fifth to one-third length of elytron. Flight wings fully developed in most specimens, some [Document text truncated for crawler view.]