SLIME-MOLD OF THE GENUS AGATHIDIUM PANZER IN NORTH AND CENTRAL AMERICA, PART I. COLEOPTERA:

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])

KELLY B. MILLER Department of Entomology, Cornell University Current address: Department of Integrative Biology Brigham Young University Provo, Utah 84042 ([email protected])

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 290, 95 pp., 53 ®gures, 1 table Issued March 24, 2005

Copyright ᭧ American Museum of Natural History 2005 ISSN 0003-0090 Male Agathidium pulchrum LeConte habitus, dorsal and lateral. Original by Byron Alexander. CONTENTS Abstract ...... 4 Introduction ...... 4 Introduction to Series ...... 6 Methods and Concepts ...... 6 Phylogenetic Concept ...... 6 Disarticulation and Dissection Techniques ...... 8 Observation of Specimens ...... 8 Natural History ...... 9 Collecting Agathidium ...... 16 Morphology ...... 17 Adult ...... 17 Male Sexual Dimorphism ...... 24 Male Genitalia ...... 28 Female Sexual Dimorphism ...... 30 Female Genitalia ...... 30 Pupa ...... 30 ...... 30 Key to Genera of Agathidiini of North and Central America ...... 33 Cainosternum Notman ...... 33 Anisotoma Panzer ...... 34 Stetholiodes Fall ...... 34 Agathdium Panzer ...... 34 Key to Species Groups of Agathidium ...... 37 Agathdium sexstriatum species group ...... 39 Key to A. sexstriatum Species Group ...... 41 A. bistriatum Horn...... 41 A. sexstriatum Horn...... 43 A. estriatum Horn ...... 45 Agathidium brevisternum Species Group ...... 48 Key to A. brevisternum Species Group ...... 48 A. brevisternum Fall ...... 49 A. rhinocerellum Wheeler and Miller, n.sp...... 52 A. dioperculum Wheeler and Miller, n.sp...... 54 Agathidium revolvens Species Group ...... 58 Key to A. revolvens Species Group ...... 58 A. revolvens LeConte ...... 59 A. jasperanum Fall ...... 63 A. depressum Fall ...... 65 A. dubitans Fall ...... 67 A. dubitanoides Wheeler and Miller, n.sp...... 69 A. omissum Fall ...... 70 A. cavisternum Fall ...... 72 A. virile Fall ...... 74 A. conjunctum Brown ...... 79 A. angustoperculum Wheeler and Miller, n.sp...... 81 A. falcatoperculum Wheeler and Miller, n.sp...... 83 Checklist of Agathidium Species: Part 1 ...... 85 Acknowledgments ...... 85 References ...... 86 Appendix ...... 94

3 ABSTRACT The external morphology of the adult, egg, larva, and pupa of Agathidium oniscoides Beau- vois is described. The natural history of the genus is reviewed, including associations with Myxomycetes and fungi and techniques for ®eld collection and laboratory study. The phylo- genetic species concept is discussed and applied in this study. A key and brief diagnoses are provided for the genera of Agathidiini (Coleoptera: Leiodidae: ) of North and Central America. Volvoxis Kugelann is regarded as a nomen oblitum, and its junior synonym Agath- idium Panzer is regarded as the valid name (nomen protectum) for this taxon by invoking article 23.9 of the International Code of Zoological Nomenclature. A key to species groups of Agathidium Panzer is presented. Species are revised for the A. sexstriatum, A. brevisternum, and A. revolvens species groups, including keys, diagnoses, descriptions, and ®gures of im- portant morphological features. Lectotypes are designated for A. bistriatum Horn and A. sexst- riatum Horn. The following new species are described: A. angustoperculum, n.sp., A. diop- erculum, n.sp., A. dubitanoides, n.sp., A. falcatoperculum, n.sp., and A. rhinocerellum, n.sp.

INTRODUCTION cal) species of the genus are far less convex. The genus Agathidium Panzer (Coleop- Interestingly, Agl. laevis has been found in tera: Leiodidae: Leiodinae) includes minute association with slime-mold plasmodia (Ste- to small beetles (1±6 mm) that are generally phenson et al., 1994), whereas Neotropical round, convex, shiny, usually concolorous, relatives of that species appear to be feeders and frequently capable of contracting their on agarics (Wheeler, 1979a). The contractil- body into a compact ball-like shape when ity of various species of Agathidium no disturbed. Most species for which biological doubt contributed to one common name for data exist are associated with slime-molds the Leiodidae, the ``round fungus-beetles'' (Myxomycetes or ), in either their (e.g., Arnett, 1971). mature (sporocarp) or immature (plasmodi- The genus Agathidium is the largest pos- um) stages. Unlike related genera in Agath- sibly monophyletic group of docu- idiini, Agathidium has also been associated mented to be principally slime-mold feeding. with a substantial number of fungi, primarily Such myxomycophagous habits are so wide- Basidiomycetes. The full extent of the geo- spread within the genus and related genera graphic range of the genus is not yet known, of Agathidiini (ϭ Anisotomini) that it is rea- although most species inhabit temperate or sonable to infer such associations with montane tropical forests of the Northern Myxomycetes were present in the common Hemisphere. ancestor of both the genus and the tribe When the body is contracted, the antennae (Newton, 1984; Wheeler, 1984a, 1984b). are retracted into ventral cephalic antennal Myxomycophagy has also been documented grooves, the legs are folded beneath the in Aglyptinus (see above) and in the southern body, and the broad lateral extensions of the temperate tribe Neopelatopini in the Leiodi- pronotum fold over the sides of the elytra to dae, and occurs outside leiodids sporadically achieve, in its ultimate expression, a nearly across the Coleoptera. Putative slime-mold completely spherical form. Such contractility associates include certain Rhysodidae, Clam- is seen only rarely among phylogenetically bidae, Eucinetidae, Scaphidiinae (Staphylin- unrelated Coleoptera, including Clambidae, idae), Cerylonidae, Latridiidae (ϭ Lathridi- Nitidulidae (Cybocephalinae), and Scara- idae), and Sphindidae (Blackwell, 1984; baeidae (Cryptocephalinae). Within the Lawrence and Newton, 1980; McHugh, Leiodidae, this kind of defense is approached 1993; Russell, 1979; Stephenson et al., 1994; only by the single North American species Wheeler, 1979b, 1984a, 1984b, 1987). of Aglyptinus Cockerell, Agl. laevis LeConte, Agathidium is remarkably diverse in both but not by its numerous tropical congeners. morphological structure and ecological hab- In the case of Agl. laevis, however, the body its. Species vary dramatically in size (rang- is extremely convex and hemispherical but ing from barely 1 mm to more than 6 mm not contractile, whereas other (mostly tropi- when fully extended), body shape (from con-

4 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 5 vex and noncontractile to elongate-oval and (Brown, 1928, 1930, 1933; Fall, 1901). To- partially contractile, to broadly rounded, day, more than 50 years have passed since highly convex, and fully contractile), pres- Fall's work, and a similar situation exists. ence/absence of certain male sexual dimor- Little taxonomic work has been done since phisms (i.e., enlarged basal tarsomeres on Fall's revision (Arnett, 1971; Hatch, 1957), front and middle legs, a setose fovea on the and a considerable number of undescribed metasternum, a tooth near the posteroapical species have accumulated in collections. The angle of the metafemur, and a tusk on the left current expansion in numbers of species is mandible), coloration (from reddish-brown to the result of three factors: (1) discovery of a black and, rarely, with striking color pat- large complex of ¯ightless species in the Ap- terns), and punctation (from nearly impunc- palachian Mountains; (2) species of Mexico tate to confused punctation to serial puncta- and Central America have nearly been ig- tion of elytra). Ecologically, Agathidium is nored to date; and (3) characters of the ae- one of few myxomycophagous taxa with spe- deagus, which are extremely valuable diag- cies that have been found associated with nostic features at the species level, have not both spores of mature fruiting bodies and the been previously examined. Only four species plasmodial stage of the hosts (Wheeler, have been described from Latin America 1984a, 1984b, 1987). Although ®rm host as- (Hendrichs, 1979; Matthews, 1887) despite sociations have not been made for a complex evidence from collections that suggests the of anophthalmic and microphthalmic and genus is very diverse in montane regions in ¯ightless species in the Appalachian Moun- this area. tains of the eastern United States, phyloge- Our interest in the genus was a natural netic af®nity, modi®cations of the mouth- continuation of the senior author's previous parts, and ®eld observations are consistent work on the related genus Anisotoma Panzer with plasmodial predation in these species (Wheeler, 1979b), and it was intensi®ed dur- (Wheeler, 1984a). ing studies of material at the American Mu- Palearctic species number about 200 and seum of Natural History in 1974 when he are comparatively well known (Angelini, ®rst observed an undescribed microphthal- 1995), whereas those of other regions contin- mic and ¯ightless species endemic to the ue to be described in large numbers. About southern Appalachian Mountains of the Unit- 350 species have been described from the ed States. A subsequent ®eld trip in the Oriental region where large numbers of spring of 1975 resulted in rediscovery of this ¯ightless endemic forms are known. As an and a related microphthalmic species. These example, 50 out of 52 species of Agathidium species appear to be specialized predators of known from Taiwan are endemic to that is- plasmodia, living in the rich, deep, moist leaf land (Angelini and de Marzo, 1995). Where- litter of high elevation forests with signi®- as substantial progress has been made on the cant levels of precipitation (Wheeler, 1984a, faunas of these regions, that of the New 1984b). Such aptery and microphthalmy is World has remained largely unstudied (An- highly unusual among fungivores in general gelini, 1988, 1990; Angelini and de Marzo, and slime-mold beetles in particular. Inter- 1983, 1984, 1987a, 1987b, 1988, 1990, estingly, ¯ightless species of Dasycerus 1995). Brongniart (Staphylinidae) also exist in this When Fall (1934: 99) revised the North region and feed upon a range of fungus hosts American species, he stated that ``A recent (Wheeler, 1984c; Wheeler and McHugh, critical survey of the Agathidium material in 1994). my collection has revealed the presence of a Whereas this taxonomic study will in- considerable number of undescribed species. crease our knowledge of the species diversity There is nothing surprising in this fact when of the Agathidium of North and Central it is recalled that the last published treatment America, it is no more than the next logical of the genus was that of Dr. Horn in his Sil- step in the scienti®c exploration of this fas- phidae paper of more than ®fty years ago.'' cinating clade. Knowledge of the species di- In fact, between the works of Horn and Fall, versity of Agathidium is far from complete. only four new species had been described Additional species remain to be discovered 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 in the United States, particularly in the Ap- ®eld studies in areas of the U.S., Mexico, and palachian and Ozark regions and in the coast- Central and northern South America in- al forests of northern California, Washington, crease, many additional species of Agathi- and Oregon. Further, based on the relative dium await discovery and description. Just as paucity of specimens examined from Latin recent work by Angelini (see references) has America and the proportion of undescribed greatly increased our knowledge and esti- species among them, it is clear that no more mates of the species diversity of the genus in than a small percentage of those species that Europe and Asia, this study will suggest that live in montane, temperate forests of Mexico the diversity of American Agathidium has and Central America have been collected. It been seriously underestimated in the past. can be hoped only that this work will facil- itate identi®cation of known species and en- METHODS AND CONCEPTS courage the collection of those not yet known, so that another revision may be done PHYLOGENETIC SPECIES CONCEPT: The bio- in the future to more completely document logical species concept (BSC) has dominated the diversity of this remarkable genus. the zoological literature for half a century (Mayr, 1942, 1963, 1982). Despite consid- INTRODUCTION TO SERIES erable controversy over modes of speciation based on models, the BSC has re- This is the ®rst of two publications that mained nearly unquestioned in the literature will collectively constitute a taxonomic re- on insects. Never fully embraced by bota- vision of the species of Agathidium of North nists, the BSC has recently been widely crit- and Central America. This part provides an icized on general conceptual grounds (Cra- introduction to the genus Agathidium, a de- craft, 1983; Cronquist, 1978; de Queiroz and tailed account of the external morphology of Donoghue, 1988; Donoghue, 1985; Mishler a representative highly contractile species (A. and Donoghue, 1982; van Valen, 1976). In oniscoides), a key to species groups, and re- particular, the emergence of rigorous phylo- visions of the A. brevisternum, A. revolvens, genetic theories has indicated the need for a and A. sexstriatum species groups. The sub- species concept more in line with the goals sequent part will include taxonomic treat- of phylogenetic analyses and classi®cations ments of the remaining species groups. (Wheeler and Meier, 2000). One conse- All species in this study will be interpreted quence of the adoption of phylogenetic the- within the phylogenetic species concept as ory is the need to recognize elements ana- discussed by Nixon and Wheeler (1990, lyzable by cladistic methods and not ones 1992), Wheeler and Nixon (1990), Wheeler that are themselves divisible into smaller and Platnick (2000a, 2000b), and Platnick units with this property (Nixon and Wheeler, and Wheeler (2000) as modi®ed from Eld- 1990). Stated another way, we need to iden- redge and Cracraft (1980), Nelson and Plat- tify the smallest units for which unequivocal nick (1981), and Cracraft (1983) (see also historical evidence of phylogeny may be re- Cracraft, 1992). This concept, discussed be- trieved. Thus, since the appearance of Hen- low, can be anticipated to increase the num- nig's phylogenetic theory, cladists have ber of species where the biological species sought with increasing intensity a ``phylo- concept has been subjectively applied and genetic'' concept of species. used to recognize as subspecies those popu- Hennig (1966) modi®ed the BSC to cor- lations that are holomorphologically distinct rect for ambiguity associated with time of and more or less allopatric (Nelson and Plat- speciation. The result, an ``inter-nodal'' con- nick, 1981). In the case of Agathidium, how- cept, represented an advance (Ridley, 1989) ever, species have historically been based on but fell short of a clearly phylogenetic con- a morphological concept and closely approx- cept. Rosen (1978) introduced an autapo- imate phylogenetic species. morphic concept describing species as mini- This monograph will serve as a stepping- mum autapomorphic units (see also de Quei- stone toward a complete accounting of the roz and Donoghue, 1988; Hill and Crane, species of the genus for the New World. As 1982). Autapomorphic species, however, are 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 7 based on the inappropriate use of phyloge- case of sexually reproducing , traits netic analyses and application of concepts of vary among individuals within a population monophyly and synapomorphy below the whereas characters are constant among all in- level of species. Such attempts to analyze de- dividuals of one or more populations. How- monstrably tokogenetic relationships through ever, constancy does not equate to invari- cladistic analyses are misguided at best ability. A subapical tooth is frequently pre- (Wheeler and Nixon, 1990). sent on the metafemur of male Agathidium, This study uses the phylogenetic species yet the structure of that tooth varies from concept (PSC), introduced by Eldredge and small straight structures to large curved ones. Cracraft (1980) and Nelson and Platnick As Cracraft (1983) stated, another goal of (1981), and more recently discussed by Cra- a species concept is to recognize the end craft (1983, 1992) and ampli®ed and extend- products of evolution. Because evolutionary ed by Nixon and Wheeler (1990, 1992) and history results from unpredictable interac- Wheeler and Platnick (2000b). Stated simply, tions among many biotic and abiotic forces, phylogenetic species are ``the smallest aggre- references to ``the'' evolutionary process are gation of populations (sexual) or lineages illusory. Given the existence of many evo- (asexual) diagnosable by a unique combina- lutionary processes, the advantages of rec- tion of character states in comparable indi- ognizing historical patterns independent of viduals (semaphoronts)'' (Nixon and Wheel- any one of them become apparent. Regard- er, 1990). The PSC is consistent with phy- less of the mode of speciation (e.g., allopat- logenetic theory, but independent of phylo- ric, parapatric, sympatric, allochronic), the genetic analysis. This is necessary since one distribution of characters provides a mecha- goal of the PSC is to identify the elements nism for recognizing their end products. for cladistic studies. Thus, the PSC serves also as a unit species Hennig (1966) distinguished between to- concept, making comparisons of numbers of kogenetic and phylogenetic relationships. To- species in different groups, and resulting kogenetic or birth relationships are shared by from different speciation processes, mean- individuals within sexually reproducing pop- ingful. Because clonal systems result in new ulations and result in more or less reticulate species with each mutation, numbers of spe- patterns of relationship. Phylogenetic rela- cies in asexual groups have been grossly un- tionships are shared among species, involve derestimated by the BSC. It is logical, how- character transformation, and are hierarchic ever, that such a system would result in in pattern. In evolutionary history, hierarchic greater numbers of end products than would relationships exist because characters of an sexual systems. ancestral species are inherited by its descen- The PSC has numerous advantages. Most dants, either in their original or some modi- importantly, it is based on observable char- ®ed form (Platnick, 1979). Application of acters so that the PSC is imminently testable cladistic analysis to populations may result and it provides logical elements among in a ``cladogram''. However, where evidence which cladistic relationships may be ana- or inference of tokogenetic relationships ex- lyzed. For those who are compelled to build ists, such a diagram may be artifactual rather some ontological construct for species, the than phylogenetic. The assumption that the PSC provides a logical interface between the outcome of such an analysis may be inter- reticulate patterns of descent observed preted as a hierarchy is not valid. Thus, a among individuals within populations of sex- species concept is desired that can distin- ually reproductive organisms and the unique guish among species prior to the cladistic sequence of patterns of common ancestry analysis. shared among species belonging to a clade. Nixon and Wheeler (1990, 1992) drew a The PSC gives to us both the maximum to- strict contrast between the terms character kogenetic construct and the minimum phy- and trait. Traits are heritable attributes that logenetic element. For us, the epistemologi- vary within terms of a cladogram whereas cal linkage of the PSC to characters is not a characters are heritable attributes that are philosophical weakness, but a strength. Its constant (®xed) throughout each term. In the de®nition makes clear its connection in prac- 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 tice to empirical data. Just as for cladistic treated similarly except for the following. analysis, the only ontological concern that The entire abdomen was removed from the must be addressed is the assumption that relaxed specimen and heated in 10% KOH. phylogenetic species share a common history While rinsing in H2O, the apical two seg- (i.e., that evolution has occurred). ments of the abdomen were pulled from the DISARTICULATION AND DISSECTION TECH- basal segments. This effectively removes NIQUES: Con®rmation of species identity for both the ovipositor and spermatheca. After many Agathidium beetles requires examina- additional clearing, these structures were tion of the male aedeagus. Males can usually gently removed from the terminal segments be sorted from females by one or more of and studied under cover glass. the following dimorphisms: (1) enlargement For detailed comparative morphological of pro- and mesobasotarsomeres and/or pres- work, entire specimens were often disartic- ence of spatulate ventral setae on basotarso- ulated. In such cases, the entire specimen meres; (2) presence of an apical or preapical was placed in KOH and the major somatic tusk or horn on the dorsum of the left man- segments were teased apart. Components of dible; (3) presence of a metasternal fovea, the mouthparts, antennae, and legs were re- varying from single or paired punctiform moved and examined. The sclerites of such structures to much larger transverse oval preparations, thoroughly rinsed in water, eroded areas bearing more or less long and were stored in glycerin in microcups cut frequently dense setae; or (4) presence of a from glass vials and stored in ``Watrous preapical or apical tooth on the posterior trays'' (Wheeler and McHugh, 1987). margin of the metafemur. Examination of the OBSERVATION OF SPECIMENS: The convex protarsus is usually suf®cient. In their ex- shape and subtle differences in dorsal punc- treme form, the basal segments are distinctly tation of many Agathidium make their study expanded with large, dense patches of spat- and illustration dif®cult. Certain conventions ulate setae beneath. In their least developed were adopted for this study that standardize form, however, the tarsi are only indistinctly both our observations and the presentation of laterally expanded. Even in these individuals, certain characters in scienti®c illustrations. however, a small number of ventral, spatulate Mention of these conventions will facilitate setae are visible at moderate magni®cations. effective use of this publication and compar- Once identi®ed, male specimens were re- ison of species. laxed by heating them in water with a mild An outline drawing showing the dorsal detergent. Once thoroughly relaxed, the ely- body shape is presented for each species. The tra were teased apart, exposing the membra- foreshortening of the curvature of the body nous tergum of the abdomen. The tergum combined with the degree of contraction of was then incised by a minuten nadeln, and the body in its state of preservation make the aedeagus was removed from the abdo- simultaneous observation of both the head men by the points of minute forceps, a and the elytra nearly impossible for many hooked insect pin, or minuten nadeln. The species. Thus, the dorsal drawings are actu- elytra were then replaced over the incision ally composites. For the head, the specimen and the specimen was dried and remounted. was tilted backward until the base of the la- Meanwhile, the aedeagus was heated gently brum was just visible. For the elytra, the in a 10% potassium hydroxide solution to specimen was tilted forward until the apex macerate adhering muscle tissues. It was then of the lateral bead was just visible. By so rinsed completely in distilled water and doing, the shape of the cranium and elytra placed in glycerin or a 50:50 % mixture of are both communicated in a single drawing, glycerin:glycerin jelly. Structures were then although in practice orientation of the spec- placed in a depression slide and examined imen will have to be changed to examine with a Leitz Dialux 20 or 22 compound mi- these structures. croscope with Nomarski differential interfer- The appearance of the punctation is in¯u- ence contrast illumination. enced to a considerable extent by the quality Female specimens, when dissected for of lighting. Our observations and drawings study of ovipositor and spermatheca, were were done using a very bright ®ber optic 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 9 light source. A Mylar ®lm shield was used than they are to true fungi. Given cues that to diffuse the light. Under such light, it is far can vary among myxomycete clades and spe- easier to discern serial rows of punctures that cies, the plasmodium wells upward, forming nearly blend in with the intervening, random- mature sporocarps over a period of a few to ly distributed punctures under less diffuse 24 or more hours. light. We strongly recommend viewing punc- Agathidium are, in general, found in moist tation under such conditions when the pres- forests. On a broad geographic scale, Agath- ence or absence of subtle punctation differ- idium are found in temperate forests of north- ence is in question. ern latitudes and southern montane regions. To achieve such diffuse lighting, two ap- The genus is diverse in most forested areas proaches were used. For illustrations, where of the northern hemisphere. On a local, hab- maximum diffusion was important and where itat level, Agathidium are most easily found specimens did not have to be moved once set in dense forests where logs and litter are into position for drawing, a tube was fash- damp at the ground level. Whereas some oth- ioned from Mylar and set around the speci- er slime-mold beetles (e.g., Latridiidae, men. Light was then shown through these Sphindidae) frequent dry microhabitats also, curved walls. For routine observation and such as in exposed ®elds, Agathidium seems description, when the specimen had to be less tolerant of low humidity situations. manipulated by hand constantly, and when North American Agathidium are found from less diffuse light was suf®cient to observe sea level to nearly 9000 feet elevation, in the punctation patterns, a piece of Mylar was case of A. estriatum in the Pinaleno Moun- bent into a loosely bowed shape and attached tains of Arizona. to the tips of the arms of the ®ber optic il- Like other myxomycophagous Coleoptera, luminator. This results in suf®cient diffusion, most documented host associations for with no interference in specimen manipula- Agathidium are with mature sporocarps. Un- tion. like most other slime-mold beetles, relation- ships with plasmodia have been documented NATURAL HISTORY in species of apparently distantly related groups of the genus (cf. Newton, 1984; Ste- Beetles of the genus Agathidium are hy- phenson et al., 1994; Wheeler, 1984a, 1984c, pothesized to be predominantly and ances- 1987). Plasmodial and sporocarpic associa- trally associated with plasmodial slime- tions have been noted also in the sister genus molds, the Myxomycetes or Mycetozoa. Anisotoma (Russell, 1979; Wheeler, 1980), Plasmodial or true slime-molds include about although plasmodial feeding is less frequent- 500 described species, most of which are ly observed and, considering the well-devel- more or less cosmopolitan in distribution oped mandibular mola structure, may be fac- (Martin and Alexopoulos, 1969). Large num- ultative. In contrast, at least those highly con- bers of mature spores are liberated by wind, tractile species related to the common North water, and into the air and envi- American species A. oniscoides may be ob- ronment where, given appropriate conditions, ligate associates of plasmodia (Newton, they germinate, releasing microscopic ¯ag- 1984; Wheeler, 1984a, 1984c). The only ellated amoeboid cells. These myxamoebae hints of similarly obligate plasmodial rela- later coalesce to form the large, multinucle- tions elsewhere are for certain cerylonids, ated plasmodium typical of Myxomycetes. eucinetids, and possibly rhysodids (Stephen- The Myxomycetes move about under bark, son et al., 1994; Vit, 1977). Generalizations between lamellae of decaying wood or de- are problematic, however, because of the ex- composing leaves, or within matrices of rot- tremely secretive habits of plasmodia under ting vegetative matter like giant amoebae, natural conditions. Plasmodia of many spe- sometimes attaining a diameter of several cies remain hidden throughout their some- feet (e.g., Wheeler, 1987). The comparison times ephemeral existence, emerging only with amoebae may be more than analogous hours before fruition. Because plasmodia oc- since one current view is that Myxomycetes cur among wet decaying leaves, within piles are more closely related to certain Protista of decomposing vegetation, between lamel- 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 ), Metatrichia ). Fuligo ), sessile sporangia ( Badhamia , Arcyria , ), plasmodiocarp, and plasmodium ( Lycogala , Fuligo ), aethalia ( showing major variations of form and life stages of Myxomycetes associated with adult and larval Ceratiomyxa Agathidium ), exosporate ( Reticularia Fig. 1. Slime-mold hosts of pseudoaethalium ( beetles. Schematic legend to taxa depicted in ®gure 2. Includes stalked sporangia ( 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 11 , schematic legend. See ®gure 1. Agathidium Fig. 2. Slime-mold hosts of 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 lae of rotting logs, and under loose-®tting In the following synopsis of reported host bark on trunks of dead trees, they are seldom associations, host data are presented in de- observed by collectors of beetles. Further, scending taxonomic order, and the geograph- once disturbed, actual feeding is rarely ob- ic regions covered by the records are indi- served. Agathidium have been reared on cated by a parenthetical letter following the plasmodia in the laboratory (Wheeler, 1987), Agathidium species (O ϭ Oriental; N ϭ Ne- but ®eld studies lag far behind circumstantial arctic; P ϭ Palearctic). Sources are cited for evidence for widespread, specialized plas- published records. Number of independent modial predation. It has been elsewhere records for each host greater than one indi- (Wheeler, 1984a, 1984c) speculated that cated in parentheses following ``fungus'' ¯ightless species of Agathidium in the south- name. Table 1 shows a summary of these ern Appalachians of the United States are data. such plasmodial feeders based on their phy- logenetic relationships and modi®cations of A. angulare (N): Myxomycetes: Brefeldia maxi- the mandibular mola that suggest a change ma, Fuligo intermedia, F. septica (2), Stemon- itis sp. (2), sp. (Cooter, 1978; Lawr- from ancestral spore-crushing (see also ence and Newton, 1980; Newton, 1984); ``fun- Lawrence, 1977; Lawrence, 1989). The se- gus'' (Newton, 1984). Comatricha sp., Stemon- nior author most recently collected such spe- itis axifera. ``Cream colored'' and cies in the summer of 1994 from wet rho- ``orange-yellow'' plasmodia. Basidiomycetes: dodendron leaf litter. Only once was he able Polyporus sp. to directly observe one of these ¯ightless A. angustoperculum, n.sp. (N): Basidiomycetes: beetles sitting upon a plasmodium. However, Polyporus anceps; P. sulfurous. such dense, wet layers of leaves are dif®cult A. annulatum (P): Basidiomycetes: Fomes fom- microhabitats to investigate. entarius (Hisamatsu, 1957). Slime-mold hosts utilized by Agathidium A. arcticum (P): Myxomycetes: undet.; Fuligo septica (Lundberg, 1960; Palm, 1951). Basid- are numerous and phylogenetically diverse, iomycetes: Hansenia abietina (2), Polyporus encompassing most higher taxa of Myxo- pinicola, Lactarius piperatus, polypore (Saalas, mycetes. These hosts include nearly the full 1917; Saalas and Schaufuss, 1923); ``fungus'' range of sizes, shapes, life stages, and spo- (3), ``mycelium under bark'' (Cooter, 1978; rocarp forms documented among slime- Franz, 1943; Jansson and Palm, 1936; Jansson molds (®gs. 1, 2), including stalked and ses- and SjoÈberg, 1932; Palm, 1951, 1959; Rehfous, sile sporangia, cushionlike aethalia, densely 1955). packed pseudoaethalia, and plasmodia. A. aristerium (N): Myxomycetes: Physarum po- The possibility of spore dispersal by lycephalum (Wheeler, 1987). Agathidium and other slime-mold beetles re- A. athabascanum (N): ``toadstools'', ``mush- mains poorly investigated. Simple, external, rooms''. A. atronitens (N): Basidiomycetes: Pleurotus os- mechanical spore dispersal has been experi- treatus (Newton, 1984). Specimens identi®ed in mentally demonstrated (see Blackwell, many collections as A. politum are actually A. 1984). Because some slime-mold spores are atronitens. It is possible that host records for known to show increased germination rates Hydnum septentrionale, Polyporus betulinus, given a pH shock, ingestion by beetles might and ``fungi'' therefore apply to this species result in the survival of fewer spores, among (Minch, 1952; Park, 1931; Hamilton, 1895). which are at least some with enhanced ger- A. atrum (P): Myxomycetes: (Jansson, 1946). mination potential. Regardless of active bee- Basidiomycetes: Polyporus squamosus, Armil- tle dispersal success, the simple act of eating laria mellea, Hypholoma fasciculare, Tricho- a sporocarp has the effect of disrupting the loma nudum, ``agaricus'' (Benick, 1952; Reh- peridium and releasing large numbers of fous, 1955; Singer, 1955); ``fungus'' (6), ``fun- gus dump'', ``fungusy branches'' (Beare, 1899; spores into the immediate environment and Donisthorpe, 1911, 1939; Fowler, 1889; Joy, into the air as well. Considering the cosmo- 1904; Polentz, 1938; Rueschkamp, 1928; Sing- politan distribution of many species of er, 1955). Myxomycetes (Martin and Alexopoulos, A. badium (P): Myxomycetes: Fuligo septica 1969), it is evident that most disperse quite (Rehfous, 1955). Basidiomycetes: Hansenia well unassisted. abietina (2), Trametes gibbosa, Stereum hirsu- 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 13

TABLE 1 Host Records for Agathidium Species (Shaded areas indicate occurrence of species on host.) 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

tum (Rehfous, 1955; Saalas, 1917; Saalas and ferruginosa, Cribraria argillacea, C. macro- Schaufuss, 1923); ``fungus'' (11), ``tree fun- carpa (?), C. vulgaris, Cribraria sp., Stemonitis gus'' (Bagnall, 1905; Donisthorpe, 1911; Kol- axifera, S. hyperopta, Trichia decipiens, undet. tze, 1901; Palm, 1951, 1959; Roubal, 1926, (Newton and Stephenson, 1990). 1927; Rueschkamp, 1928; von Wanka, 1908; A. kimberlae, n.sp. (part II) (N): Myxomycetes: West, 1925, 1940). leaf litter in proximity of plasmodium (Wheel- A. bicolor (P): ``fungus'', ``mycelium under er, 1984). bark'' (Palm, 1951, 1959). A. laevigatum (P): Myxomycetes: undet. (Jans- A. brevisternum (N): Myxomycetes: Fuligo sep- son, 1946). Basidiomycetes: Boletus variega- tica (Russell, 1979). tus, Fomes fomentarius (2), Polyporus sp. A. californicum (N): Myxomycetes: Fuligo sep- (Donisthorpe, 1939; Gusmann, 1925; Lindroth tica (Russell, 1979). and Palm, 1934; Platonoff, 1942); ``fungus'' A. compressidens (N): Myxomycetes: Arcyria (7) (Johnson, 1890; Lentz, 1879; Polentz, 1938; denudata; Comatricha-like. Richards, 1926; Roubal, 1927; von Wanka, A. confusum (P): Myxomycetes: Reticularia ly- 1908). coperdon (Donisthorpe, 1935). Basidiomy- A. maculosum (N): Basidiomycetes: Fomitopsis cetes: Polystictus versicolor, Coriolus hirsutus, pinicola, Phellinus gilvus (Newton, 1984). Trametes gibbosa, Pholiota mutabilis, ``poly- A. mandibulare (P): Basidiomycetes: Polyporus pore'' (Donisthorpe, 1939; Palm, 1951; Reh- sp. (Luigioni, 1920); ``fungus'', ``fungus on as- fous, 1955); ``fungus'', ``mycelium under pen'' (Jansson, 1924; Meinert, 1893; Palm, bark'' (Champion, 1907; Fowler, 1889; Gus- 1951, 1959; Roubal, 1927; Rueschkamp, 1928; mann, 1914; Horion, 1949; Joy, 1932; Koch, Schiùdte, 1862; Wiepken, 1886; Woerndle, 1961; Palm, 1951, 1959; SjoÈberg, 1928; West, 1950). 1940; Woerndle, 1950; Woodroffe, 1968). A. marginatum (P): Myxomycetes: (Jansson, A. contiguum (N): Myxomycetes: Fuligo septica 1946); ``fungus'', ``fungusy bark'' (3) (Jans- (Russell, 1979). son, 1946; Polentz, 1938; Rueschkamp, 1928). A. convexum (P): ``fungus'' (Cooter, 1978). A. mollinum (N): Myxomycetes: Leocarpus fra- A. dentatum (P): Myxomycetes: Fuligo septica gilis (Lawrence and Newton, 1980); Cribraria (Rehfous, 1955); ``fungus'' (2) (Fleischer, purpurea; Diderma ¯oriforme; Stemonitis axi- 1910; Uhmann, 1925). fera. A. depressum (N): Myxomycetes: Stemonitis fus- A. nigrinum (P): Myxomycetes (2): Reticularia ca; Badhamia. lycoperdon (Dajoz, 1966; Palm, 1951). Basid- A. difforme (N): Myxomycetes: Stemonitis sp.; iomycetes: Polyporaceae (Palm, 1951); ``fun- Arcyria pomiformis, undet. (Newton, 1984). gus'', ``mycelium under bark'' (8) (Fowler, A. discoideum (P): Myxomycetes: Fuligo septica, 1889; Jansson, 1918b; Palm, 1951, 1959; Rye undet. (Lundberg, 1960; Palm, 1951). Basid- and Sharp, 1865; Sharp, 1871; Thomson, 1862; iomycetes: Polyporus pinicola, Armillaria mel- Wiepken, 1886). lea (Lundberg, 1960; Palm, 1951); ``fungus'' A. nigripenne (P): Basidiomycetes (5): Phellinus (10) (Ammann and Knabl, 1922; Gerhardt, igniarius (1), Hansenia abietina (2), Polyporus 1910; Heiss, 1971; Ihssen, 1939; Palm, 1959; squamosus (1), Polyporaceae (Hincks and Reitter, 1910; Roubal, 1926; Rueschkamp, Shaw, 1951; Jansson and SjoÈberg, 1932; Nuss, 1928; von Wanka, 1915; Woerndle, 1950). 1975; Platonoff, 1942; Saalas, 1917). Asco- A. estriatum (N): Myxomycetes: Fuligo septica mycetes: Daldinia concentrica (1) (Hingley, (Lawrence and Newton, 1980). 1971); ``fungus'', ``mycelia under bark'', ``fun- A. exiguum (N): Myxomycetes: Comatrichia ni- gusy bark'' (14) (Bagnall, 1905; Barner, 1922; gra, Reticularia splendens, , Donisthorpe, 1939; Ihssen, 1939; Lentz, 1879; Tubifera ferruginosa, plasmodium (Lawrence Palm, 1951, 1959; Palmen, 1946; Poppius, and Newton, 1980; CUIC, unpubl.); ``fungus'' 1905; Saalas, 1917; Sick, 1930; 1939; Singer, (2) (Cooper, 1935; Hamilton, 1895); ``shelf fun- 1955; West, 1940). gus''. A. oniscoides (N): Myxomycetes: Arcyria denu- A. fawcettae n. sp. (Part II) (N): Myxomycetes: data, Craterium minutum, Diderma ¯oriforme, Badhamia; Fuligo; Hemitrichia; Leocarpus; Fuligo septica, Stemonitis fusca, plasmodium Physarum; Tubifera; ``fungus''. (2), undet. (Stephenson et al., 1994; CUIC, un- A. globosum (P): ``fungus'' (Tomlin, 1915). publ.). Basidiomycetes: Fomes applanatus, A. haemorrhoum (P): ``fungus'' (2) (Jansson, Hydnum septentrionale (Park, 1931; Park et al., 1946; TheÂrond, 1975). 1931); ``fungus'' (2) (Blatchley, 1910; Hamil- A. kashmirense (O): Myxomycetes: Arcyria ci- ton, 1895); Ceratiomyxa fruticulosa, ``yellow nerea, A. denudata, Trichia favoginea, Tubifera '', and ``clear myxomycete''. 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 15

A. pallidum (P): ``fungus'', ``fungusy bark'', Lindroth and Palm; 1934); ``fungus'' (8) (Al- ``mycelium under bark'' (3) (Palm, 1947, len, 1954; Jansson, 1924; Jansson and SjoÈberg, 1951). 1932; Johnson, 1962; Lindberg, 1933, 1937; A. piceum (P): Basidiomycetes: Boletus verie- SjoÈberg, 1928; West, 1940). gatus (2) (Benick, 1952; Gusmann, 1925); A. temporale (N): Myxomycetes: Fuligo septica ``fungus'' (3) (Hansen, 1922b; TheÂrond, 1975; (7) (S.L. Stephenson, unpubl. record). West, 1940). A. tribulograndum, n.sp. (part II) (N): ``fungusy A. plagiatum (P): ``fungus'' (4) (Horion, 1949; wood chips''. Palm, 1959; Uhmann, 1925; von Wanka, 1915). A. tumidiventre, n.sp. (part II) (N): ``gilled mush- A. pulchrum (N): Myxomycetes: Arcyria denu- room''. data, A. incarnata, A. nutans, A. versicolor, A. varians (P): Basidiomycetes: Anisoporus odo- Comatricha suksdor®i, Fuligo septica, Lyco- rus, Polyporus fomentarius, P. squamosus, Po- gala epidendrum, , S. ¯avo- lyporus sp., Pholiota mutabilis, Trametes gib- genita, S. sp. (2), Trichia decipiens, Tubifera bosa, Tricholoma terreum (Jansson, 1918a; ferruginosa, Tubifera sp., undet. (Lawrence and Nuss, 1975; Rehfous, 1955; Rueschkamp, Newton, 1980; Russell, 1979; Stephenson et al., 1928); ``fungus'', ``fungus on aspen'', ``fun- 1994) Basidiomycetes: Coriolus versicolor, gusy branches'', ``moldy leaves'' (10) (Ger- Lenzites betulina (Newton, 1984); fungus (Fall, hardt, 1890, 1910; Jansson, 1918b; Morley, 1901). 1899; Palm, 1951, 1953, 1959; Roeben, 1907; A. reitteri (P): Myxomycetes: Reticularia lyco- Rueschkamp, 1928). perdon (Donisthorpe, 1935); ``fungus'' (Don- Agathidium spp.: Myxomycetes: undet. plasmo- isthorpe, 1929). dium, sclerotium (Stephenson et al., 1994); A. rhinoceros (P): Myxomycetes: Reticularia ly- Amaurochaete ferruginea, Arcyria denudata, A. coperdon, Symphytocarpus ¯accidus (Donis- nutans, A. pomiformis, A. stipata, Badhamia thorpe, 1935; Ing, 1967). Basidiomycetes: Fo- sp., Ceratiomyxa fruticulosa, Comatricha alpi- mes sp. (Angus, 1965); ``fungus'' (2) (Har- na, C. nigra, C. typhoides, Cribraria intricata wood, 1925; Roubal, 1909). (2, 1?), C. splendens (?), C. purpurea, C. rufa, A. rotundatum (P): Myxomycetes: Arcyria den- Cribraria sp. (2), Dictydium cancellatum, Di- udata, Lycogala epidendrum, undet. (Ing, 1967; derma ¯oriforme, D. niveum, Fuligo interme- Palm, 1951; Russell, 1979). Basidiomycetes: dia, F. septica (2), Hemitrichia clavata, Leo- Fomitopsis ungulata (2), Hansenia abietina, carpus fragilis, Lindbladia tubulina, Lycogala Polyporus fomentarius, Polyporus sp. (Jansson epidendrum, Physarum viride, Reticularia and Palm, 1936; Platonoff, 1942; Saalas, 1917; splendens?, Stemonitis axifera, S. ¯avogenita, Saalas and Schaufuss, 1923); fungus (14) (Ben- nett, 1893, 1899; Cooter, 1978; Fowler, 1889; S. hyperopta, Stemonitis sp., Tubifera ferrugi- Harwood, 1925; Joy, 1904; Koltze, 1901; Palm, nosa (Lawrence and Newton, 1980; Newton 1959; Roubal, 1926; Rye and Sharp, 1865; Saa- and Stephenson, 1990; Stephenson et al., 1994; las, 1917; Strand and Hanssen, 1932; Walker, Wheeler, 1984a). Basidiomycetes: Hansenia 1874, 1875). abietina (Saalas, 1917); ``fungus'' (5) (Anon- A. rubellum (N): Myxomycetes: Tubifera ferru- ymous, 1889; Lundberg, 1984; Rottenberg, ginosa, ``mushrooms''. 1864; Roubal, 1924; Weber, 1916). A. seminulum (P): Myxomycetes: Fuligo septica From these data and those for Anisotoma (2), Reticularia lycoperdon, Trichia cinnabari- na (3) (Donisthorpe, 1935, Perris, 1851; Reh- (Blackwell, 1984; Lawrence and Newton, fous, 1955; Weber, 1925; West, 1940) Basid- 1980; Russell, 1979; Wheeler, 1979b, 1984a, iomycetes: Auricularia auricula; Coriolus ver- 1984c) it is reasonable to conclude with sicolor, Lenzites betulina, Polyporus caudici- Newton (1984) that whereas Anisotoma spe- nus, P. squamosus, Trametes gibbosa, Collybia cies appear to be obligate slime-mold feed- platyphylla, Hypholoma fasciculare (Benick, ers, evidence for Agathidium is less conclu- 1952; Eisfelder, 1963; Rehfous, 1955; Saalas, sive. Of 27 species reported from Myxomy- 1917); ``fungus'', ``mycelium under bark'' (13) cetes, about 56% are reported also from Ba- (Ammann and Knabl, 1922; Derksen, 1941; sidiomycetes. And of 21 species noted from Donisthorpe, 1939; Koltze, 1901; Palm, 1951, 1959; Polentz, 1938; Roussin, 1947; Ruesch- Basidiomycetes, 38% are known also from kamp, 1928; Saalas, 1917; Saalas and Schau- Myxomycetes. Thirty-two species are report- fuss, 1923; von Wanka, 1908; Williams, 1924). ed from Myxomycetes, 23 from Basidiomy- A. sphaerulum (P): Basidiomycetes: Polyporus cetes, and 14 are noted from both. Eighteen fomentarius, Polyporus sp. (Brundin, 1934; species of Agathidium are recorded from 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Myxomycetes exclusively, whereas nine spe- search image) for the hosts. Sporocarps of cies are recorded only from Basidiomycetes. Myxomycetes are so diverse in size, shape, Some species have been reported so many and color that learning to recognize all or times from diverse Basidiomycetes that an most of them as slime-molds is in itself an association with these higher fungi is strong- accomplishment (®gs. 1, 2). Fruiting bodies ly suggested. This is reinforced by reports of vary from minute, stalked forms less than 1 spores of Polyporus squamosus in the gut of mm in height to enormous cushionlike fru- Agathidium seminulum (HlisnikovskyÂ, itions of Fuligo or dense patches of hairlike 1964). Further, whereas species of diverse Stemonitis that may be more than a foot in species groups (including three subgenera as diameter. Similarly, plasmodia vary from mi- generally recognized: Agathidium s.s., Cy- croscopic, transparent, amoeboid forms to phoceble, and Neoceble) have been associ- the world's largest polynucleated cells. ated with Myxomycetes, species of these Agathidium aristerium, for example, was ®rst same groups have been repeatedly associated discovered breeding on a plasmodium of with Basidiomycetes also. For at least some Physarum that was estimated to be larger of these species, however, there may be an than 4 feet in diameter! Some slime-molds alternative explanation. are easily recognized, even in the plasmodial Agathidium aristerium was observed feed- stage. Most are positively identi®ed only ing on plasmodia of the myxomycete Phy- with more careful, microscopic study (Martin sarum polycephalum growing on the surface and Alexopoulos, 1969). of oyster shell fungi (Pleurotus ostreatus: Once located, slime-molds may be re- Tricholomataceae) on a beech tree in New moved with a piece of their substrate for York state (Wheeler, 1987). This epimycetic closer examination. It is useful to hold a growth of Physarum polycephalum on Pleu- white sheet or a white box under the fruiting rotus is a phenomenon known well to my- body while it is cut away from its substrate cologists, and the plasmodia of this and other with a sharp knife blade. Sporocarps may be slime-molds grow upon the surfaces of de- returned to the laboratory, where they can be clining sporocarps of higher fungi. For at spread upon a light-colored background un- least some basidiomycete host reports for der the heat of a desk lamp for sorting. Bee- Agathidium it seems possible that plasmodia tles covered by host spores are amazingly could have been present but undetected by dif®cult to see. However, once heated, bee- collectors. In the future, special care should tles will become active and easily spotted be taken to examine host Basidiomycetes and sorted from the mass of host spores. carefully to ascertain if and when this may Vouchers of sporocarps may be kept by glu- be the case. ing them to the inner surface of the lid of a The occurrence of apparently both slime- fungus specimen box and drying them com- mold and higher fungus feeding in Agathi- pletely. dium, combined with the occurrence of Because plasmodia are easily damaged or slime-mold feeding only in the related genus desiccated, it is sometimes easier to simply Anisotoma, reinforces the hypothesis that examine them closely in the ®eld. Both adult slime-mold associations may have been an- and larval beetles may be seen upon plas- cestral for the two clades combined and, by modia since they lack the cover of spores extension, for Agathidiini as a whole (New- afforded by mature hosts. Some plasmodia, ton, 1984; Wheeler, 1984a, 1984c). particularly certain pigmented ones (such as the common yellow Physarum and Fuligo COLLECTING AGATHIDIUM species), can be induced to mature by ex- posing them to intense light as, for example, Specimens of Agathidium species may be setting them in a Petri dish in direct sunlight collected either by hand directly from host on a window sill. They can be collected and slime-molds or indirectly by means of certain kept in the laboratory on either agar plates mechanical contrivances or traps. The most or damp ®lter paper onto which are placed dif®cult part about searching for Agathidium one or two ¯akes of old-fashioned oatmeal. on Myxomycetes is developing an eye (i.e., Upon fruition, such forms may be positively 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 17 identi®ed. Mycologists sometimes have clues slime-mold associations of A. oniscoides to plasmodial identi®cation that can be very makes conjectures about the functional sig- helpful to the entomologist. Mustard yellow ni®cance of its morphology possible. For plasmodia that stain brick red upon ``bruis- theoretical reasons, one might prefer a spe- ing'' (that is, touching them with one's ®n- cies less removed from the ground plan for ger) are usually members of the genus Fuli- the genus Agathidium. For example, differ- go, though we do not know that similar ences between species of the A. sexstriatum bruising does not exist in other genera. Some species group and related taxa, notably Ani- plasmodia, including Fuligo, have a charac- sotoma species, are few and as such one of teristic odor that can be quite strong when these species might have made a good the plasmodium is expansive. Occasionally a choice. However, the morphology of Aniso- plasmodium can be smelled before locating toma has already been discussed elsewhere it visually. It has a sweet, earthy smell. Non- (Wheeler, 1979b), and A. oniscoides is more pigmented plasmodia are more dif®cult to typical of the highly contractile forms gen- see, such as the gray, jellylike Stemonitis erally thought of as representative Agathi- plasmodium that can be quite substantial in dium. size. The morphology of adult and pupal stages Flighted species may be collected using of A. oniscoides is given below. The eggs large-area ¯ight-intercept traps of the kind that have been seen in the course of female described by Peck and Davies (1980). Both dissections have been oval, shiny, whitish ¯ighted and ¯ightless species may be col- colored, without striking structural details at lected by sifting leaf litter, decaying wood, low magni®cations, and notable only in their and similar substrates likely to include slime- large size relative to the abdominal cavity. molds and placing the siftate in a tullgren or Agathidium larvae are active, eruciform, and Berlese funnel. This is particularly effective setose and are rather broad and dorsoven- for collecting ¯ightless species that are com- trally compressed in shape (Wheeler, 1990a). mon in the Appalachians and Ozarks. In fact, There are three postembryonic larval instars during wet times of the spring and fall that are rich in external characters, particu- months in the Appalachians, sifting samples larly with regard to chaetotaxy. The compar- of mixed rhododendron-hardwood litter ative morphology and ontogeny of larval se- along mountain streams predictably yields maphoronts of three North American species one or more such species. In such circum- of Agathidium were described elsewhere in stances leiodids often are exceeded in num- detail (Wheeler, 1990a), including all post- ber of individuals only by carabids. embryonic instars of A. oniscoides. These data were used in a critical evaluation of Nel- MORPHOLOGY son's (1978) biogenetic rule published sepa- rately (Wheeler, 1990b). Additional descrip- A detailed account of the morphology of tions of larvae are given by Kilian (1998). Agathidium oniscoides is presented below. This is done to indicate through accurate il- ADULT lustrations the structures referred to in formal Figures 3±37 descriptions, and to establish a basis for com- parative morphological studies within and BODY FORM (®gs. 3±5): Very broadly outside of Agathidium. Agathidium oniscoi- round, combined elytra about as wide as des was selected as a representative species long; very convex, hemispherical; pronotum for biological and practical reasons. It is one laterally expanded; sterna, particularly pros- of the most common and abundant species of ternum and metasternum, short; capable of northeastern North America, making many contracting body into compact, ball shape specimens available for study. Its compara- concealing venter and appendages; length tively large size made illustration of certain about 4.0±6.0 mm, extended; moderately structures easier. Because its eggs, larvae, heavily sclerotized; dorsum smooth, shiny, and pupae are known, adult structures may with at most microscopic setose punctures, be seen in ontogenic context. Knowledge of but under low to moderate magni®cation 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 3±5. Agathidium oniscoides Palisot de Beauvois, habitus: 3, dorsal; 4, ventral; 5, lateral. 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 19

(e.g., 10±25ϫ) nearly devoid of punctation MFD Metasternal fovea dimensionÐ or setation; color at maturity dark reddish- greatest width. brown to black. Antennomeres Length of antennomeresÐmea- I±XI sured along dorsal midline. MORPHOMETRICS (®gs. 6±8): If few speci- mens were available, then all of them were CRANIUM (®gs. 9±12): Prognathous, very measured. If more material was available, broadly quadrate (cf. ®gs. 11, 12); dorsoven- then four or ®ve of the smallest and largest trally compressed (®gs. 9, 10), height less individuals were measured to give an indi- than one-third cranial width; wider than long cation of the size range for the species. Mea- (OHW/MDL ϭ 1.4); vertex large, ¯at; with surements were made on a Wild M3C dis- distinct supraocular ridge from edge of clyp- secting microscope at 64ϫ magni®cation us- eus nearly to posterior margin of head; pos- ing a Lasico digital micrometer. Morphomet- tocular temporum short, not prominent in rics referred to in this study include the dorsal view; eyes anterolaterally positioned, rounded in dorsal view, more angulate in following primary measures, some of which ventral view, ®nely faceted; genae broad, are presented in the text in the form of rel- each nearly as wide as clypeus; clypeus ative ratios: quadrate; frontoclypeal suture faint medially, OHW Ocular head widthÐmeasured at incompletely separating clypeus from frons; midpoint of longitudinal axis of gular sutures convergent posteriorly and eye. slightly divergent at apices; ventral antennal PHW Postocular head widthÐmea- grooves present between eyes and gular su- sured at posterior margin of eye. tures, convergent posteriorly, shallowly im- MDL Median head lengthÐmeasured pressed; occipital foramen small, about one- from base of labrum to posterior margin of cranium. third width of cranium (®g. 9); tentorium PNW Pronotal widthÐmeasured at with distinct anterior and posterior bridges midpoint of longitudinal axis, (®g. 9); antenna inserted anterior to eye, be- measured in practice in ventral neath supraocular ridge (®g. 10). view to avoid problems of cur- Members of other species of Agathidium vature in dorsal orientation. (especially the A. concinnum and A. pul- PNH Pronotal heightÐdistance along chrum groups) often have a moderately dis- imaginary axis from midpoints of tinct crease extending transversely across the dorsum and lateral margin, mea- medial portion of the head. With the head sured in lateral view. fully extended forward, this crease receives PNL Pronotal lengthÐmeasured along the anterior, curved margin of the pronotum. midline. Agathidium oniscoides lacks this crease. ELW Elytral widthÐcombined elytral widths at longitudinal midpoint. ANTENNA (®gs. 13, 14): Short, with abrupt SEL Postscutellar elytral lengthÐ 3-segmented club; antennomere I large, nar- length of elytra along suture from row and geniculate at base, wider distally, posterior margin of scutellum to with sparse setae; antennomere II short, apex. wide; antennomere III elongate, much longer ELH Elytral heightÐmeasured along than II, slightly expanded at apex, with imaginary axis between halfway sparse setae, including subapical circle of points of elytral dorsum and lat- larger setae; antennomeres IV and V short, eral margin in lateral view. wider in apical half, with sparse ®ne setae MSL Mesosternal lengthÐmeasured at and subapical circle of larger ones; anten- midline. nomeres VII and VIII short and compara- MTL Metasternal lengthÐmeasured at tively more transverse, with comparable se- midline. MTW Metasternal widthÐmeasured tation; antennomeres IX and X subquadrate, between posterolateral corners. similarly proportioned, with dense ®ne seta- TBL Total body lengthÐthe sum of tion, and subapical ``ring'' of larger setae; head length (MDL) ϩ pronotum antennomere XI elongate, narrowed apically, length (PNL) ϩ postscutellar ely- surface similar to antennomeres IX and X. tral length (SEL). Antennomeres IX and X with two Hamann's 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 6±8. Schematic drawings of Agathidium oniscoides Palisot de Beauvois showing various body measurements: 6, dorsal; 7, ventral; 8, lateral. See text for explanation of abbreviations. 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 21

Figs. 9±12. Agathidium oniscoides Palisot de Beauvois, cranium: 9, posterior; 10, lateral; 11, ventral; 12, dorsal. organs present, opening into periarticular asperities basally; dense erect setae at apex, gutter (®g. 14). obscuring apical row of spines and apical MAXILLA (®g. 15): Cardo small, quadroid; tooth; galea broad, apex rounded, bearing stipes short, broad, with single lateral seta; dense long ®ne setae, basal two-thirds cov- lacinia broad, apex truncate; dense minute ered by dense minute asperities; palpifer 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 13, 14. Agathidium oniscoides Palisot de Beauvois, antennae: 13, antenna; 14, apical anten- nomeres. 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 23

Figs. 15±19. Agathidium oniscoides Palisot de Beauvois, mouthparts: 15, maxilla; 16, mandible; 17, labrum, dorsal; 18, labrum, ventral; 19, labium, ventral. small; palpus with three palpomeres; pal- penicillus of long, ®ne setae; subbasal pros- pomere I narrow at base, campanulate, with theca covered in very dense, ®ne, short setae; few setae; palpomere II broader, widened mesobasal mola with poorly de®ned parallel apically, covered by long setae; palpomere transverse and slightly serrate lines. III large, narrowed apically, covered by long LABRUM AND EPIPHARYNX (®gs. 17, 18): setae, apex with numerous minute sensilla. Labrum transverse, with median emargina- MANDIBLE (®g. 16): Broad, short, robust; tion fringed with dense, minute setae; dorsal with single apical dens; subapical orthogonal surface with 3 pairs of short, mesal, subapi- 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 cal setae, about 7 pairs of large lateral, dorsal tibial insertion when ¯exed, metafemur of setae, about 2 pairs of minute setae near mid- male sometimes with subapical tooth (see be- line of dorsal surface, and a pair of sensilla low); tibia narrow, with anterior carina, rows near base (®g. 17). Adoral surface with short, of setae; apex fringed with stout crown of broad paired setae near apex, paired median spines; tibial spurs small on prothoracic leg, ®elds of sensilla, and paired median fringe of larger on mesothoracic leg, very large on microtrichiae. Epipharynx membranous, with metathoracic leg. Tarsomeres sexually di- very dense, ®ne microtrichiae (®g. 18). morphic: ( 5±5±4 / & 5±4±4. Pro- and me- LABIUM (®g. 19): Mentum broad at base, sobasotarsomeres (especially I ϩ II) expand- narrowed distally, with paired sensilla near ed in male, bearing dense ventral spatulate middle, numerous asperities, and moderately setae. dense setae; ligula broad apically, very shal- ABDOMEN (®g. 35): Short, broad; sternite lowly emarginate, with median sclerotized III (®rst visible sternite) longer, with deep, plate bearing setae and dense apical setae broad coxal depressions, pointed intercoxal merging with hypopharyngeal setae. process; IV±VII gradually narrowed; VIII PRONOTUM (®gs. 3, 20): Wide, very con- small. vex; prohypomeron very broad, convex, re- ceiving and overlapping humeral angles of MALE SEXUAL DIMORPHISM elytra when body contracts; procoxa trans- verse; procoxal cavity externally closed pos- Male and female Agathidium differ in a teriorly, small, and round; prosternum short; number of secondary sexual characters. cervical opening broad; prothoracic spiracle Males have enlarged basal tarsomeres on the elliptical. front and middle pairs of legs and dense, MESOSTERNUM (®gs. 4, 21): Short; divided spatulate ventral setae on the expanded tar- by lateral line into anterior and posterior someres, a metasternal fovea, a metafemoral parts; with median carina; mesocoxal cavity tooth, and (in some) mandibular tusks. Each large, transverse, closed externally. is described for male A. oniscoides and dis- METASTERNUM (®gs. 4, 21, 24): With cussed in general below. oblique line (``linee ®morali'' of Angelini PRO- AND MESOBASOTARSOMERES: Three and de Marzo, 1980); metepisternum narrow; basal tarsomeres of front (®g. 30) and middle intercoxal process narrow (®g. 21). With me- leg moderately laterally expanded, with dian fovea in male (see below). Metendos- dense ventral pad of elongate, spatulate setae. ternite compact, furcal arms broad, separated METASTERNAL FOVEA (®g. 21): Small, by distinct median lamina (®g. 24). round, located medially, bearing tuft of long, ELYTRON (®gs. 3, 22): Combined elytra golden setae. about as broad as long; surface appears im- The metasternal fovea is present in almost punctate at low magni®cation, with micro- all male Agathidium, although its placement, scopic setose punctures, barely visible at size, and shape vary considerably. It is al- higher magni®cations (e.g., 60±80ϫ); hu- ways found at the midline. It is usually lo- meral angle obtuse, poorly de®ned; epipleu- cated near the center of the metasternum, but ron broad, narrowed posteriorly; subcuticular is sometimes placed near its anterior or pos- serial vestiges visible posteriorly in cleared terior margin. The fovea is single or, more specimens under transmitted illumination. rarely, paired. It can be minute and puncti- HIND WING (®g. 23): With reduced vena- form, larger and circular, or very large and tion; costa and subcosta fused (C/Sc), con- transversely arcuate in shape. The fovea usu- ®ned to basal 0.40 length anterior margin of ally bears a tuft of long, ®ne, golden setae wing; median vein (M) well de®ned in apical that each stand individually erect or are, fre- half; sclerotized ®eld present between medi- quently, pasted together as a brush. an line and costa-subcosta; cubitus (Cu) well Similar foveae are found in other male de®ned in apical half of wing; two well-de- Agathidiini, including Anisotoma species veloped anal veins present (A). (Wheeler, 1979b). Foveae are also sometimes LEGS (®gs. 25±31): Elongate, straight; fe- found on the labium in this taxon. Faustini mur broad, partially excavate near apex at (1980) and Faustini et al. (1981) examined 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 25

Figs. 20, 21. Agathidium oniscoides Palisot de Beauvois: 20, prothorax, ventral; 21, pterothorax, ventral. 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 22±24. Agathidium oniscoides Palisot de Beauvois: 22, elytron, dorsal; 23, Hindwing hingw- ing; 24, metepisternite, dorsal. similar structures in a range of Coleoptera MANDIBULAR TUSK: Absent, left mandible and concluded that they function as external not modi®ed. surface areas for the dissemination of secre- The left mandibles of a number of other tions. male Agathidium are modi®ed into a horn or METAFEMORAL TOOTH: The posterior mar- tusk (e.g., ®gs. 89, 90). The development of gin of the male metafemur protrudes poste- the tusk varies, both between and within spe- riorly as a broad, blunt tooth. cies, from a small bump to a large horn. The 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 27

Figs. 25±29. Agathidium oniscoides Palisot de Beauvois, legs: 25, proleg, male; 26, mesoleg, male; 27, metaleg, male; 28, proleg, female; 29, tarsal claw detail, male. 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

MALE GENITALIA Figures 32±34

AEDEAGUS: The aedeagus in members of Agathidium is trilobate and characteristic of ``primitive'' staphylinoids (Crowson, 1955). The basic structure consists of a tubular and curved median lobe, paired lateral lobes which are usually nearly as long as the me- dian lobe and joined basally by the phallo- base (formed by a ring like structure encir- cling the base of the median lobe), and a sub- apical opening on the ventral face of the me- dian lobe (the distal ori®ce). In Agathidium the distal ori®ce is usually partially covered by a sclerotized ¯ap (the operculum or ``ven- tral piece''; Wheeler, 1979b). This structure is highly variable in shape between species and is often an important source of taxonom- ic information. It is usually clearly articula- ble basally, but in some cases is apparently not. For the purposes of this revision, any structure extending over the distal ori®ce from its base is regarded as an operculum, though not all these structures may be nec- essarily homologous. Internal to the median lobe is a membranous, saclike endophallus which bears a variety of armature in the form Figs. 30, 31. Agathidium oniscoides Palisot of spines, hairs, and sclerotized patches in de Beauvois, protarsi: 30, male protarsus detail; addition to apical or subapical membranous 31, female protarsus detail. lobes in the form of elongate digits or lan- ceolate structures. The median lobe, at least near the apex, and the lateral lobes exhibit a variety of patterns in the distribution of horn is sometimes posteriorly curved and ``pores'' and ``ducts'' (see below). bears a tuft of apical setae. At least two ap- MEDIAN LOBE (®gs. 32±34): In A. oniscoi- parently distinct kinds of horns occur that des the median lobe is elongate, narrow, ven- may not be homologous with one another. In trally curved through its midlength (most one form the apex of the mandible is pro- sharply near the basal piece), and recurved longed and dorsally recurved into a hornlike in the apical one-®fth. The apical portion structure (e.g., A. aristerium; Wheeler, 1987). (one-®fth) is dorsoventrally compressed rel- In others, such as A. pulchrum, a tusk arises ative to the basal portions of the median lobe. from the dorsal surface of the mandible, pos- The basal piece is completely fused to the teriorly removed from the apex, which re- median lobe dorsally. mains unmodi®ed (see Frontispiece). It is ``PORES AND DUCTS'' OF MEDIAN LOBE (®g. conceivable that one represents a state mod- 32): The terms ``pore'' and ``duct'' are used i®ed from the other. From a strictly morpho- advisedly. ``Pore'' has been applied broadly logical perspective, however, it seems more in the larval literature for a variety of struc- reasonable to suppose that these represent in- tures, including campaniform sensilla (Ashe dependent modi®cations of the left, male and Watrous, 1984). In Agathidium genitalia mandible, but perhaps with some shared de- the exact nature of the pores has not been velopmental genetic coding considering the determined. However, based on compound unusual asymmetry of the structure. microscopy and a limited amount of scan- 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 29

Figs. 32±34. Agathidium oniscoides Palisot de Beauvois, aedeagus: 32, median lobe apex, ventral; 33, median lobe apex, lateral, endophallus partially everted; 34, lateral. ning electron microscopy, these do not seem cells, feeding through the visible ``ducts'' to to always have any obvious surface struc- a median canal. These may be glandular, but tures associated with them. ``Pore'', then, is we hesitate to strongly hypothesize this until purely a descriptive word, since this is what more detailed ®ne structure work has been they appear to be under compound micros- done. copy. They may be reservoirs of secretary In A. oniscoides the ducts of the median 30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 lobe radiate from a central region, lining the Agathidiini (ϭ Anisotomini auctorum ), male sides and rounding the apex of the apical and female tarsal formulae differ. Whereas one-®fth of the median lobe (®g. 32). the numbers of tarsomeres vary among LATERAL LOBES (®g. 32): The lateral lobes males, females consistently have a reduced of A. oniscoides are slightly widened at the number in comparison to the male of the middle and are narrower at their apices. same species (e.g ( 5±5±4 / & 5±4±4 or ( Pores are irregularly distributed over the sur- 5±4±4 / & 4±4±4). face, but they are more dense apically and absent basally. Two subequal setae are pre- FEMALE GENITALIA sent subapically. Figures 36, 37 OPERCULUM (®gs. 32±34): The operculum is narrowed gradually. Its apex is abruptly Female genitalia (®g. 37) short, membra- wider, forming an apical knob. Pores are pre- nous; proctiger broadly rounded; coxite nar- sent at the apex only. There is some variation row, elongate, cylindrical with moderate in the operculum within the species, partic- number of small pores, few small subapical ularly in the degree of pronouncement of the setae; stylus small, narrow, elongate with sin- apical knob. gle long apical seta, pair of short subapical ENDOPHALLUS (®g. 33): The endophallus in setae. A. oniscoides, like in other Agathidium,is SPERMATHECA (®g. 36): Spermatheca elon- typically inverted, and details of its structure gate, curved; with capsule large, bulbous at are dif®cult to discern. When everted, the en- base; neck narrow, curved; apex rounded, dophallus can be seen to be complex (®g. blunt; walls thick and lumen paralleling cap- 33). In A. oniscoides there are few scleroti- sule shape; very long, narrow spermathecal zations except for a number of long tentacle- duct inserted into bulb; membranous, amor- like subdivisions of the sac. There are two phous gland inserted near base of spermathe- bifurcate structures that are wrinkled in ap- cal duct. pearance and several smaller ones, including two smooth, very sharply pointed processes. PUPA Details of endophallic structure have been Figures 38±40 dif®cult to assess in the genus. No reliable Pupa broad, rounded, and whitish colored. technique was found for its eversion. Some- Head inferior to laterally expanded prono- times gentle heating of the ¯uid, particularly tum. Terminal dorsal abdominal segment of glycerin, will lead to eversion, but this is with paired, pointed, posteriorly-directed not entirely predictable and certainly cannot spines. be counted on for species where large num- bers of specimens are not available for dis- TAXONOMY section. When such techniques or large enough numbers of specimens to support tri- An apparent synapomorphy of the tribe al-and-error approaches do exist, many ad- Agathidiini (ϭ Anisotomini) is the sexually ditional characters will certainly be found. dimorphic tarsal formulae. The number of We predict that the number of endophallic tarsomeres is greater in males than in females features may well surpass the number of ex- with the infrequent exception of when 4±4± ternal genitalic ones when they are examined 4 tarsi (or, even more rarely, 4±4±3) occur in detail. The ®ngerlike subdivisions of the in both sexes of certain species. Whereas sac in A. oniscoides, for example, are present male tarsomere numbers are variable, they in closely related species and are potentially are usually heteromerous and greater than fe- synapomorphic. male tarsomere numbers whether heterom- erous or not, for example, ( 5±5±4 / & 5± FEMALE SEXUAL DIMORPHISM 4±4 and ( 5±4±4 / & 4±4±4. Agathidiines are small, about 1±6 mm in length, round, The only dimorphism found in females in- and convex. Most are glabrous or very nearly volves a reduction in the number of tarso- so, and with rare exceptions (e.g., Anisotoma meres. As in other members of the tribe basalis, Agathidium pulchrum, A. maculos- 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 31

Figs. 35±37. Agathidium oniscoides Palisot de Beauvois, female: 35, abdomen, ventral; 36, sper- matheca; 37, genitalia. um) are uniformly reddish-brown to black. diini belong to the Leiodinae, one of six sub- Serial punctation is symplesiomorphically families in Leiodidae (Newton, 1998) (Lawr- scattered through the tribe and is reduced or ence and Newton, 1995). Like most leiodids, lost in many species of Agathidium. Agathi- many agathidiines have the eighth antennom- 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 38±40. Agathidium oniscoides Palisot de Beauvois, pupa: 38, dorsal; 39, ventral; 40, lateral. ere narrower than either the seventh or ninth terpreted as part of a 5-segmented antennal (which are generally more equal in size), cre- club, the postocular temporum and supraocu- ating a characteristically interrupted 5-seg- lar carinae are prominent and conspicuous, a mented antennal club. Most Agathidium, character combination not found in related however, have the seventh antennomere re- taxa like Anisotoma. duced in size also, and thus they typically Species of Agathidium were ®rst recog- possess an abrupt 3-segmented club. In some nized by Linnaeus (1758) and Fabricius species of Agathidium the seventh antennom- (1792). The genus was proposed by Panzer ere is slightly larger than the eighth, but both (1797) for Silpha seminulum Linnaeus (ϭ are still much narrower than the ninth. In the Agathidium globosum Panzer). The ®rst New few cases when Agathidium has the seventh World species of Agathidium described was antennomere large enough to possibly be in- A. oniscoides Palisot de Beauvois (1817), be- 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 33 ing one of the largest and most common spe- present, series often obscured by con- cies native to the United States. The ®rst fused punctation between; postocular comprehensive revision of the genus was temporum present; tarsi ( 5±5±4, 4±4±4 done by Horn (1880) in his classic treatment or 4±3±3/ & 5±4±4, 4±4±4 or 4±3±3 . . of the Silphidae. The second and most recent ...... Agathidium Panzer taxonomic revision was completed by Fall Cainosternum Notman (1934). Four described genera of Agathidiini are Cainosternum Notman, 1921: 147 (Type species: currently recognized in North and Central Cainosternum imbricatum Notman, 1921, by America and may be distinguished by the monotypy); Hatch, 1929; Wheeler, 1986; New- following diagnostic key. Keys to the world ton, 1998. genera are provided by Angelini and Peck DISCUSSION: The genus Cainosternum was (2000), along with a cladogram for genera of originally described from western New York the tribe, and by Peck et al. (2000) for Cen- state by Notman (1921) based upon a single tral and South American genera. In this last specimen of Cainosternum imbricatum Not- treatment the authors also included two pu- man. The senior author has made numerous tatively undescribed genera which we will be trips to the type locality in Chautauqua treating separately. A catalog and complete County in the vicinity of West®eld in search bibliographies and synonymies of world gen- of this unusual species but failed to collect it era are provided by Newton (1998), along from either slime-molds or sifted leaf litter. with comments on some of the numerous However, while collecting ¯ightless Agathi- problems with classi®cation of the group. dium species in the Appalachian Mountains of western North Carolina, C. imbricatum KEY TO GENERA OF AGATHIDIINI was rediscovered (Wheeler, 1986). No iden- OF NORTH AND CENTRAL AMERICA ti®able inclusions were observed in the gut of the North Carolina material, and habits of 1. Elytron with many (ca. 20) impressed rows of this elusive species remain unknown. Sexu- punctures, integument between rows with ally dimorphic tarsal formulae suggest that transverse striations; mesosternum keel-like, interrupted by large excision visible in lat- the genus belongs within Agathidiini. If this eral view; apex of lateral lobe membranous, is so, then the widespread and putatively an- setose; eastern U.S...... cestral associations of Agathidiini with ...... Cainosternum Notman Myxomycetes suggest possible slime-mold Ð Elytron with 9 or fewer rows of punctures, predation. Plasmodial feeding would be con- frequently with only irregularly distributed sistent with both the absence of microscopi- punctures or none; mesosternum with or cally recognizable material in the gut and ab- without carina, never in form of excised sence of specimens on exposed fruiting bod- keel; apex of lateral lobe not membranous ies. Although this inference is not fully jus- ...... 2 ti®ed by available data, such speculation 2(1). Antennal club distinctly 5-segmented, an- tennomere VIII narrower than VII; Hol- does provide one logical place to search for arctic, in New World from Alaska and Cainosternum in the future. Canada south to Guatemala ...... Cainosternum was excluded from Agathi- ...... Anisotoma Panzer diini and placed in the tribe Pseudoliodini by Ð Antennal club 3-segmented, if antennomere Newton (1998) and Peck (1998). While VII slightly wider than VIII, VII much pointing out the similarity of this taxon to narrower than IX and/or head with prom- members of Pseudoliodini, Newton (1998) inent postocular temporum ...... 3 also remarked that Cainosternum could rep- 3(2). Clypeus not excavate; elytron with distinct resent the basal member of the tribe Agath- punctured striae; postocular temporum idiini. This taxon does possess an unusual absent (in American species); tarsi ( 5± 5±4 / & 5±4±4 or 4±4±4 ...... character combination. Since there is no ...... Stetholiodes Fall comprehesive cladistic analysis of members Ð Clypeus excavate or not; elytron usually of the various tribes of Leiodinae and, there- without punctures or, if punctation pre- fore, no good hypotheses for the relative ple- sent, usually confused, if serial punctation siomorphy or apomorphy of characters used 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 to group these taxa, we have elected to retain Blatchley, 1910; Wheeler, 1981; Newton, 1982, this taxon within Agathidiini until such anal- 1998; Angelini and de Marzo, 1983, 1987b; yses can be presented. Perkovskii, 1990. Agathodes Portevin, 1926: 80 (Type species: Anisotoma Panzer Agathodes striatipenne Portevin, 1926, by mon- Figure 41 otypy) preoccupied, nec Agathodes GueÂneÂe, 1854, Lepidoptera. Anisotoma Panzer, 1797: 8 (Type species: Tritoma Agathidiodes Portevin, 1926: 169 (new name for glabra Fabricius, 1787, by subsequent desig- Agathodes Portevin, 1926); Newton, 1983 (syn- nation of Hatch, 1929b, but see Newton, 1998); onymized with Stetholiodes). Wheeler, 1979a; Angelini and de Marzo, 1988; Newton, 1998. DISCUSSION: The genus Stetholiodes was Pentatoma Schneider, 1792: 339 (Type species: described from northern Indiana by Fall Sphaeridium humerale Fabricius, 1792, by sub- (1910), based on material collected from sequent designation of Hatch, 1929b), preoc- moss (Blatchley, 1910). Wheeler (1981) re- cupied, nec Pentatoma Olivier, 1789, Hemip- described S. laticollis Fall and discussed this tera. single North American member of the genus. Eucyrta Portevin, 1927: 82 (Type species: Eucyr- Additional species are known from Asia and ta didymata Portevin, 1927, by subsequent des- eastern Europe (Angelini and de Marzo, ignation of Hatch, 1929b), preoccupied, nec Eu- cyrta Felder, 1874, Lepidoptera; Wheeler, 1987b). Monophyly of the genus has not 1979a (synonymized with Anisotoma). been ®rmly established. Habits remain un- known, though slime-mold associations in DISCUSSION: Species of Anisotoma are related genera suggest similar relationships among the most common, widespread, and in this taxon. abundant slime-mold-associated Coleoptera in North and Central America. In general Agathidium Panzer habitus, some species of Agathidium (e.g., Ag. sexstriatum and Ag. concinnum groups) Volvoxis Kugelann, 1794: 535 (Type species: Te- strongly resemble Anisotoma species (e.g., tratoma globosa Herbst, 1792 (ϭ A. seminulum (Linnaeus, 1758), by subsequent designation of ®g. 41), from which they may be distin- Newton, 1998), not preoccupied by Volvox Lin- guished by their distinct 3-segmented anten- naeus, 1758, Chlorophyta. NOMEN OBLITUM. nal club or continuation of the supraocular Agathidium Panzer, 1797: 13 (Type species: Te- carina posterior of the eye, setting off a dis- tratoma globosa Herbst, 1792 (ϭ Silpha semi- tinct posterior temporum. nulum LinneÂ, 1758, by monotypy); Illiger, Anisotoma species of North America were 1798; Palisot de Beauvois, 1817; Sturm, 1807; revised by Wheeler (1979b). Since that re- Stephens, 1829, 1839; Leach, 1832; Erichson, vision, a few species have been added from 1845; Redtenbacher, 1845, 1849, 1858, 1874; MeÂxico and Panama (Wheeler, 1980, 1983). Fairmaire and LaboulbeneÂ, 1854; JacquelõÂn du Based on the paucity of specimens collected Val, 1857; Thomson, 1859, 1862; LeConte, 1861; Brisout, 1872; Cox, 1874; Seidlitz, 1891; from MeÂxico and Central America to date Horn, 1880; LeConte and Horn, 1883; Fowler, and the high proportion of undescribed spe- 1889; Everts, 1899; Ganglbauer, 1899; Stierlin, cies found among them, it seems likely that 1900; Schaufuss in Calwer, 1916; Jacobson, the number of species of Anisotoma will be 1910; Blatchley, 1910; Kuhnt, 1913; Portevin, signi®cantly greater than that now known. 1914, 1926, 1927, 1929; Barthe, 1920±1922; Most slime-mold host records of Anisoto- Hansen, 1922a; Houlbert, 1922; Hatch, 1929a, ma are in association with mature sporocarps 1929b, 1957; Joy, 1932; Fall, 1934; Javorek, (Blackwell, 1984; Lawrence and Newton, 1947; Horion, 1949; HlisnikovskyÂ, 1964; Ar- 1980; Stephenson et al., 1994; Wheeler, nett, 1971; von Peez, 1971; Hendrichs, 1980; 1979b, 1984a), although plasmodial feeding Angelini and De Marzo, 1984, 1987a, 1987b, 1988, 1990, 1995; Angelini, 1984, 1988, 1990; has been observed (Newton, 1984; Russell, Nunberg, 1987; Wheeler, 1987, 1990a; Newton, 1979; Wheeler, 1980). 1998; Angelini and Peck, 2000; Peck et al., 2000. NOMEN PROTECTUM. Stetholiodes Fall Agathidium Illiger, 1798: 81, unnecessary replace- Stetholiodes Fall, 1910: 4 (Type species: Stetho- ment name for Volvoxis Kugelann, 1794. liodes laticollis Fall, 1910, by monotypy); Chaetoceble Sainte-Claire-Deville, 1899: 292 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 35

Fig. 41. Anisotoma horni Wheeler, habitus, dorsal. A common and typical species of genus in eastern United States. 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

(Type species: Agathidium pilosum Sainte-Clai- bodies capable of rolling into compact, ball- re-Deville, 1899, by original designation) (sub- like shapes when disturbed. genus). A complete bibliography and synonymic Cyphoceble Thomson, 1859: 59 (Type species: list for Agathidium is provided by Newton Anisotoma staphylaeum Gyllenhal, 1810 (ϭ A. (1998) along with a discussion of some no- nigrinum Sturm, 1807, by original designation) menclatural problems including the obser- (subgenus). Saccoceble des Gozis, 1886: 17 (Type species: vation that the original name for this taxon, Agathidium discoideum Erichson, 1845, by Volvoxis, is not a homonym as supposed. original designation) (synonym of Cyphoceble). Volvoxis is the oldest name for the genus, Euryceble HlisnikovskyÂ, 1964: 123 (Type species: and historically the International Code of Agathidium antennatum HlisnikovskyÂ, 1964, by Zoological Nomenclature (the Code) would original designation) (subgenus). have required either resurrection of the name Macroceble Angelini, 1993: 30 (Type species: Volvoxis for this genus or a special ruling by Agathidium shermathangense Angelini and De the Commission for conservation of the ju- Marzo, 1981) (subgenus). nior name Agathidium and suppression of Microceble Angelini and De Marzo, 1986: 439 Volvoxis. However, according to article 23.9 (Type species: Agathidium grouvellei Portevin, of the newest Code (ICZN, 1999), the pre- 1907, by orginal designation) (subgenus). vailing usage of a junior name may be main- Neoceble des Gozis, 1886:16 (Type species: tained when the senior name has not been Agathidium marginatum Sturm, 1807, by orig- inal designation) (subgenus). used as a valid name after 1899 (to our Stigmoceble Hlisnikovsky 1964: 119 (Type spe- knowledge true of Volvoxis) and the junior cies: Agathidium longicorne Portevin, 1908, by name has been used as a presumed valid original designation); Angelini, 1986 (synony- name in at least 25 works published by at mized with Neoceble). least 10 authors in the preceding 50 years Rhabdoelytrum HlisnikovskyÂ, 1964:31 (Type spe- and a span of not less than 10 years (true of cies: Agathidium sexstriatum Horn, 1880, by Agathidium, see bibliography above). There- original designation) (subgenus). fore, of the two names Volvoxis Kugelann and Agathidium Panzer, the second is the val- DIAGNOSIS: Agathidium may be distin- id name. guished from other Agathidiini in North and Agathidium has historically been divided Central America by the following combina- into subgenera, several of which were pro- tion of characters: antennal club usually dis- posed as recently as the 1980s and 1990s tinctly 3-segmented (5-segmented in a few (Angelini, 1993; Angelini and De Marzo, taxa, including A. rusticum Fall); anterior 1986). Subgenera recognized for the Palearc- clypeal margin usually not extending beyond tic fauna can be used to organize the Nearctic anterolateral margins of frons (extending be- fauna, though with considerable dif®culty in yond frons in the A. sexstriatum group and some cases, and they may be identi®ed using several species of the A. pulchrum and A. Angelini's (1995) key. However, because our concinnum groups); supraocular carina ex- fauna remains poorly known, and monophyly tending from margin of clypeus or clypeal of the subgenera as currently classi®ed re- region posteriorly to well behind eye delim- mains equivocal, we have decided to not rec- iting a postocular temporum that may be ognize subgenera and instead arrange the short and inconspicuous or long and very species treated here into informal species conspicuous; elytral punctation usually con- groups. Our proposed species groups would fused or absent, occasionally serial, but with loosely belong to previously recognized sub- fewer than nine series and series often ob- genera in the following way: A. brevisternum scured by dense confused punctures between; group ϭ A.(Neoceble) des Gozis; A. onis- prothorax convex; elytra usually convex, coides group ϭ A.(Agathidium); A. revol- sometimes dorsoventrally compressed. vens group ϭ A.(Neoceble) (centered around DISCUSSION: The name Agathidium is de- A. nigripenne Fabricius); A. concinnum rived from ``agathidion'', Greek for ``small group ϭ A.(Cyphoceble) Thomson; A. pul- ball'', and is quite appropriate for many spe- chrum group ϭ A.(Neoceble). Our A. iota cies of the genus which have broad convex and A. compressidens species groups do not 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 37 appear to ®t well into any of the described different from Kilian's (1998). Further anal- subgenera. yses incorporating more comprehensive tax- Especially problematic are those species on and character sampling are likely to result described by Horn and later placed in their in different results from each of these. Thus, own subgenus, A.(Rhabdoelytrum), by Hlis- we would regard any nomenclatural change nikovsky (1964). According to our scheme based on Kilian's (1998) phylogeny to be in- of species groups, the A. sexstriatum group judicious at this time. In addition, whereas would belong to this subgenus. According to we would agree that such a conclusion as she the characters used by Angelini and de Mar- presents would call for synonymy of the two zo (1987b) to distinguish Agathidium from names, doing so would likely serve little ex- Stetholiodes, the species of A.(Rhabdoely- cept to make Agathidium even more hetero- trum) would key to Stetholiodes. For now, geneous in character combinations than it al- these controversial and primitive forms are ready is. We think that were Agathidium or retained in Agathidium although there is mer- Anisotoma eventually found to be paraphy- it in considering a different combination. letic, it would be more appropriate to break Considering only our fauna, such a combi- up these two huge groups of species into nation would appear to make Agathidium genera rather than synonymizing them. species monophyletic, based in part on the Agathidium are sometimes misidenti®ed in relatively excavate clypeus or clypeal region, collections with the following taxa, from while making monophyly of Stetholiodes un- which they may be distinguished by charac- certain and based primarily on symplesiom- ters given parenthetically: Anisotoma (®g. orphic character states. 41: Agathidium with head not narrowed im- There have been two cladistic analyses in- mediately behind eye [except in some A. pul- cluding members of this tribe. Considering chrum and A. sexstriatum species groups] the world genera of the tribe, and a small and with antennal club rarely approaching 5- data set of adult characters, Angelini and segmented); Clambidae (Agathidium never Peck (2000) proposed the following relation- with enlarged hind coxal plates); cybocepha- ships: (Anisotoma ϩ (Amphicyllis ϩ (Lio- line Nitidulidae (Agathidium with front cox- dopria ϩ (Cyrtoplastus ϩ (Sphaeroliodes ϩ ae conical, not transverse); acanthocerine (Stetholiodes ϩ (Afroagathidium ϩ (Pseu- Scarabaeidae (Agathidium with antennae not doagathidium ϩ (Besuchetionella ϩ (Agath- lamellate); sphaeridiine Hydrophilidae idium))))))))). Relationships among non- (Agathidium with club not so compact). Agathidium taxa are beyond the scope of this During this revision three species previ- work, but see Miller and Wheeler (in press) ously placed in Agathidium, A. cognatum for a discussion of their conclusions. Matthews, 1887: 77, A. parile Fall, 1934: Kilian (1998) also conducted a cladistic 117, and A. parvulum LeConte, 1878: 598, analysis of agathidiine taxa, but one using were discovered to belong to a separate ge- larval instead of adult characters. She sug- nus along with several undescribed Central gested synonymizing Anisotoma with Agath- and South American species. These species idium based on the conclusion that Agathi- are treated in a separate revision (Miller and dium species are nested in several places Wheeler, in press). within Anisotoma. However, Kilian's analy- sis was based on a relatively limited data set KEY TO SPECIES GROUPS OF including only characters from larvae and AGATHIDIUM only those from the relatively few species of agathidiines known from larvae (including 1. Mesosternum subhorizontal, in approximately only several species of Agathidium and An- same plane as metasternum (®g. 43) . . . 2 Ð Mesosternum concave posteriorly, anterior isotoma and one species of Liodopria). An- portion clearly in different plane from me- other recent analysis by Angelini and Peck tasternum (®g. 46) ...... 4 (2000) (see above) used adult characters cod- 2(1). Size extremely small (TBL ϭ 1.15±1.51 ed from many more taxa representing a mm); tarsal formula 4±4±3 in both male greater diversity within Agathidiini. They and female; without oblique femoral ca- found a solution to the phylogeny radically rinae on metasternum; male without me- 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 42±51. Characters of Agathidium: Figs. 42, 43. Mesosternum, horizontal and anteriorly cari- nate: 42, ventral view; 43, lateral view. Fig. 44, head with short postocular temporum and with protruding clypeus. Figs. 45, 46. Mesosternum, anteriorly concave and noncarinate, subvertical between mesocoxae: 45, ventral view; 46, lateral view. 47. Head with long postocular temporum and without protruding clypeus. 48. Metasternum with paired oblique lateral lines (``femoral lines''). Figs. 49±51, body forms: 49, noncontractile and posterolateral angles of pronotum de®ned; 50, partially contractile and postero- lateral angles of pronotum partially de®ned but apically rounded; 51, highly contractile and posterolateral angles of pronotum not de®ned.

tasternal fovea; southern Mexico (this 3(2). Body highly contractile (®g. 51); metaster- group based on an undescribed species to num with paired oblique femoral carinae be treated in part 2 of this revision) .... which often meet medially along poste- ...... A. iota group rior margin (®g. 48), though these carinae Ð Size larger (TBL ϭ 1.76±5 mm); tarsal for- may be indistinct; usually dorsally im- mula ( 5±5±4, & 5±4±4; with or without punctate or nearly so; humeral angle of oblique femoral carinae on metasternum; elytron broadly rounded ...... male generally with metasternal fovea ...... A. oniscoides group though it may be small; widespread . . 3 Ð Body moderately contractile (®g. 50); me- 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 39

tasternum without paired oblique femoral uated slightly posterad of middle .... lines or with faint traces; frequently with ...... A. compressidens group conspicuous, confused dorsal punctation; Ð Mesosternum generally strongly concave humeral angle of elytron angulate . . . posteriorly, rising nearly vertically be- ...... A. revolvens group tween mesocoxae, anterior portion nar- 4(1). Clypeus distinctly protruding beyond an- rower along center line than posterior terolateral margins of frons, with one- portion and usually without prominent third to one-half of clypeus occuring be- longitudinal carina medially; metaster- yond sides of front of cranium (®g. 44); num relatively broader (MTL/MTW ϭ body not strongly convex dorsally nor 0.19±0.40); with or without horn on male contractile ...... A. sexstriatum group left mandible; male metasternal fovea Ð Clypeus not or only slightly protruding be- generally smaller and situated medially or yond anterolateral margins of frons, at slightly anterad of middle ...... most with less than one-third of clypeus ...... A. pulchrum group protruding beyond sides of front of cra- nium; body generally dorsally convex and AGATHIDIUM SEXSTRIATUM SPECIES moderately to strongly contractile . . . 5 GROUP 5(4). Metasternum extremely short, meso- and metacoxae nearly touching; male often DIAGNOSIS: The A. sexstriatum species with prominent mandibular horn (®gs. 80, group is distinguished by the following com- 81); anterior portion of mesosternum bination of characters: head narrowed behind moderately broad along center line com- eye, without prolonged postocular temporum pared to posterior portion, but without (®g. 44); body not strongly convex dorsally longitudinal carina medially, posterior and not contractile (®gs. 49); elytron with 2± portion of mesosternum moderately con- cave; western United States ...... 8 serial series of punctures; aedeagus with ...... A. brevisternum group ventral operculum divided into two plates; Ð Metasternum not extremely shortened, male tarsal formula 5±5±4, female 4±4±4; meso- and metacoxae at least moderately pronotum with evident posterolateral angles separated by metasternum; male mandib- (®g. 52); clypeus large, separated from frons ular horn present or absent; anterior por- by impressed frontoclypeal suture, protrud- tion of mesosternum generally much ing anteriorly, from one-third to one-half of shorter than posterior portion, if broad clypeus anterad of lateral margins of frons then with prominent longitudinal carina (®g. 52). medially; widespread ...... 6 DISCUSSION: The A. sexstriatum species 6(5). Postocular temporum long and prominent, group includes three similar species ®rst de- when viewed in dorsal aspect extending scribed by Horn (1880). Hlisnikovsky (1964) lateral margin of head posterad of eye about 0.5 to 1.0 times eye length at which proposed the subgenus Rhabdoelytrum for point head is narrowed (®g. 47), in some these species. As discussed above, it is con- species protruding very prominently lat- ceivable that these species are related to erally beyond eye in rounded bulge when Stetholiodes or other genera (Angelini and de head viewed in dorsal aspect (®g. 47) . . Marzo, 1987b; Angelini and Peck, 2000)...... A. concinnum group They are conservatively maintained in Ð Postocular temporum short, extending lat- Agathidium until analyses are completed. eral margin of head posterad of eye less Species of the group are distributed in the than 0.5 times eye length in dorsal aspect western United States. (®g. 44), head narrowed immediately pos- This group includes species that exhibit no terad of eye ...... 7 observed unequivocal synapomorphies, al- 7(6). Mesosternum moderately concave posteri- though the rather short and broad form of the orly, anterior portion as broad as or broader along center line than posterior median lobe of the aedeagus, serial series of portion and with prominent longitudinal punctures, and prominently protruding clyp- carina medially; metasternum relatively eus make these species separable from other short (MTL/MTW ϭ 0.12±0.16); without Agathidium included in this study. The latter horn on male left mandible; male metas- character refers to forms in which the clyp- ternal fovea generally very large and sit- eus protrudes clearly beyond the anterior an- 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Fig. 52. Agathidium sexstriatum Horn, habitus, dorsal (note: differences in angle of observation result in differences in shape and proportion of structures in this image and that in ®g. 67). gles of the frons located dorsad of the anten- symplesiomorphic similarities, these species nal insertions and each side of the clypeus. share a general habitus and many speci®c It is particularly useful in distinguishing the characters with the related genus Anisotoma A. sexstriatum group species from most sim- (cf. ®g. 41, see also Wheeler, 1979b) and oth- ilarly small-sized Agathidium with short pos- er genera. However, the abrupt 3-segmented tocular tempora. Due to a large number of antennal club and distinct supraocular cari- 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 41

nae distinguish these Agathidium from all TYPE LOCALITY: United States, western Anisotoma species and other genera. Nevada. Circumscription of these species is based DIAGNOSIS: This species can be distin- on external characters, including color and guished from other members of the group by punctation patterns, and male aedeagal the presence of only 2±3 evident series of shape. Although external characters are used punctures medially on elytron, punctures extensively in the key, these features are more confused laterally, punctures moderate- somewhat less reliable than male aedeagal ly large and dense (®g. 56); head and elytron shape and, when possible, male specimens dark reddish-brown, pronotum more pale should be dissected and aedeagal shape com- red; and aedeagus with median lobe attenu- pared with the illustrations for the most re- ated apically (®g. 61). liable identi®cations. DESCRIPTION: Body broadly round, glo- bose, noncontractile (®gs. 58, 59); TBL ϭ KEY TO A. SEXSTRIATUM SPECIES 3.2 mm. Clypeal region, pronotum, and ap- GROUP pendages pale reddish; head, elytra dark red- dish-brown. 1. Punctures serially arranged only medially, 2 or Head narrow (®g. 58); OHW/MDL ϭ 1.4; 3 series clearly evident; median lobe apically dorsal surface with moderately dense punc- attenuate, apex narrowly rounded (®g. 61) 2 ...... A. bistriatum Horn tures (about 10 per 0.01 mm ) (®g. 54); pos- ± Punctures arranged in series, at least six serial tocular temporum short, head narrowed be- rows present and more or less evident be- hind eyes; frontoclypeal suture present, ®ne; coming obscure laterally; median lobe not clypeus with sparse punctures, half as dense attenuate apically ...... 2 as on head (ca. 5 per 0.01 mm2); antenna 2(1). Body elongate oval (®gs. 52, 67); bicolored, short with 3-segmented club; length ratio of pronotum reddish, elytron dark reddish to antennomere II:III ϭ 1:1.4; width ratio of an- nearly black; serial punctures of elytra tennomeres VII:VIII:IX ϭ 0.8:1:1.3; labrum coarse, deeply impressed (®g. 65); male similar in form to that of A. sexstriatum. metasternal fovea paired, large (®g. 66); Pronotum short, not convex; PNW/PNL ϭ median lobe broadly truncate apically (®g. 70) ...... A. sexstriatum Horn 2.1; PNL/PNH ϭ 1.3; PNW/PNH ϭ 2.9; dor- Ð Body broadly oval (®g. 76); concolorous or sal surface with irregularly distributed, pronotum reddish and paler than elytra; sparse and very poorly de®ned punctures, serial punctures of elytra ®ne, obscured generally smaller than those of head and by dense confused punctures between much less distinct (®g. 55). Elytra convex, (®g. 74); male metasternal fovea single, wider than long; ELW/SEL ϭ 1.2, SEL/ELH small (®g. 75); median lobe slightly con- ϭ 1.3, ELW/ELH ϭ 1.5; dorsal surface with stricted subapically, gradually narrowed about 2 or 3 poorly de®ned series of punc- apically to moderately rounded apex (®g. tures near midline, punctation becoming in- 79)...... A. estriatum Horn creasingly confused laterally; punctures larg- er than those of head, surface between punc- Agathidium bistriatum Horn tures with minute, almost imperceptible Figures 53±62 punctules; punctures moderately dense and 2 Agathidium bistriatum Horn, 1880: 304; Leng, large (ca. 4 punctures per 0.01 mm , puncture 1920; Fall, 1934; HlisnikovskyÂ, 1964. ca. 0.04 mm diameter) (®g. 56). Mesoster- num subhorizontal anteriorly; narrow anteri- TYPE MATERIAL: Lectotype (designated or area set off; carinate posteriorly only; ver- here to clarify association of this name with tical between mesocoxae; surface alutaceous; the species), ( in MCZC (G.H. Horn Col- about half length of metasternum. Metaster- lection) labeled ``Nev [with red line on left num with impunctate areas both in front of edge of `N']/ LectoTYPE 3021 [red label]/ and behind center, punctures through middle A. bistriatum Horn/ LECTOTYPE, Agathi- and laterally, and alutaceous pattern laterally. dium bistriatum Horn, des. Q.D. Wheeler & MTL/MTW ϭ 0.2. K.B. Miller, 2001 [red label with black line Male tarsi 5±5±4; pro- and mesobasotar- border]''. someres slightly expanded; metasternal fo- 42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Fig. 53. Agathidium sexstriatum species group geographic distribution: A. bistriatum ϭ ᭡ (᭝ ϭ state record only); A. estriatum ϭ ● A. sexstriatum ϭ Ⅵ (Ⅺ ϭ state record only). 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 43

Figs. 54±59. Agathidium bistriatum Horn: Figs. 54±56, punctation detail: 54, head; 55, pronotum; 56, elytron. 57. Male metasternal fovea. Figs. 58, 59, habitus: 58, dorsal; 59, lateral. Bars ϭ 0.5 mm.

veae paired, large, circular, with pencil of Agathidium sexstriatum Horn ®ne golden setae (®g. 57), MFD ϭ 0.06 mm; Figures 52, 53, 63±71 metafemur not modi®ed; left mandible not Agathidium sexstriatum Horn, 1880: 303; Leng, modi®ed. Aedeagus elongate, narrow; medi- 1920; Fall, 1934; HlisnikovskyÂ, 1964. an lobe attenuated apically (®g. 61); apex re- curved (®gs. 60, 61); operculum platelike, di- TYPE MATERIAL: Lectotype (designated vided (®g. 61); lateral lobes with two sub- here to clarify association of this name to the apical setae nearly equal in length (®gs. 60, species), ( in MCZC (G. H. Horn Collec- 61). tion) labeled ``Nev [with red line on left edge Female tarsi 4±4±4. Spermatheca bulb of N]/ Lectotype 3025 [red label]/ A. sexst- small; neck very long, strongly curved and riatum Horn/ LECTOTYPE, Agathidium attenuated to ®ne point apically (®g. 62). sexstriatum Horn, des. Q. D. Wheeler & K.B. DISTRIBUTION: Specimens were examined Miller, 2001 [red label with black line bor- from California and Nevada (®g. 53). der]''. TYPE LOCALITY: United States, western SPECIMENS EXAMINED: UNITED STATES: Cal- Nevada. ifornia: Lassen Co.: 4 Jun 1913, paralectotypes DIAGNOSIS: This species can be distin- (2, MCZC). Nevada: paralectotypes (11, MCZC). guished from A. sextriatum group members DISCUSSION: This species has been rela- by the body being elongate oval (®gs. 52, 67) tively rarely collected, and we did not ex- and dorsoventrally compressed (®g. 68) rel- amine any recently collected specimens. ative to body form of other known species Thus, its status is unknown and nothing is of the group, the head and pronotum gener- known of its biology. ally distinctly red in color, contrasted with 44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

®ne pubescence. Pronotum short and wide, not strongly convex; PNW/PNL ϭ 2.0; PNL/ PNH ϭ 1.2; PNW/PNH ϭ 2.26; dorsal sur- face similar to head with dense, small punc- tures (ca. 10 punctures per 0.01 mm2) (®g. 64); surface shiny. Elytra broad, about as wide as long; ELW/SEL ϭ 1.0; dorsal sur- face with about six serial rows of punctures set in shallow longitudinal impressions; in- tervals with dense punctures nearly equal to those of series; series obscure laterally, punc- tation confused (about 6 punctures per 0.01 mm2), punctures larger than those of prono- tum and head (®g. 65). Mesosternum vertical between mesocoxae; almost indistinct, weak carina medially; length about 0.2 mm; sur- face minutely alutaceous; with narrow ante- rior area set off by suture. Metasternum sur- face smooth, shiny, with dense, irregularly distributed punctures; MTL/MTW ϭ 0.4. Male tarsi 5±5±4; pro- and mesobasotar- someres moderately expanded; metasternal foveae comprised of paired shallow depres- Figs. 60±62. Agathidium bistriatum Horn, sions with dense pencils of setae (®g. 66); genitalia: Figs. 60, 61, aedeagus: 60, lateral; 61, MFD ϭ max. about 0.07 mm; metafemur un- ventral. 62. Spermatheca. modi®ed; left mandible unmodi®ed. Aedea- gus with median lobe broad, slightly re- curved distally in lateral view (®g. 69); black elytra, the elytra with more than 6 dis- broadly truncate (®g. 70); apical pores fan- tinct rows of serial punctures, coarse and ning out from wide central area (®g. 70); deeply impressed, and the aedeagus with me- operculum divided, each part narrow, trun- dian lobe broadly truncate (®g. 70). cate (®g. 70); lateral lobes shorter than me- DESCRIPTION: Body elongate oval, not con- dian lobe, terminating just beyond operculum tractile, only little convex (®g. 68); TBL ϭ (®gs. 69, 70). 2.3±2.5 mm. Color of head, prothorax, and Female tarsi 4±4±4. Spermatheca bulb appendages pale red; of elytra dark reddish- narrow and neck subapically widened (®g. brown; head sometimes darker than prono- 71). tum. DISTRIBUTION: Specimens were examined Head narrow, more strongly narrowed be- from Nevada, several sites in California, and hind eyes (®g. 67); OHW/MDL ϭ 1.3; dorsal southwestern British Columbia (®g. 53). surface with moderately dense (ca. 10 per 0.01 mm2), small, irregularly distributed SPECIMENS EXAMINED: CANADA: British Co- punctules (®g. 63); postocular temporum lumbia: Squamish, 25 Jul 1980, FIT, SA Marshall, RS Anderson (1, CNCI). short; frontoclypeal suture nearly absent; sur- UNITED STATES: California: state only (5, face of clypeal region paler in color than rest MCZC); Yosemite Valley, 26 May 1928, FE of dorsum and nearly impunctate; antenna Blaisdell (1, CASC); Yosemite Valley, 8 Jun 1930 short, not reaching posterior margin of pron- (4, CASC); Marin Co.: SP Taylor State Park en- otum; length ratio of antennomeres II:III ϭ trance, 9 Mar 1963 (2, EMEC); Tehama Co.: 2 1:1.5; width ratio of antennomeres VII:VIII: mi SW Lassen Lodge, Paynes Ck, 6 Dec 1986, IX ϭ 0.7:1:1.4; labrum narrowed apically 3700Ј, Douglas ®r leaf litter, DS Chandler (3, (length/width at base ϭ 0.55; at apex ϭ 0.9); CASC); 6 mi W Log Springs Mendocino Natl apex shallowly emarginate medially; with Forest, 29 Nov 1986, 5200Ј, oak leaf litter, DS 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 45

Figs. 63±68. Agathidium sexstriatum Horn: Figs. 63±65, punctation detail: 63, head; 64, pronotum; 65, elytron. Fig. 66, male metasternal fovea. Figs. 67, 68, body form: 67, dorsal; 68, lateral. bars ϭ 0.5 mm.

Chandler (1, CASC). Nevada: state only, paralec- inconspicuously arranged into series, the totypes (3, MCZC). confused elytral punctation consists of two DISCUSSION: This species has been collect- kinds of punctures, small, dense punctures ed from oak and Douglas ®r forest litter. El- and minute punctules on integument between evation records are from 3700 to 5200 ft. (®g. 74), the elytral integument with weak alutaceous pattern, the male metasternum Agathidium estriatum Horn with single, small, round anteromedian fovea Figures 53, 72±79 (®g. 75), and the aedeagus elongate, median lobe gradually narrowed apically in ventral Agathidium estriatum Horn, 1880: 304; Leng, view (®g. 79). 1920; Fall, 1934; HlisnikovskyÂ, 1964. DESCRIPTION: Body broadly rounded, con- TYPE MATERIAL: Holotype, ( in MCZC vex, not contractile (®gs. 76, 77); TBL ϭ 2.2 (G. H. Horn Collection) labeled ``Garland mm. Color dark reddish-brown, pronotum, Col 30.6 [`30.6' handwritten]/ HoloTYPE clypeus, labrum, and appendages, except an- 3023 [red label, `3023' handwritten]/ A. es- tennal club, paler reddish. triatum [handwritten]/ HOLOTYPE, Agathi- Head narrow, narrowed behind (®g. 76); dium sexstriatum Horn, 1880 [red label with dorsal surface with moderately dense punc- black line border]''. Because only a single tures (ca. 20 per 0.01 mm2); punctures larger specimen was available to Horn (1880), it is than those of pronotum, smaller than those a holotype by monotypy. of elytra; with minute secondary punctules TYPE LOCALITY: United States, Colorado, irregularly distributed between punctures Garland (probably Fort Garland, Costilla (®g. 72); punctures smaller and less dense on Co.). clypeus; postocular temporum indistinct; DIAGNOSIS: This species is distinguishable frontoclypeal suture distinct; length ratio of by the elytron with confused punctures, only antennomere II:III ϭ 1.3:1; width ratio of an- 46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

region with dense, poorly de®ned alutaceous pattern. Male tarsi 5±5±4; pro- and mesobasotar- someres moderately expanded; metasternal fovea small, single, round, anteromedial in position, with short tuft of setae (®g. 75); MFD ϭ 0.03 mm; metafemur not modi®ed; left mandible not modi®ed. Aedeagus elon- gate and subparallel-sided (®gs. 79); median lobe narrowed apically, with slight subapical constriction (®g. 79); operculum consisting of two small plates (®g. 79); lateral lobes short, with unequal pair of subapical setae (®gs. 78, 79). Female tarsi 4±4±4. DISTRIBUTION: This is the most broadly distributed member of the group. Specimens were examined from British Columbia, Sas- katchewan, Arizona, California, Colorado, Montana, New Mexico, Oregon, and Wash- ington (®g. 53).

SPECIMENS EXAMINED: CANADA: British Co- lumbia: 2 mi E Squamish (1, CNCI); Midday Val- ley, Merritt, 18 May 1925, J Stanley (1, CASC); Manning Pk E Gate, 2 Jun 1984, lodgepole pine Figs. 69±71. Agathidium sexstriatum Horn: forest, FIT, D Miller (1, CNCI); Vancouver, B Ste- Figs. 69, 70, aedeagus: 69, lateral; 70, ventral. 71. phens (1, CASC). Saskatchewan: Cedar Hills Spermatheca. Prov. Park (1, CNCI). UNITED STATES: Arizona: Barfoot Park, Chiricahuas, 18 Jul 1963, PSB (1, CASC); Bar- foot Park, Chiracahuas, 18 Jul 1963, PSB (4, tennomeres VII:VIII:IX ϭ 0.9:1:1.1; labrum CASC); Barfoot Park, Chiricahuas, 18 Jul 1963, separated by broad membrane; subquadrate, PSB (1, CASC); Cochise Co.: Rustler Park, 7 Sep apically emarginate; with very sparse, ®ne 1976, FG Andrews (9, FGAC); Graham Co.: Hos- pubescence; punctures smaller and less dense pital Flat Pinaleno Mts, 2 Aug 1965, 8950Ј,on on clypeus. Pronotum short, wide, and not Daldinia-like fungus, HB Leech (6, CASC). Cal- very convex (PNW/PNH ϭ 2.2; PNL/PNH ifornia: Amador Co.: 1 mi W Pine Grove, 24 Jun 1975, leaf litter mixed hardwood conifer forest, A ϭ 1.6); dorsal surface with moderately dense Newton, M Thayer (1, MCZC); Calaveras Co.: punctures, with minute secondary punctules 2.7 mi N Camp Connell, 4 May 1976, litter, ber- (®g. 73). Elytra very broad and convex; lese, SC Kuba (1, FGAC); Eldor Co.: 2 mi E Lake ELW/SEL ϭ 1.0, SEL/ELH ϭ 2.0, ELW/ Edson Dam, 12 Jul 1971, Ponderosa pine duff, DS ELH ϭ 1.9; dorsal surface with dense irreg- Chandler (1, CASC); Lassen Co.: Norval Flats, ular punctures larger than those of head; ar- 13 May 1920, 5500Ј, JO Martin (2, CASC); Mono eas between punctures with minute irregular- Co.: White Mts S Fk Cottonwood Crk, 18 Jun ly distributed punctules; punctures dense (ca. 1987, 9200Ј, pitfall, D Giuliani (1, FGAC); Shasta 9 per 1 mm2) (®g. 74). Mesosternum ¯at, Co.: Buckhorn Summit, 15 Feb 1983, oak duff, nearly vertical between mesocoxae; weak, TR Haig (1, FGAC); Trinity Co.: Junction City, indistinct carina medially; anterior part set 12 Mar 1981, TR Haig (2, FGAC); Douglas City, 23 Jan 1980, oak duff, berlese, TR Haig (1, off by distinct suture; surface densely and FGAC); Douglas City, 6 Mar 1975, TR Haig (1, minutely alutaceous. Metasternum large, FGAC); Tulare Co.: Sequoia Natl Park, Mineral MTL/MTW ϭ 0.8; punctures indistinct to King, 25 May 1984, 7700Ј, R Baranwoski (4, absent anteriorly, moderately dense posteri- LUND); Kaweah, 10 Apr 1907 (1, CASC); Se- orly and laterally; impunctate anteromedial quoia Natl Park, Halstead Crk, 23 May 1984, 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 47

Figs. 72±77. Agathidium estriatum Horn: Figs. 72±74, punctation detail: 72, head; 73, pronotum; 74, elytron. 75. Male metasternal fovea. Figs. 76, 77, habitus: 76, dorsal; 77, lateral. Bars ϭ 0.5 mm.

7000Ј, R Baranwoski (1, LUND); Tuolene Co.: 5 and A. bistriatum, given proper illumination mi W Dardanelle, 18 Aug 1953, white ®r and ce- six or more serial rows of punctures can gen- dar, GA Marsh (1, CASC). Colorado: Garland, 30 erally be seen among the dense and slightly Jun, Hubbard and Schwarz (3, USNM); Garland, smaller punctures present between them. 30 Jun (5, MCZC); Ouray, 1 Jul 1897, 8000Ј,HF Wickham (2, USNM); Ouray, 1 Jul 1897, 8000Ј Some specimens, possibly teneral ones, have (1, MCZC). Montana: Madison Co., Hidden Lake the head and especially pronotum reddish in Ranch (1, USNM). New Mexico: Torrance Co.: 6 contrast to distinctly darker reddish elytra. In mi SW Manzano, Red Can. Camp, 17 Jun 1979, some of these bicolored specimens (e.g., 9000Ј, spruce-®r litter, S and J Peck (3, PECK). Ouray, Colorado; Norval Flats, California) Oregon: Lake Co.: Quarts Mt, 14 Jun 1984, 1600 the apex of the median lobe is more gradu- m, R Danielsson (1, LUND); Wasco Co.: Wap- ally narrowed than in most other specimens initia Pass, 11 Jul 1975, 4000Ј,onFuligo septica, and the ventral operculum is slightly more A Newton, M Thayer (1, MCZC). Washington: Pierce Co.: Mt Rainier Natl Park, 4.7 mi W Long- elongate. In the most extreme cases, speci- mire, 20 Jul 1975, 2200Ј,onFuligo septica,A mens from Cochise Co., Arizona, have the Newton, M Thayer (3, MCZC); Skaminia Co.: pronotum rather brightly orange-red in sharp Gifford Pinchot Natl Forest, 20 Jul 1984, old contrast to the dark head and elytra. Another growth, D Thomas (1, PECK); Yakima Co.: 8 mi specimen, from Douglas City, Trinity Co., SW Tieton RS Snoqualmie NF Bear Crk, 11 Jun California, is similar, but with reddish clyp- 1973, WJ Turner (1, WSUC). eus and labrum also. The latter material is so DISCUSSION: Depending upon variation brilliantly bicolored as to be confused with among individuals and the angle of light un- A. sexstriatum initially. The dense elytral der which specimens are examined, evidence punctation, smaller serial punctures, and less of serial punctation is variable. Whereas se- parallel-sided elytra distinguish such speci- ries are not as prominent as in A. sexstriatum mens from A. sexstriatum. Unfortunately, all 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

AGATHIDIUM BREVISTERNUM SPECIES GROUP

DIAGNOSIS: The A. brevisternum group species may be recognized by the following combination of characters: minute size, ex- treme convexity, strong contractility (®gs. 80, 81), tusked left male mandible (not pre- sent on all males) (®gs. 80, 81), exception- ally short metasternum with meso- and me- tacoxae nearly touching, aptery, dispropor- tionately large and broad head (®g. 80), ar- ticles of antennal stem very elongate, narrow, parallel-sided with subapical whorl of setae. DISCUSSION: Although the extremely short- ened metasternum of this group is undoubt- edly apomorphic and unusual within the ge- nus, its potential synapomorphic signi®cance is less convincing. This is because such shortened metasterna are correlated with ¯ightlessness in other Coleoptera and, like other loss characters, can mask convergence. Nonetheless, this short metasternum com- bined with the minute size, disproportionate- ly large cranium, and horned left male man- dible combine to make the species assigned Figs. 78, 79. Agathidium estriatum Horn, ae- deagus: 78, lateral; 79, ventral. to this group distinct and easily identi®ed. The ventral operculum of the median lobe is very thin and fragile and, after clearing, is easily overlooked. Care must be taken in pre- these specimens are females so that their paring the aedeagus so as to not make the conspeci®city with other A. estriatum rests operculum too transparent. A pair of sclero- on these external features alone. Considering tized structures on the endophallus are more this structural variation, the broad geographic evident in cleared specimens than is the oper- distribution of the species, and the paucity of culum. existing specimens, infraspeci®c variation The majority of these species are identi®- and the possibility of more than one species able at this time based only upon the shape here should be reconsidered when suf®cient of the median lobe. numbers of specimens have accumulated. Other specimens, however, are more uni- KEY TO A. BREVISTERNUM SPECIES formly darkly colored, aside from pale mark- GROUP ings of the pronotum. A series of specimens from Arizona (Hos- 1. Rami of operculum broad, not strongly diver- pital Flat) are labeled as having been col- gent apically (®g. 84) ...... lected from a ``daldinia-like fungus'', which ...... A. brevisternum Fall could easily describe Lycoperdales slime- Ð Rami of operculum narrow, strongly divergent apically (®gs. 92, 93, 95) ...... 2 molds. The species has also been reported 2(1). Rami of operculum with medial fold, apices from the myxomycete, Fuligo septica (Lawr- entirely separated (®gs. 92, 93) ..... ence and Newton, 1980). Specimens have ...... A. rhinocerellum, new species been collected from a variety of forest litter Ð Rami of operculum without medial fold, types, including ponderosa pine, white ®r, ce- apically connected by truncate, hyaline dar, spruce, lodgepole pine, and others. Ele- shield (®g. 95) ...... vation records are from 2200 to 9200 ft...... A. dioperculum, new species 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 49

Figs. 80. Agathidium brevisternum Fall, habitus, dorsal.

Agathidium brevisternum Fall ten]/ M.C.Z. Type 24030 [red label]/ HO- Figures 80, 81, 82±85 LOTYPE, Agathidium brevisternum Fall, Agathidium brevisternum Fall, 1934: 122; Hatch, 1934 [red label with black line border]''. 1957. TYPE LOCALITY: United States, California, Humboldt Co., Eureka. TYPE MATERIAL: Holotype, ( in MCZC la- DIAGNOSIS: This species is distinguished beled ``Eureka 29.5 Cal [`29.5' handwritten]/ from other members of the group by male HS Barber Collector/ (/ H. C. FALL COL- genitalia with deeply emarginate operculum LECTION/ TYPE brevisternum [`TYPE' of median lobe with apices convergent (®g. with red underline, `brevisternum' handwrit- 84). The carina bordering the anterior margin 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Fig. 81. Agathidium brevisternum Fall, habitus, lateral. of the cranium is less distinctly sinuate above dering eye; antenna with abrupt 3-segmented the bases of the antennae than in A. rhino- club; antennomere III elongate, narrow, lon- cerellum which, in combination with the rel- ger than II, subequal to length of I; II:III ϭ atively simple, platelike halves of the oper- 1:1.8; VII, VIII subequal, small, submonili- culum distinguish A. brevisternum from that form; VII:VIII:IX ϭ 0.7:1:1.4; frontoclypeal species. suture absent, extremely faint hint of suture DESCRIPTION: Body very broadly rounded, under high magni®cation; surface of head strongly convex, and partially contractile shiny, impunctate; under high magni®cation, (®gs. 80, 81); TBL ϭ 2 mm. Color reddish- sparse, microscopic punctules exist bearing testaceous; dorsum somewhat reddish; anten- very ®ne setae; labrum narrow, elongate, na pale, concolorous; venter and appendages with blunt apex at most slightly emarginate testaceous. medially. Pronotum very broad, convex; an- Head large, very wide, broadly rounded in terior margin broadly rounded and protrud- front, more or less quadrate and dorsoven- ing anterad medially; posterior angles weakly trally ¯attened (®gs. 80, 81); OHW/MDL ϭ de®ned; surface shiny, nearly impunctate, as 2; eyes very large, elongate, situated on sides on head, except punctures slightly less dense. of head, with large, coarse facets numbering PNW/PNL ϭ 1.6; PNL/PNH ϭ 1.1; PNW/ less than 36; postocular temporum short, in- PNH ϭ 1.8. Combined elytral width, in dor- conspicuous; OHW/PHW ϭ 1.1; anterior sal view, about equal to length (ELW/SEL ϭ marginal carina conspicuous from edge of la- 1); in posterior view, elytral apex attenuate; brum to point posterior to eye, narrowly bor- sutural stria evident in apical half; surface 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 51

Fig. 82. Agathidium brevisternum group geo- Figs. 83±85. Agathidium brevisternum Fall, graphic distribution: A. brevisternum Fall ϭ ⅷ; A. genitalia: Figs. 83, 84, aedeagus: 83, lateral; 84, dioperculum, n.sp. ϭ Ⅵ; A. rhinocerellum, n.sp. ventral. 85. Spermatheca. ϭ ᭡. to one another; lateral lobes narrow, termi- shiny, nearly impunctate, as on pronotum; nating at point of median lobe constriction at epipleuron very wide, bordered by sharp ca- about apex of operculum, with paired sub- rinae, ending shortly before apex of elytron. apical setae and moderately dense, irregular- Mesosternum large, simple; without lateral ly distributed pores along length (®g. 83±84); carinae; without median carina; surface mi- operculum broad, ¯at, paired, together nar- croscopically alutaceous; MSL/MTL ϭ 3.2. rowed apically, slightly emarginate between Metasternum very short medially, sides at at apex, each with about 6 subapical pores, widest point shorter than mesosternum me- 4 in oblique row (®g. 84); endophallus ap- dially, narrowed to bands between meso- and parently with two membranous apical digits; metacoxae, narrower than one-half maximum with paired, median, slightly C-shaped scler- width of metacoxa; surface with microscopic ites with minute, blunt, toothlike median alutaceous pattern; MTL/MTW ϭ 0.12. ridges. Male tarsi 5±5±4; pro- and mesobasotar- Female tarsi 5±4±4; spermatheca with someres slightly expanded, with spatulate neck very long, slender, and twisted (®g. 85). ventral setae; metasternal fovea median, DISTRIBUTION: This species is known from punctiform, with few long setae; metafemur areas of northern California, particularly in narrow, without tooth; left mandible usually the northwest corner of the state, and western with dorsally directed, curved, apically blunt Oregon (®g. 82). tusk arising dorsally from side near base of SPECIMENS EXAMINED: UNITED STATES: Cal- mandible (®gs. 80, 81); with tuft of apical ifornia: Del Norte Co.: Crescent City, 24 Apr setae. Aedeagus with median lobe short, bul- 1991, TR Haig (7, CASC); 2 mi N Ft Dick, 21 let-shaped, pointed, narrowed before apex Nov 1953, VD Roth (1, CASC); 6 mi NE Cres- (®g. 84); apical pores number approximately cent City, 25 Jun 1978, J Schuh, L and N Herman 16, ducts longitudinally oriented and parallel (1, AMNH); Humboldt Co.: Eureka, 7 Jun, HS 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 86±90. Agathidium rhinocerellum, n.sp.: Figs. 86±88, punctation detail: 86, head; 87, prono- tum; 88, elytron. Figs. 89, 90, habitus: 89, dorsal; 90, lateral. Bars ϭ 0.5 mm.

Barber (1, MCZC); Arcata Comm. Forest, 4 Dec ington Co.: 10 mi N Forest Grove, 9 Apr 1967, 1962, berlese, J Pinto (4, CASC); Berry Summit, EM Benedict (1, CASC). 12 Dec 1973, oak duff, berlese, TR Haig (2, DISCUSSION: All measurements except CASC); Kneeland, 1 Sep 1979, redwood duff, berlese, TR Haig (8, CASC); Weott, 17 Mar 1976, those of antennomere lengths and widths redwood duff, TR Haig (1, CASC); Willow were taken from the holotype. Antennal mea- Creek, 15 Mar 1979, oak duff, TR Haig (1, surements are from a female from Eureka, CASC); Mendocino Co.: Westport, 14 Dec 1973, California (MCZC). This species has been bark and moss, berlese, TR Haig (2, CASC); Trin- collected from a variety of different litter ity Co.: 8.5 mi E Salyer, 12 Apr 1971, oak litter, types including oak, ®r, pine, alder, maple, berlese, TR Haig (1, CASC); Del Loma, 12 Mar redwood, and general wood and leaf litter. 1980, Douglas ®r duff, TR Haig (3, CASC). Elevation records are from 600 to1900 ft. Oregon: New Era, 30 Apr 1954, deciduous duff, V Roth (1, RLWE); nr Bridge, 13 Oct 1954, VD Agathidium rhinocerellum Wheeler and Roth (1, MCZC); Curry Co.: 3 mi N Shasta Creek Miller, new species Rogue River, 10 Mar 1972, 600Ј, oak duff, E Ben- edict (2, CUIC); Jackson Co.: Applegate, 19 Jun Figures 82, 86±90 1958, duff, K Goeden (1, RLWE); 3.5 mi S Ruch TYPE MATERIAL: Holotype, ( in OSUC la- 2 mi from Hwy 238, 13 Nov 1971, 1500Ј, pine beled ``Ore., Lane Co. Spensers Butte 2 mi oak ®r duff, E Benedict (1, CUIC); Josephine Co.: S Eugene 23 Nov. 1959 John D. Lattin/ HO- O'Brien, 18 Dec 1971, 1600Ј, live oak duff, E LOTYPE Agathidium rhinocerellum Wheel- Benedict (1, CUIC); Lane Co.: Dolly Varden For. er and Miller, 2002 [red label with black line Cpgd., 4 Mar 1972, 1100Ј, alder maple, E Bene- dict (1, CASC); Dolly Varden Forest Cpgd, 4 Mar border]''. 1972, 1100Ј, alder maple, E Benedict (4, CUIC); TYPE LOCALITY: United States, Oregon, Leaburg, 10 Nov 1974, G Cooper (1, CASC); Lane County, 2 miles S Eugene. Linn Co.: 2 mi S 2 mi E Cascadia, 29 Apr 1972, DIAGNOSIS: This species is distinguished 1900Ј, maple duff, E Benedict (3, CUIC); Wash- from other A. brevisternum group members 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 53

rowly bordering eye, sharply angled and ter- minating immediately posterior to eye, sin- uate above base of antenna; antenna with abrupt 3-segmented club; antennomere III elongate, narrow, longer than II, subequal to length of I; II:III ϭ 1:1.3; VII, VIII subequal, small, submoniliform; VII:VIII:IX ϭ 0.8:1: 1.6; frontoclypeal suture absent, extremely faint hint of suture under high magni®cation; surface of head shiny, impunctate; under high magni®cation, sparse, microscopic punctules exist bearing very ®ne setae (®g. 86); labrum short, wide, blunt at apex. Pron- otum very broad, convex; anterior margin posterior to head emarginate, nearly straight; posterior angles weakly de®ned; surface shiny, nearly impunctate, as on head, except punctures slightly less dense (®g. 87); PNW/ PNL ϭ 0.7; PNL/PNH ϭ 1.2; PNW/PNH ϭ 1.7. Combined elytral width, in dorsal view, slightly greater than length; ELW/SEL ϭ 1.02; in posterior view, apex attenuate; su- Figs. 91±93. Agathidium rhinocerellum, n.sp., tural stria evident in apical half; surface aedeagus: 91, lateral; 92, ventral; 93, apex of me- dian lobe, ventral. shiny, nearly impunctate, as on pronotum (®g. 88); epipleuron very wide, bordered by sharp carinae, ending shortly before apex of by the male genitalia with the ventral oper- elytron. Mesosternum large, simple; without culum of the median lobe large, deeply emar- lateral carinae; without median carina; sur- ginate and with the halves narrow, strongly face microscopically alutaceous; posterior divergent, and with a distinct medial, irreg- portion between mesocoxae triangular; MSL/ ular fold (®gs. 91±93). The carina bordering MTL ϭ 1.4. Metasternum very short medi- the anterior margin of the cranium is more ally, sides at widest point shorter than me- distinctly sinuate above the bases of the an- sosternum medially, narrowed to bands be- tennae than in A. brevisternum which, in tween meso- and metacoxae, narrower than combination with the sinuate pair of sclerites one-half maximum width of metacoxa; sur- of the endophallus of the aedeagus (®gs. 91± face with microscopic alutaceous pattern; an- 93), distinguishes A. rhinocerellum from that terior margin between mesocoxae elevated as species. transverse carina; MTL/MTW ϭ 0.14. DESCRIPTION: Body very broadly rounded, Male tarsi 5±5±4; pro- and mesobasotar- strongly convex and partially contractile someres slightly expanded, with spatulate (®gs. 89, 90); TBL ϭ 2.25 mm. Color dark ventral setae; metasternal fovea median, reddish-brown to black, eyes pale, venter and punctiform, with few long setae; metafemur appendages yellowish-brown, antenna pale, narrow, without tooth; left mandible usually concolorous. with dorsally directed, curved, apically blunt Head large, very wide, broadly rounded in tusk arising dorsally from side near base of front, more or less quadrate and dorsoven- mandible, with tuft of apical setae (®gs. 89, trally ¯attened (®gs. 89, 90); OHW/MDL ϭ 90). Aedeagus with median lobe short, par- 1.8; eyes very large, elongate, situated on allel-sided, narrowed at apex (®gs. 91, 92); sides of head, with large, coarse facets num- apical pores reaching margin numbering ap- bering less than 36; postocular temporum proximately 24, with similar number not at- short, inconspicuous; OHW/PHW ϭ 1.02; taining margin, pores more or less longitu- anterior marginal carina conspicuous from dinally oriented and parallel to one another; edge of labrum to point posterior to eye, nar- lateral lobes narrow, terminating at point of 54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 median lobe constriction at about apex of operculum, with paired subapical setae, and moderately dense, irregularly distributed pores along length (®g. 92); endophallus with two membranous apical winglike ¯aps; with paired, median, strongly sinuate scler- ites (®gs. 92, 93). Female tarsi 5±4±4; spermatheca not ob- served. ETYMOLOGY: This species is named from rhino, Greek for ``nose'', keros, Greek for ``horn'', and the diminutive suf®x ``ellum''. The name re¯ects both the rhinoceroslike horn on the left male mandible and the di- minutive size of beetles of this species. DISTRIBUTION: This species is only known from western Oregon (®g. 82). PARATYPES: UNITED STATES: Oregon: Doug- las Co.: 20 mi ENE Glide, 23 Mar 1961, 2450Ј, moss and ground litter, D Fellin (1, CASC); Lane Co.: Spensers Butte, 2 mi S Eugene, 23 Nov 1959, JD Lattin (1, CASC). DISCUSSION: One female from Douglas Co., Oregon, is tentatively assigned to this species because the supraantennal carina is Figs. 94, 95. Agathidium dioperculum, n.sp., more strongly sinuate above the antennal in- aedeagus: 94, lateral; 95, ventral. sertion. All measurements were taken from the holotype with the exception of antennom- eres VII, VIII, and IX. Because the apical vided into two elongate plates diverging api- portion of the antenna of the type was not cally, with halves connected by a broad, trun- visible, apical antennomeres of the paratype cate, hyaline plate (®g. 95). were measured instead. One specimen was DESCRIPTION: Body very broadly rounded, collected from ``moss and ground litter''. A strongly convex, and partially contractile; single host record is from Fuligo septica TBL ϭ about 2.2 mm. Color reddish-brown; (Russell, 1979). antenna pale, concolorous; venter and ap- pendages testaceous. Agathidium dioperculum Wheeler and Head large, very wide, broadly rounded in Miller, new species front, more or less quadrate and dorsoven- Figures 82, 94, 95 trally ¯attened; OHW/MDL ϭ 2.0; eyes very large, elongate, situated on sides of head, TYPE MATERIAL: Holotype, ( in AMNH with large, coarse facets numbering less than (deposited from California Department of 36 in dorsal view; postocular temporum Food and Agriculture) labeled ``CALIF:Del short, inconspicuous; OHW/PHW ϭ 1.1; an- Norte Co. Crescent City IX-19±1978 terior marginal carina very conspicuous from T.R.Haig,Coll. [date handwritten]/ Berlesed edge of labrum to point posterior to eye, nar- from Redwood Duff/ HOLOTYPE Agathi- rowly bordering eye; antenna with abrupt 3- dium dioperculum Wheeler and Miller, 2002 segmented club; antennomere III elongate, [red label with black line border]''. narrow, longer than II, subequal to length of TYPE LOCALITY: United States, California, I; II:III ϭ 1:1.8; VII, VIII subequal, small, Del Norte Co., Crescent City, redwood duff. submoniliform; VII:VIII:IX ϭ 0.7:1:1.4; DIAGNOSIS: This species is separable from frontoclypeal suture absent, extremely faint other A. brevisternum group members by the hint of suture under high magni®cation; sur- male median lobe with operculum deeply di- face of head shiny, nearly impunctate; under 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 55

Fig. 96. Agathidium depressum Fall, habitus, dorsal. high magni®cation, sparse, microscopic Pronotum very broad, convex; anterior mar- punctules exist bearing very ®ne setae; la- gin broadly rounded; posterior angles weakly brum narrow, elongate, distally tapered with de®ned; surface shiny, nearly impunctate, as blunt apex slightly emarginate medially. on head, except punctures slightly less dense; 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 97, 98. Agathidium depressum Fall: 97, habitus, lateral; 98, male metasternal fovea.

PNW/PNL ϭ 1.8; PNL/PNH ϭ 1.6; PNW/ region between two sclerotized ``arms'' con- PNH ϭ 2.8. Combined elytral width, in dor- nected by truncate, transparent, extremely sal view, about equal to length (ELW/SEL ϭ ®ne, membranelike structure (®g. 95); lateral 1.3); in posterior view, extreme apex atten- lobes gradually narrowed apically, subapical uate; sutural stria evident in apical half; sur- setae similar in size (®gs. 94, 95) (foreshort- face shiny, nearly impunctate, as on prono- ened due to angle of vision in ventral draw- tum; epipleuron very wide, bordered by ing, ®g. 95). sharp carinae, ending shortly before apex of Female tarsi 5±4±4; spermatheca not ex- elytron. Mesosternum large, simple; without amined. lateral carinae; without median carina; sur- ETYMOLOGY: This species is named from face microscopically alutaceous; MSL/MTL di, Latin for ``two'', and operculum, Latin for ϭ 2.1. Metasternum very short medially, ``lid'', in reference to the deeply divided sides at widest point shorter than mesoster- operculum of the median lobe in males of num medially, narrowed to bands between this species. meso- and metacoxae, narrower than one- DISTRIBUTION: This species is only known half maximum width of metacoxa; surface from the type locality in extreme northwest- with microscopic alutaceous pattern; MTL/ ern California (®g. 82). MTW ϭ 0.11. PARATYPES: UNITED STATES: California: Del Male tarsi 5±5±4; pro- and mesobasotar- Norte Co.: Crescent City, 24 Apr 1991, pine duff, someres slightly expanded, with spatulate berlese, TR Haig (4, CASC, deposited from Cal- ventral setae; metasternal fovea median, ifornia Department of Food and Agriculture). punctiform, with few long setae; metafemur DISCUSSION: This form is hypothesized to narrow, without tooth; left mandible with be a distinct species largely because the con- dorsally directed, curved, apically blunt tusk nective structure between the two sclerotized arising dorsally from side near base mandi- lateral arms of the operculum was not seen ble; with tuft of apical setae. Aedeagus with in individuals of related species. Because this short, stout median lobe, narrowed at apex in form appears to by sympatric with related ventral view, curved ventrad with apex re- species and because variation in some struc- curved dorsad (®g. 95); operculum with tures have been observed, including the num- sclerotized basal and lateral areas forming ber of pores on the operculum, it is possible deeply emarginate, divided parts, the apices that this is simply a variant of A. brevister- of which bear two pores and the medial sur- num. The type series was collected from pine faces of which are not excavated, the distal duff. 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 57

Fig. 99. Geographic distribution of Agathidium revolvens group species: A. depressum Fall ϭ ⅷ; A. dubitanoides, n.sp. ϭ ૽; A. dubitans Fall ϭ Ⅵ; A. jasperanum Fall ϭ ᭡. 58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

AGATHIDIUM REVOLVENS SPECIES KEY TO A. REVOLVENS SPECIES GROUP GROUP

DIAGNOSIS: Body convex, somewhat elon- 1. Elytron with evident serial rows of punctures gate, elytra sometimes distinctly dorsoven- ...... 2 trally compressed; pronotum more or less Ð Elytral punctation confused, without serial convex; body partially contractile; color rows, or, at most, with indistinct serial rows nearly uniform, pale to dark reddish-brown, of punctures largely obscured by dense, rarely nearly black; head with postocular confused intervening punctures ...... 6 2(1). Body relatively broadly ovate (®g. 108); temporum more or less prominent, always punctures of head (®g. 104) small and much shorter than eye; tarsal formula male: sparse, slightly smaller and less dense female 5±5±4:5±4±4; mesosternum nearly than those of pronotum which are also horizontal between mesocoxae, divided into relatively small and weakly impressed anterior and posterior parts, anterior part with (®g. 105) ...... A. revolvens LeConte distinct median carina; metasternum without Ð Body relatively elongate, dorsoventrally paired oblique lines or, if present, no more compressed; density of punctures of head than very short, indistinct traces; elytra usu- and pronotum approximately equal or ally distinctly punctate, punctures sometimes punctures of head larger than those of in more-or-less evident serial rows. pronotum ...... 3 3(2). Punctures of head larger than those of pron- DISCUSSION: Our A. revolvens species group appears to be related to certain Pale- otum (®gs. 110, 111); median lobe in ventral aspect slender, abruptly narrowed arctic members of the subgenus A.(Neoce- subapically, apex narrowly rounded (®g. ble) centered around A. nigripenne Fabricius. 117) ...... A. jasperanum Fall The group, as interpreted here, includes sev- Ð Punctures of head and pronotum approxi- en species recognized by Fall (1934) plus mately same size; median lobe broader, several species described as new. Fall (1934) not abruptly narrowed subapically, with separated these species into two informal apex more broadly rounded ...... 4 groups. One group included species that, like 4(3). Elytron with serial rows of punctures often A. revolvens, are relatively broadly oval. The irregularly double, punctation very dense, other group included putatively related spe- most punctures separated by no more than cies that are comparatively dorsoventrally 0.5±1.5ϫ puncture diameter; median lobe compressed and elongate, a form most fully in ventral aspect subapically broad, not expressed in A. depressum. The distinction constricted, evenly narrowed to rounded apex (®g. 120) ..... A. depressum Fall between the comparatively broader, more Ð Elytron with rows of punctures comprised convex forms and the elongate, more de- of series of single punctures, punctures pressed forms is not consistent, however, and less dense, most punctures separated by this supposed contrast is useful only in com- 1±2ϫ puncture diameter; median lobe in parative descriptions of pairs of species. ventral aspect subapically distinctly con- In general, the species in this group pos- stricted, apex more narrowly rounded . . sess easily discerned differences in shape of ...... 5 the aedeagus, and dissection of males pre- 5(4). Endophallus with apical arms of median ar- sents the best avenue for species identi®ca- mature blunt, divergent, without distinct tion. However, there are a number of external tooth between or with tooth inconspicu- differences, including subtle variation in the ous, basal and lateral arms blunt (®g. 130) type and degree of punctation on head, pron- ...... A. dubitans Fall otum and elytron, body shape, and the male Ð Endophallus with apical arms of median ar- mature acutely pointed, convergent, with metasternal foveae. These characters are less distinct, sharply pointed tooth between consistent and require critical examination of basal and lateral arms acutely pointed specimens. The following key relies heavily (®g. 131) ...... A. dubitanoides, n.sp. on external characters. However, most cer- 6(1). Punctures of head less dense than those of tain species identi®cation is best done by dis- pronotum ...... 7 section of male specimens and comparison of Ð Punctures of head about as dense as those aedeagi with the illustrations. of pronotum ...... 8 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 59

7(6). Body broadly ovate (®g. 136); male with TYPE MATERIAL: Holotype (by monotypy), metasternal fovea absent or, at most, & in MCZC [LeConte Collection] labeled weakly present (®g. 135); punctures of ``[small green disc]/ Type 3178 [red and head larger than those of pronotum (®gs. white label, number handwritten]/ A. revol- 132, 133); rami of operculum of median vens Lec./ HOLOTYPE, Agathidium revol- lobe broad, each half with large subapical laterally directed hook (®g. 139) .... vens LeConte, 1850 [red label with black line ...... A. omissum Fall border]''. Ð Body relatively elongate (®g. 145); male TYPE LOCALITY: Thought to be north shore with metasternal fovea in large depressed of Lake Superior (United States, Minnesota, anteromedial area of metasternum (®g. or Canada, Ontario) (exact locality un- 144); punctures of head smaller than known). those of pronotum (®gs. 141, 142); rami DIAGNOSIS: This species is distinguishable of operculum of median lobe relatively from others in the group by the body broadly broad, but without subapical laterally di- ovate (®g. 107), the punctures on head some- rected hook (®g. 148) ...... what smaller and less dense than those of ...... A cavisternum Fall 8(6). Elytron with punctures large, shallowly im- pronotum, which are also small and weak pressed, comparatively sparse (®g. 151); (®gs. 104, 105), the punctation of the elytron metasternal fovea in large, round shallow (®g. 106) coarser and denser than that of depression (®g. 152); median lobe gen- pronotum and head, the elytral punctation erally slender or arrowhead-shaped and confused, but with more-or-less evident se- narrowly rounded apically (®g. 156); rami rial arrangement of punctures (®g. 106), the of operculum narrow medially, broadly sutural stria, and approximately three addi- expanded and truncate apically (®g. 156) tional parallel, incompletely impressed striae...... A. virile Fall The head has a single pair of setose punc- Ð Elytron with punctures ®ne, dense, more tures near the posterolateral corners of the deeply impressed; fovea small, not in large depression; median lobe and oper- poorly de®ned clypeus, with these punctures culum various in shape ...... 9 being much larger than others on the head. 9(8). Elytral punctures variable in size, relatively DESCRIPTION: Body broadly elongate oval, small (®g. 159); fovea circular (®g. 160); partially contractile, only moderately convex median lobe generally broadly arrow- (®gs. 107, 108); TBL ϭ 3.6 mm. Color of head-shaped and narrowly rounded api- head and elytron dark reddish-brown, nearly cally (®g. 165); rami of operculum very black, pronotum and clypeal spot paler red- broad throughout length, lateral margins dish-brown, mouthparts and appendages pale approximately parallel (®g. 164) ...... A. conjunctum Brown reddish-testaceous. Ð Elytral punctures nearly uniform in size, rel- Head large, broad, subquadrate (®g. 107); atively large and coarse; fovea transverse with distinct, angulate, short postocular tem- ...... 10 porum; OHW/MDL ϭ 1.8; OHW/PHW ϭ 10(9). Apex of median lobe broad, bluntly round- 1.0; dorsal surface with small, sparse, irreg- ed (®g. 168); rami of operculum con- ularly distributed punctures, surface between stricted medially, apically broad, trun- smooth, shiny with sparse, microscopic cate (®g. 167) ...... punctules (®g. 104); frontoclypeal suture ...... A. angustoperculum, n.sp. nearly absent, incomplete and faintly im- Ð Apex of median lobe narrowed, constrict- ed subapically, apex relatively narrowly pressed, head with single pair of setose punc- rounded (®g. 177); rami of operculum tures much larger than other cranial puncta- with medial margins concave, apices tion situated behind posterolateral ``corners'' curved medially, apicomedial apices of clypeal area; clypeal punctation as on pointed (®g. 177) ...... frons. Antennae of holotype damaged, apical ...... A. falcatoperculum, n.sp. portions missing; antennomere III much lon- ger than II, ratio II:III ϭ 1:2.7; labrum small, Agathidium revolvens LeConte apex rounded (not emarginate), dorsum with Figures 100, 104±109 sparse setae; eye large, conspicuous, faintly Agathidium revolvens LeConte, 1850: 222, 1853; faceted; postocular temporum short, about Horn, 1880; Leng, 1920; Fall, 1934. 0.2ϫ length eye; supraocular ridge from edge 60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Fig. 100. Geographic distribution of Agathidium revolvens group species: A. cavisternum Fall ϭ Ⅵ; A. falcatoperculum, n.sp. ϭ ⅷ; A. revolvens LeConte ϭ ૽; A. omissum Fall ϭ ᭡. 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 61

Figs. 101±103. Geographic distribution of Agathidium revolvens group species: 101, A. virile Fall; 102, A. conjunctum Brown (୍ ϭ state record only); 103, A. angustoperculum, n.sp. 62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 104±108. Agathidium revolvens LeConte: Figs. 104±106, punctation detail: 104, head; 105, pronotum; 106, elytron. Figs. 107, 108, body form: 107, dorsal; 108, lateral. Bars ϭ 0.5 mm. of labrum to point far beyond eye; pair of dense (®g. 105); surface anteromedially with setae above ridge, near anterior end of eye. microscopic, faint alutaceous pattern. Elytra Pronotum very wide, convex; PNW/PNL ϭ broadly rounded; ELW/SEL ϭ 0.9; dorsal 1.6; PNL/PNH ϭ 1.2; PNW/PNH ϭ 1.9; surface with large, irregularly distributed, punctation similar to cranium except less weakly impressed punctures, with very poor- ly de®ned series of slightly larger punctures (®g. 106); sutural stria impressed in apical half; surface between punctures smooth, shiny, with microscopic, faint, sparse, trans- verse lines. Mesosternum relatively large; anterior portion with distinct longitudinal median carina, surface alutaceous, subdivid- ed by oblique line. Metasternum with ®ne, broadly distributed ®nely setose punctures; surface between punctures ®nely alutaceous. Male unknown. Female tarsi 5±5±4. Spermatheca (®g. 109) with neck narrowed at base, swollen medially, and apically pointed; bulb globose, with duct insertion terminal on short protu- berance. DISTRIBUTION: This species is known only from the type locality (®g. 100). Fig. 109. Agathidium revolvens LeConte, DISCUSSION: As Fall (1934) reported, spec- spermatheca. imens later added to the LeConte collection 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 63

Figs. 110±115. Agathidium jasperanum Fall: Figs. 110±112, punctation details: 110, head; 111, pronotum; 112, elytron. 113. Male metasternal fovea. Figs. 114, 115, habitus: 114, dorsal; 115, lateral. Bars ϭ 0.5 mm. as ``A. revolvens'' are not conspeci®c with [red label]/ H.C. FALL COLLECTION/ HO- the unique type specimen taken from the LOTYPE, Agathidium jasperanum Fall, northern shore of Lake Superior by LeConte, 1934 [red label with black line border]''. nor are specimens in the Horn collection de- TYPE LOCALITY: Canada, Alberta, Jasper termined to be ``A. revolvens'' actually as- Park. signable to this species. No additional spec- DIAGNOSIS: This species is distinguishable imens attributable to this species were ex- from others in the group by the body some- amined during this study, either. Thus, A. re- what elongate (®g. 114), the punctures on the volvens remains known from the female type head distinctly larger than those of pronotum, specimen alone. Apical portions of the an- but similar in density (®gs. 110, 111), the tennae of this specimen are missing, and the elytral punctation coarser and denser than on ventral sclerites were not measured. head and pronotum and with some punctures more or less serially arranged (®g. 112), and Agathidium jasperanum Fall the median lobe of the aedeagus elongate and Figures 99, 110±118 narrow, subapically abruptly narrowed, with Agathidium jasperanum Fall, 1934: 109. the apex slender and apically narrowly Agathidium jasperinum Hatch, 1957: 35 (incorrect rounded (®g. 117). Agathidium jasperanum subsequent spelling). is less elongate and has a somewhat less con- TYPE MATERIAL: Holotype, ( in MCZC la- spicuously serially arranged elytral puncta- beled ``Jasper Pk. Alb. VIII 4 `24/ (/ TYPE tion than does A. depressum. jasperanum [`TYPE' underlined in red, `jas- DESCRIPTION: Body elongate oval, strongly peranum' handwritten]/ M.C.Z. Type 24038 dorsoventrally depressed, attenuate posteri- 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

rum, gradually narrowed immediately behind eye, separated from frons by distinct supra- ocular carina running from lateral margin of clypeal region to posterior margin of crani- um; without a larger puncture above each eye; antennomere III longer than IV ϩ V; ratio length II:III ϭ 1:2; VII twice as long, slightly wider than VIII, much smaller than IX; ratio width VII:VIII:IX ϭ 1.1:1:1.4; with abrupt 3-segmented club. Pronotum broad, transversely oval, not strongly convex; with very shallow posterior emargination medial- ly; width/length ϭ 1.6; length/height ϭ 1.2; punctures smaller, slightly more dense than those of head, irregularly distributed (®g. 111); area between punctures smooth, shiny, with sparse, irregularly distributed indistinct micropunctules. Elytra elongate oval, ELW/ SEL ϭ 0.84; punctures coarser, larger, less dense than those of head or pronotum; punc- tures larger than those of head; with several poorly de®ned serial rows of punctures, about a single puncture in width, blending with slightly smaller, irregularly distributed punctures between; smaller punctures some- times nearly forming parallel series; surface between punctures smooth, shiny, with irreg- Figs. 116±118. Agathidium jasperanum Fall, ularly distributed, very sparse, very indistinct aedeagus: 116, lateral; 117, ventral; 118, ventral micropunctules (®g. 112); sutural stria im- operculum. pressed in apical three-fourths; area between sutural striae elevated as slight median ``ca- orly, weakly contractile (®gs. 114, 115); TBL rina''; sutural stria con¯uent with suture be- ϭ 3.7 mm. Color reddish-brown, except el- fore apex; humeral angle broadly rounded, ytra darker, castaneous; cranium with pale obtuse. Mesosternum relatively large; ante- central spot, antenna including club testa- rior portion with longitudinal median carina, ceous; pronotum paler than elytra, slightly surface lightly alutaceous, subdivided by paler than head; venter and tibiae reddish- oblique line. Metasternum with small, very brown. ®ne, widely distributed setose punctures; sur- Head subquadrate, dorsoventrally ¯at- face between densely, ®nely alutaceous to tened, wider than long (®gs. 114, 115), imbricate. OHW/MDL ϭ 1.8; surface shiny, with mod- Male tarsi 5±5±4; pro- and mesobasotar- erately dense, irregularly distributed, small someres enlarged, with spatulate setae be- punctures (®g. 110); labrum small, trans- neath; left mandible without tusk; metafemur verse, emarginate medially, surface punctate with apical posterior tooth, relatively small, with long, ®ne setae; frontoclypeal suture ab- broad, not well de®ned, more or less de®ned sent laterally, present only as lightly im- by apical angle of femur; metasternal fovea pressed, basal arc; mandible with evident anteromedial, moderate-sized, round, de- dorsal suture; dorsal surface shiny, with pressed, with tuft of long, ®ne setae (®g. moderately dense, irregularly distributed 113). Aedeagus elongate, narrow; median small punctures; area between punctures with lobe apically abruptly narrowed, apex slen- irregularly distributed, indistinct micropunc- der, narrowly rounded at apex (®g. 117); tules; eye large, conspicuous, anterolateral, operculum with rami relatively short and ®nely faceted; with short postocular tempo- rounded (®g. 118); lateral lobes wide at base, 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 65

Fall, 1934 [red label with black line bor- der]''. TYPE LOCALITY: Canada, British Colum- bia, Terrace. DIAGNOSIS: Agathidium depressum can be separated from other species of the A. revol- vens group by the following combination of characters: body elongate-oval, dorsoventral- ly ¯attened (®gs. 96, 97); color dark reddish- brown; elytral punctation coarser, denser than pronotal punctation, with several poorly de- ®ned serial rows of punctures, series often comprised of an irregular double row of punctures; aedeagus with median lobe swol- len subapically (®g. 120), operculum with deep, rounded emargination, each lobe of operculum with conspicuous, rounded ter- minal disk (®g. 121); and lateral lobes api- cally ¯attened, spatulate (®gs. 119, 120). DESCRIPTION: Body elongate oval, strongly dorsoventrally depressed, attenuate posteri- orly, weakly contractile (®gs. 96, 97); TBL ϭ about 3.6 mm. Color dark reddish-brown; cranium with pale central spot, antenna red- dish-brown, club slightly darker; pronotum Figs. 119±121. Agathidium depressum Fall, paler than elytra, particularly and broadly aedeagus: 119, lateral; 120, ventral; 121, ventral around margins; elytra paler at apices and operculum. along narrow marginal band; venter dark red- dish-brown, legs paler. Head subquadrate, dorsoventrally ¯at- gradually narrowed, apex spatulate, bisetose; tened, wider than long (®gs. 96, 97), OHW/ endophallus with small pair apical digits, MDL ϭ 1.3; labrum small, transverse, emar- paired larger subapical arms, lateral blunt ginate medially, surface punctate with long, plates (®gs. 116, 117) ®ne setae; frontoclypeal suture incomplete, Female 5±4±4. lightly impressed basal arcuate line only, DISTRIBUTION: Specimens were examined with no vestige of lateral portions of suture; from Alberta and the Yukon, Canada (®g. surface shiny, with moderately dense, irreg- 99). ularly distributed punctures, area between SPECIMENS EXAMINED: CANADA: Alberta: Jas- punctures with irregularly distributed micro- per Pk., 4 Aug 1924, paratype (1, MCZC) Yukon: punctules; a single, shallow puncture larger White Horse, 21 Jun 1924, paratype (1, MCZC). than others of head located above each eye, posterior to middle of eye, just above supra- Agathidium depressum Fall ocular carina; eye large, conspicuous, antero- Figures 96±99, 119±121 lateral, ®nely faceted; with short postocular temporum, gradually narrowed immediately Agathidium depressum Fall, 1934: 108; Hatch, 1957. behind eye, separated from frons by distinct supraocular carina running from lateral mar- TYPE MATERIAL: Holotype, ( in MCZC la- gin of clypeal region to posterior margin of beled ``Terrace BC Mrs WW Hip pisley/ (/ cranium; antennomere III about as long as IV TYPE depressum [`TYPE' underlined in red, ϩ V, ratio of length of II:III ϭ 0.8:1; VII `depressum' handwritten]/ M.C.Z. Type distinctly longer, but about same width as 24035 [red label]/ H.C. FALL COLLEC- VIII, much smaller than IX, ratio of width TION/ HOLOTYPE, Agathidium depressum VII:VIII:IX ϭ 1.1:1:1.3; with abrupt 3-seg- 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 mented club. Pronotum broad, transversely Quebec. Another specimen from Urbana, Il- oval, not strongly convex; with shallow pos- linois, was ``doubtfully referred'' to the spe- terior emargination medially; PNW/PNL ϭ cies by Fall. We did not examine this speci- 1.7, PNL/PNH ϭ 1.6; punctures smaller, less men, but it may be better assigned to A. dub- dense than those of head, irregularly distrib- itans. Fall also assigned two specimens from uted; area between punctures smooth, shiny, California of the type series of A. revolvens with sparse, irregularly distributed micro- in the LeConte collection to A. depressum. punctules. Elytra elongate, dorsoventrally The species is the most broadly distributed compressed; ELW/SEL ϭ 0.83; with coarser, of the species in this group (®g. 99). denser punctation than pronotum; punctures SPECIMENS EXAMINED: CANADA: Alberta: Sib- larger than those of head; with several poorly bold Flats Rec Area, 6 Sep 1981, FIT, RS Ander- de®ned serial rows of punctures, about a sin- son (2, QDWC); Banff, 7 May 1930, O Bryant gle puncture wide, blending with slightly (1, CASC); Whitford L., 18 May 1924, O Bryant smaller, irregularly distributed punctures be- (3, CASC); Demmitt, 3 km W 90 km NW Grand tween; smaller punctures sometimes nearly Prairie, 12 Jun 1984, poplar forest, FIT, S and J forming parallel series; surface between Peck (1, PECK); 11 mi NE Robb, 4 Aug 1985, punctures smooth, shiny, with irregularly dis- lodgepole pine forest, FIT, RS Anderson (1, tributed, sparse micropunctules; sutural stria CNCI); Edmonton, 21 Aug 1917, FS Carr, para- type (1, MCZC). British Columbia: Stanley, K impressed in apical three-fourths; area be- Graham (1, CASC); Stanley, 15 Aug 1932, K tween sutural striae elevated as slight roo¯ike Graham (2, CASC); Stanley, 1932, K Graham (1, median prominence; sutural stria con¯uent CASC). Manitoba: The Pas, 28 May 1930, O with suture shortly before apex; humeral an- Bryant (3, CASC). Quebec: Montreal, paratypes gle broadly rounded. Mesosternum relatively (2, MCZC); Duparquet, 3 Jun 1938, GS Smith (1, large; anterior portion with longitudinal me- CASC); Rigaud, paratype (1, MCZC); Duparque, dian carina, surface alutaceous, subdivided 26 Aug 1934, under Poplar bark, GS Smith (1, by oblique line. Metasternum with small, CASC). moderately dense, setose punctures; surface UNITED STATES: Alaska: McKinley Park, 8 between densely, ®nely alutaceous. Jul 1924, paratypes (2, MCZC); Pligrim Hot Springs nr Nome, 6 Aug 1995, on Stemonitus fus- Male tarsi 5±5±4; pro- and mesobasotar- ca, SL Stephenson (2, CUIC); Pligrim Hot someres enlarged, with spatulate setae be- Springs nr Nome, 6 Aug 1995, on Badhamia sp., neath; left mandible without tusk; metafemur SL Stephenson (6, CUIC); Pligrim Hot Springs nr with apical posterior tooth, relatively small, Nome, 6 Aug 1995, decaying log, SL Stephenson broad, not well de®ned; metasternal fovea (1, CUIC); Seward, 15 Jun 1911, WS McAlpine, very small, anteromedial, transverse, with a paratype (1, MCZC). California: Santa Barbara few setae slightly longer than those of me- Mts, 21 Jul 1892, paratypes (2, MCZC); El Do- tasternal surface (®g. 98). Aedeagus elon- rado Co.: 10 mi E Kyburz, 29 Jun 1975, Ponde- gate, median lobe with apex laterally mi- rosa pine duff, FG Andrews (4, CASC); Inyo Co.: nutely serrate, small, pointed, ventrally re- Sierra Nevada Mts S Fk Lubkin Ck, 5 Apr 1982, 6300Ј, D Giuliani (1, CASC); Nevada Co.: Sa- curved (®gs. 119, 120); apical pores dense, gehen Creek nr Hobart Mills, 24 Jun 1964, SG in double row (®g. 119); operculum broad at Seminoff (1, CASC); Independence Lake, 13 Sep base, deeply bifurcate, emargination round- 1969, FG Andrews (1, CASC); San Bernadino ed, each lobe with distinct elevated oval api- Co.: 30 mi ENE Redlands, South Fk Camp- cal area (®gs. 120, 121), apices spatulate in groundd, Santa Ana River, 13 May 1981, 6300Ј, lateral view (®g. 119); lateral lobe stout, L Herman (1, AMNH); San Bernardino Co.: Lake elongate, apex laterally ¯attened, spatulate Fulmore, Nov 1967, litter, Tullgren funnel, D with setae nearly terminal (®gs. 119, 120); Hagstrum (1, CASC); Snow Creek nr Forest endophallus with paired, large, heavily scler- Home, San Bern. Mts, 16 Jul 1961, in wash, EL otized spines. Sleeper (1, CASC); Siskiyou Co.: Mt Shasta, 30 Oct 1969 (1, CASC). Colorado: Rabbit Ears Pass, Female tarsi 5±4±4. 1 Jul 1953, 8000Ј, under stone, Bryant (1, CASC); DISTRIBUTION: Agathidium depressum was Hinsdale Co.: Martin Ranch, Weminuche Valley, reported by Fall (1934) from British Colum- 14 Aug 1977, FG Andrews (1, CASC). Illinois: bia, Seward and McKinley Park, Alaska, Ed- Urbana, 1 May 1921, CP Alexander, paratype (1, monton, Alberta, and Montreal and Rigaud, MCZC). Nevada: Eureka Co.: Monitor Range, 21 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 67 mi S 28 mi W Eureka, 1987±1988, 8000Ј, D Giu- ϭ about 3.2 mm. Color reddish-brown; cra- liani (1, CASC); Nye Co.: 5 mi N 10 mi E Cur- nium with pale central spot, antenna testa- rant, Currant Summit, 28 Mar 1987, 7000Ј,D ceous, club reddish-brown; pronotum paler Giuliani (1, CASC). Washington: Spokane Co.: than elytra, yellowish-red; venter and tibiae Bald Knob Cmgr. Mt Spokane SP, 24 Jul 1978, reddish-brown, femora testaceous. 4800Ј, WJ Turner (5, WSUC); Whitman Co.: Lyle Grove 8 mi SW Pullman, 29 Mar 1973, under Head subquadrate, dorsoventrally ¯at- bark Pinus ponderosa, SD Berkenkamp (1, tened, wider than long (®gs. 126, 127), WSUC). Wyoming: Park Co.: Yellowstone Natl OHW/MDL ϭ 1.4; labrum small, transverse, Park, Cub Creek, 2 Jul 1984, FG Andrews (1, emarginate medially, surface punctate with CASC). long, ®ne setae; frontoclypeal suture present, DISCUSSION: Specimens from Alaska, indistinct; dorsal surface shiny, with moder- Nome County, Pilgrim Hot Springs (CUIC), ately dense, irregularly distributed small were collected from Stemonitis fusca and punctures (®g. 122); area between punctures Badhamia sp. The species has been collected with randomly distributed micropunctules; from a variety of litter sources including rot- without large punctures behind frontoclypeal ting logs and pine duff. Elevation records are suture; eye large, conspicuous, anterolateral, from 4800 to 8000 ft. ®nely faceted; with short postocular tempo- rum, gradually narrowed immediately behind Agathidium dubitans Fall eye, separated from frons by distinct supra- Figures 99, 122±130 ocular carina running from lateral margin of clypeal region to posterior margin of crani- Agathidium dubitans Fall, 1934: 110. um; without a larger puncture above each TYPE MATERIAL: Holotype, ( in MCZC la- eye; antennomere III longer than IV ϩ V; beled ``(/ TYPE dubitans [`TYPE' under- ratio length II:III ϭ 1:1.7; VII distinctly lon- lined in red, `dubitans' handwritten]/ M.C.Z. ger, slightly wider than VIII, much smaller Type 24036 [red label]/ H.C. FALL COL- than IX; ratio width VII:VIII:IX ϭ 1.4:1:1.5; LECTION/ Agathidium dubitans Fall [hand- with abrupt 3-segmented club. Pronotum written, with red line around label]/ HOLO- broad, transversely oval, not strongly con- TYPE, Agathidium dubitans Fall, 1934 [red vex; PNW/PNL ϭ 1.9; PNL/PNH ϭ 1.3; label with black line border]''. with very shallow posterior emargination TYPE LOCALITY: United States, New Mex- medially; punctures smaller, slightly more ico, Otero Co., Cloudcroft. dense than those of head, irregularly distrib- DIAGNOSIS: Agathidium dubitans can be uted (®g. 123); area between punctures separated from other species in this group by smooth, shiny, with sparse, irregularly dis- the following combination of characters: tributed micropunctules. Elytra broadly body elongate-oval, dorsoventrally ¯attened ovate, SEL/ELW ϭ 1.1 with punctation (®gs. 126, 127); color reddish-brown; elytral coarser, but less dense than that of head or punctation coarser and denser than pronotal pronotum; punctures larger than those of punctation (®gs. 123, 124), with several head; with several poorly de®ned serial rows poorly de®ned serial rows of punctures, se- of punctures, more or less single puncture ries generally comprised of a single row of wide, blending with slightly smaller, irregu- punctures; aedeagus with median lobe in larly distributed punctures between; smaller ventral aspect distinctly constricted subapi- punctures sometimes nearly forming parallel cally (®g. 129); operculum deeply emargin- series (®g. 124); surface between punctures ate, each lobe of operculum apically rounded smooth, shiny, with irregularly distributed, (®g. 129); lateral lobes apically ¯attened, sparse micropunctules; sutural stria im- spatulate (®gs. 128, 129); and endophallus pressed in apical three-fourths; area between with arms of median armature blunt and sutural striae elevated as slight median ``ca- without prominent apicomedial tooth be- rina''; sutural stria con¯uent with suture be- tween apical arms (®g. 130). fore apex; humeral angle broadly rounded. DESCRIPTION: Body elongate oval, strongly Mesosternum relatively large; anterior por- dorsoventrally depressed, attenuate posteri- tion with weak longitudinal median carina, orly, weakly contractile (®gs. 126, 127); TBL surface alutaceous, subdivided by oblique 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 122±127. Agathidium dubitans Fall: Figs. 122±124, punctation details: 122, head; 123, pron- otum; 124, elytron. 125. Male metasternal fovea. Figs. 126, 127, habitus: 126, dorsal; 127, lateral. Bars ϭ 0.5 mm. line. Metasternum with small, moderately basolaterally, and basally directed arms dense, setose punctures; surface between which are short and bluntly rounded, without densely, ®nely alutaceous to imbricate. distinct, pointed tooth between apical arms Male tarsi 5±5±4; with pro- and mesoba- or with short, rounded prominence only (®g. sotarsomeres enlarged, with spatulate setae 130). beneath; left mandible without tusk; metafe- Female tarsi 5±4±4. mur with apical posterior tooth relatively DISTRIBUTION: This species is known from small, broad, not well de®ned, more or less Arizona, New Mexico, Ohio, Pennsylvania, de®ned by apical angle of femur; metasternal and Tennessee. The seemingly disjunct dis- fovea minute, punctiform, set in broad de- tribution of this species from the southwest- pression, anteromedial (®g. 125). Aedeagus ern United States and areas from Tennessee elongate, narrow, ventrally curved; median to Pennsylvania may indicate that the species lobe narrowed at apex, sharply ventrally is more widespread than current collection curved, with lateral pectinate pattern of api- information indicates. cal ducts, with minute serrations in lateral SPECIMENS EXAMINED: UNITED STATES: Ari- view (®g. 128); lateral lobes attain base of zona: 10 mi NW Flagstaff, San Franciso Mts, 18 operculum, apices expanded, abruptly spat- Jul 1979, 9500Ј, spruce ®r aspen meadow, mal- ulate in lateral view, compressed in ventral aise, S and J Peck (1, PECK); 10 mi W Greer Mt, view (®g. 129); operculum bifurcate, lobes Baldy, Ft Apache Indian Res., 3 May 1979, R bluntly rounded apically (®g. 129); endo- Baranowski (2, LUND); Pima Co.: Sta. Catalina phallus with medial armature with apically, Mts, Mt Lemmon, 5 Sep 1974, 9300Ј (1, MCZC). 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 69

in the endophallus of the aedeagus. In A. dubitanoides the apical, lateral, and basal arms of the median armature are acutely pointed and there is a distinctly pointed tooth between the apical arms (®g. 131), whereas in A. dubitans the arms are short and blunt and there is no tooth between the apical arms or this tooth is short and blunt (®g. 130) DESCRIPTION: Body elongate oval, strongly dorsoventrally depressed, attenuate posteri- orly, weakly contractile; TBL ϭ about 3.2 mm. Color dark reddish-brown; cranium with pale central spot, antenna testaceous, club reddish-brown; pronotum medially slightly paler than elytra, distinctly paler around margins; venter and tibiae reddish- brown, femora testaceous. Head subquadrate, dorsoventrally ¯at- tened, wider than long, OHW/MDL ϭ 1.4; labrum small, transverse, emarginate medi- ally, surface punctate with long, ®ne setae; frontoclypeal suture present, indistinct; dor- sal surface shiny, with moderately dense, ir- regularly distributed small punctures; area between punctures with irregularly distrib- Figs. 128±131. Agathidium revolvens group species: Figs. 128±130, A. dubitans Fall, aedea- uted micropunctules; without large punctures gus: 128, lateral; 129, ventral; 130, apex of me- behind frontoclypeal suture; eye large, con- dian lobe and armature of endophallus, ventral as- spicuous, anterolateral, ®nely faceted; with pect. 131. A. dubitanoides, apex of median lobe short postocular temporum, gradually nar- and armature of endophallus, ventral aspect. rowed immediately behind eye, separated from frons by distinct supraocular carina run- ning from lateral margin of clypeal region to Ohio: Champaign Co.: Cedar Bog, 4 mi SW Ur- bana (1, AMNH); Delaware Co.: colr. (1, posterior margin of cranium; without a larger AMNH). Pennsylvania: Jeannette, 16 Sep 2002 puncture above each eye; antennomere III (2, CMNH); Pittsburgh, 24 Aug (1, CMNH). Ten- longer than IV ϩ V; ratio length II:III ϭ 1: nessee: Gr. Smokey Mts, NP (1, AMNH). 1.7; VII distinctly longer, slightly wider than VIII, much smaller than IX; ratio width VII: DISCUSSION: Elevation records are from VIII:IX ϭ 1.4:1:1.5; with abrupt 3-segment- 9300 to 9500 ft. ed club. Pronotum broad, transversely oval, not strongly convex; PNW/PNL ϭ 1.9; PNL/ Agathidium dubitanoides Wheeler and PNH ϭ 1.3; with very shallow posterior Miller, new species emargination medially; punctures smaller, Figures 99, 131 slightly more dense than those of head, ir- TYPE MATERIAL: Holotype, ( in AMNH regularly distributed; area between punctures (deposited from MTEC) labeled ``MONT: smooth, shiny, with sparse, irregularly dis- Gallatin Co. Bozeman Crk 6200Ј 28 APR tributed micropunctules. Elytra broadly 1987 D.L. Gustafson Col./ HOLOTYPE ovate, SEL/ELW ϭ 1.1 with punctation Agathidium dubitanoides Wheeler and Mill- coarser, but less dense than that of head or er, 2002 [red label with black line border]''. pronotum; punctures larger than those of TYPE LOCALITY: United States, Montana, head; with several poorly de®ned serial rows Gallatin Co., Bozeman Creek. of punctures, more or less single puncture DIAGNOSIS: This species is nearly identical wide, blending with slightly smaller, irregu- with A. dubitans except for slight differences larly distributed punctures between; smaller 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 punctures sometimes nearly forming parallel TYPE MATERIAL: Holotype, ( in MCZC la- series; surface between punctures smooth, beled ``Glac. Pk Mont. 10Jy 29/ Cut Bank E. shiny, with irregularly distributed, sparse mi- Mank/ (/ NB.2. p.334 [handwritten]/ TYPE cropunctules; sutural stria impressed in api- omissum [`TYPE' underlined in red, `omis- cal three-fourths; area between sutural striae sum' handwritten]/ M.C.Z. Type 24040/ elevated as slight median ``carina''; sutural Agathidium omissum Fall [handwritten, red stria con¯uent with suture before apex; hu- line around label]/ H.C. FALL COLLEC- meral angle broadly rounded. Mesosternum TION/ HOLOTYPE, Agathidium omissum relatively large; anterior portion with weak Fall, 1934 [red label with black line bor- longitudinal median carina, surface aluta- der]''. ceous, subdivided by oblique line. Metaster- TYPE LOCALITY: United States, Montana, num with small, moderately dense, setose Glacier Park. punctures; surface between densely, ®nely DIAGNOSIS: This species is distinguishable alutaceous to imbricate. from other members of the group by the Male tibia 5±5±4; with pro- and mesoba- body broadly ovate (®g. 136), the punctation sotarsomeres enlarged, with spatulate setae of the head distinctly larger and less dense beneath; left mandible without tusk; metafe- than those of pronotum (®gs. 132, 133), the mur with apical posterior tooth, relatively elytral punctation not arranged serially, the small, broad, not well de®ned, more or less male metafemoral tooth prominent, pointed, de®ned by apical angle of femur; metasternal and located subapically, the male without fovea minute, punctiform, anteromedial. Ae- metasternal fovea (®g. 135), and the oper- deagus elongate, narrow, ventrally curved; culum of the median lobe deeply divided, median lobe narrowed at apex, apex sharply each half with a large apicolateral hook (®g. ventrally curved, with lateral pectinate pat- 139). tern of apical ducts, with minute serrations DESCRIPTION: Body broadly elongate oval, in lateral view; lateral lobe attains base of pronotum convex, elytra dorsoventrally com- operculum, apex expanded, abruptly spatu- pressed (®gs. 136, 137); TBL ϭ 3.4 mm. late in lateral view, compressed in ventral Color of pronotum and apex of elytron tes- view; operculum bifurcate, lobes bluntly taceous, antenna testaceous, concolorous, rounded apically; endophallus with conspic- uous medial armature with apically, basola- club slightly darker, appendages reddish- terally, and basally directed arms which are brown, head, base of elytron, ventral surface acutely pointed, with distinct, pointed tooth darker, nearly black. between apical arms (®g. 131). Head subquadrate (®g. 136) with dorsal Female not examined. transverse depression near posterior margin; ETYMOLOGY: This species is named from OHW/MDL ϭ 1.4; surface with moderately the word dubitans and the suf®x -oides for dense, poorly de®ned, shallowly impressed, the great similarity between this species and irregularly distributed punctures (®g. 132), A. dubitans. surface between punctures alutaceous; eye DISTRIBUTION: This species is known from large, conspicuous, forming anterolateral an- Montana and Idaho (®g. 99). gle of head, facets ®ne; postocular temporum short, less than 0.5ϫ eye length; OHW/PHW PARATYPES: UNITED STATES: Idaho: Cassia Co.: Howell Canyon, 20 Jun 1976, under ®r bark, ϭ 0.9; anterior marginal carina distinct, from RD Allen (1, PECK). Montana: Gallatin Co.: clypeus to postocular temporum; with abrupt Bozeman Crk, 25 Jul 1989, 6000Ј, funnel trap, DL 3-segmented club; antennomere III elongate, Gustafson (1, MTEC). narrow, twice as long as II; II:III ϭ 1:2; VII DISCUSSION: A single elevation record is and VIII short, transverse; width VII:VIII:IX from 6000 ft. ϭ 0.9:1:1.8; frontoclypeal suture absent; la- brum small, narrow, emarginate medially of Agathidium omissum Fall apical margin separated from clypeal region Figures 100, 132±140 by wide membrane. Pronotum broad, con- Agathidium omissum Fall, 1934: 113. vex, anterior margin deeply emarginate to re- Agathidium conjunctum, Hatch; 1957: 36 (mis- ceive head, base of emargination nearly identi®cation). straight; posterolateral angles not de®ned, 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 71

Figs. 132±137. Agathidium omissum Fall: Figs. 132±134, punctation details: 132, head; 133, pron- otum; 134, elytron. 135. Male metasternal fovea. Figs. 136, 137, habitus: 136, dorsal; 137, lateral. Bars ϭ 0.5 mm.

obtusely rounded; surface with punctures surface densely alutaceous; MSL/MTL ϭ smaller than those of head, separated by 1.1; MTL/MTW ϭ 0.3. about 1±3ϫ diameter individual puncture, Male tarsi 5±5±4; with pro- and mesoba- more distinctly impressed than punctures of sotarsomeres expanded with dense spatulate head (®g. 133), surface between punctures setae beneath; left mandible without tusk; shiny, with microscopically nearly invisible metafemur with large, prominent, pointed, alutaceous pattern. Elytra with narrow plicate subapical tooth; metasternal fovea absent margin extending to apex; sutural stria im- (®g. 135). Aedeagus robust, elongate; medi- pressed in apical three-fourths, area between an lobe sinuate in lateral view, curved from sutural striae elevated; elytral surface with base toward apex ®rst ventrally then dorsally dense, poorly de®ned punctures separated by (®g. 138); in ventral aspect gradually nar- about 1±2ϫ individual puncture diameter rowed then abruptly narrowed in apical ®fth (®g. 134); without serial arrangement of (®gs. 139, 140), apex with dense, irregularly punctures; surface between punctures aluta- distributed ducts in fanlike pattern (®gs. 139, ceous; epipleuron narrowed, ending before 140); operculum divided entirely in apical apex; ELW/SEL ϭ 0.8. Mesosternum large; half, partially in basal half, apex of each pro- anterior portion with median carina, surface cess in form of asymmetrical, laterally pro- densely alutaceous, subdivided by oblique duced and recurved hook, with sparse, irreg- line; posterior part with oblique lateral ridg- ularly distributed pores on apical half (®g. es, surface alutaceous. Metasternum large; 139); lateral lobes gradually narrowed, api- with very slight, short trace of oblique lines, ces slightly clavate, with 2 setae, 1 apical, 1 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

genitalia, his concept of A. conjunctum ap- pears to be the same as ours of A. omissum. This may have led to his description of a new species (A. obtusum Hatch) for specimens that we regard to be A. conjunctum (see be- low under treatment of A. conjunctum). This species has been collected from ``squirrel middens under conifer'' and ele- vations from 7700 to 9300Ј.

Agathidium cavisternum Fall Figures 100, 141±148 Agathidium cavisternum Fall, 1934: 111; Hatch, 1957.

TYPE MATERIAL: Holotype, ( in MCZC la- beled ``Sifting moss [`moss' handwritten]/ Terrace BC Mrs WW Hip pisley IV-8±1924 [date handwritten]/ (/ TYPE cavisternum [`TYPE' underlined in red, `cavisternum' handwritten]/ M.C.Z. Type 24031 [red label]/ H.C. FALL COLLECTION/ HOLOTYPE, Agathidium cavisternum Fall, 1934 [red label with black line border]''. TYPE LOCALITY: Canada, British Colum- bia, Terrace. DIAGNOSIS: This species is distinguishable Figs. 138±140. Agathidium omissum Fall, ae- from others in the group by the following deagus: 138, lateral; 139, ventral; 140, apex of character combination: body relatively large median lobe. and elongate (®g. 145); punctation of head ®ner and less dense than on pronotum (®gs. 141, 142); elytral punctation confused, with- subapical, with sparse, irregularly distributed out serially arranged punctures (®g. 143); pores (®gs. 138, 139). with large metasternal fovea placed in a de- Female unknown. pressed area anteromedially on the metaster- DISTRIBUTION: This species occurs in the num (®g. 144); and aedeagus with median central and southern Rocky Mountains (®g. lobe apically slender and elongate, apex 100). blunt (®g. 148). SPECIMENS EXAMINED: CANADA: British Co- DESCRIPTION: Body broadly oval, moder- lumbia: Midday Valley, Merritt, 14 May 1925, Pi- ately dorsoventrally compressed, partially nus ponderosa, J Stanley (1, CASC); Monashee contractile (®gs. 145, 146); TBL ϭ 4.0 mm. Mtn nr. Cherryville, 11 Aug 1982, 1400Ј, M So- Color reddish-brown; elytra darker reddish- renson (2, LUND). UNITED STATES: Colorado: Eagle Co.: 5 mi brown at base approaching black, paler api- E Avon, 25 May 1986, 7500Ј, M Sorenson (2, cally; appendages paler, slightly orange. LUND); Grand Co.: 9 mi SE Jct US 40 and CO Head large, subquadrate, dorsoventrally 14, 21 Jun 1975, 7700Ј, squirrel middens under ¯attened (®gs. 145, 146); surface with mod- conifer, A Newton, M Thayer (1, FMNH). Idaho: erately dense, small, irregularly distributed Twin Falls, Magic Mt, 15 May 1976, AD Miller punctures (®g. 141); areas between punctures (1, FMNH). New Mexico: Bernalillo Co.: 10 mi smooth, shiny, with irregularly distributed E Albuquerque Sandia Mt, Capulin Spg, 21 Aug micropunctules; OHW/MDL ϭ 1.3; OHW/ 1975, 9300Ј, S Peck (1, CUIC). PHW ϭ nearly 1.0; postocular temporum DISCUSSION: Based on Hatch's (1957: pl. short, distinct; eye large, oval, anterolateral, IV, ®g. 20) key and illustration of the male conspicuous, ®nely faceted; supraocular ca- 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 73

Figs. 141±146. Agathidium cavisternum Fall: Figs. 141±143, punctation detail: 141, head; 142, pron- otum; 143, elytron. 144. Male metasternal fovea. Figs. 145, 146, habitus: 145, dorsal; 146, lateral. Bars ϭ 0.5 mm. rina present from lateral edge clypeal region cropunctules, without serial punctures (®g. to approximately one eye-length posterior to 143); sutural stria present, extending from eye; antennomere III long, about as long as slightly forward of half elytral length to next three antennomeres combined; II:III ϭ apex, apically convergent. Mesosternum car- 1:2; antennomere VII larger than VIII, much inate, with lateral oblique lines; MSL/MTL smaller than IX; VII:VIII:IX ϭ 0.9:1:1.7; ϭ about 1.0. Metasternum large, densely mi- with abrupt 3-segmented club. Pronotum nutely alutaceous, with weak transverse ru- large, broad, convex; posterior edge with gulose lines, including in male arced line shallow median emargination; corners broad- forming posterior edge of median depressed, ly rounded; surface with punctation similar smooth area; MTL/MTW ϭ 0.3. to that of head; punctures slightly more Male tarsi 5±5±4; pro- and mesobasotar- dense, area between as on head except with someres strongly expanded, with dense spat- sparse, irregularly distributed, short pairs or ulate setae; left mandible not modi®ed; me- sets of parallel microscopic impressions (®g. tafemur with large tooth located about one- 142); PNW/PNL ϭ 1.5; PNL/PNH ϭ 1.5; fourth length femur before apex, with PNW/PNH ϭ 2.2. Elytra broadly oval, not oblique line between apex of tooth and apex sharply attenuate posteriorly; ELW/SEL ϭ of femur; metasternal fovea large, located about 1.0; humeral angles obtuse, broadly near middle in large, smooth depression, rounded; punctures larger than those of head bearing long setae (®g. 144). Aedeagus with or pronotum, dense, irregularly distributed, median lobe elongate, ventrally curved, api- area between smooth, shiny, with sparse mi- cal portion relatively slender, constricted, 74 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

moss''. Other specimens have been collected from spruce and hemlock forests. Two female specimens reported from the type locality were reported by Fall (1934), but these appear to have been returned to C.A. Frost, are not at the MCZC, and were not examined by us.

Agathidium virile Fall Figures 101, 149±156 Agathidium virile Fall, 1901: 219, 1934. Agathidium hebetatum Hatch, 1957: 35. NEW SYN- ONYM.

TYPE MATERIAL: Agathidium virile: holo- type, ( in MCZC labeled ``(/ Pom Cal Mar. 25.93 [date handwritten]/ TYPE virile [`TYPE' underlined in red, `virile' handwrit- ten]/ M.C.Z. Type 24045 [red label]/ H.C. FALL COLLECTION/ HOLOTYPE, Agath- idium virile Fall, 1901 [red label with black line border]''. Agathidium hebetatum: holotype, (, USNM, labeled ``McMinnville, ORE X-20± 41 [handwritten]/ TYPE ( Agathidium (s.str.) hebetatum 1955ÐM. Hatch [hand- Figs. 147±148. Agathidium cavisternum Fall, written, red label]/ HOLOTYPE Agathidium aedeagus: 147, lateral; 148, ventral. hebatatum Hatch, 1957 [red label with black line border]''. TYPE LOCALITY: Agathidium virile: United slightly expanded subapically, apex blunt States, California, Los Angeles Co., Pomona. (®g. 148), in lateral aspect apex recurved Agathidium hebatatum: United States, dorsally, apical quarter approximately Oregon, Yamhill Co., McMinnville. straight, irregularly shaped along ventral or- DIAGNOSIS: This variable species is distin- i®ce, with almost toothlike projection (®g. guished from others in the A. revolvens spe- 147); operculum deeply emarginate, each cies group by the following combination of part elongate, bluntly rounded (®g. 148); lat- characters: body broadly oval, convex, par- eral lobes blunt (®gs. 147, 148). tially contractile (®gs. 153, 154); dorsal Female tarsi 5±4±4. punctation dense, confused, coarsely to DISTRIBUTION: This species is known from weakly impressed (®gs. 149±151); metafe- British Columbia, Canada (®g. 100). moral tooth apical, broad; aedeagus nar- SPECIMENS EXAMINED: CANADA: British Co- rowed at apex, sometimes slightly widened lumbia: Toco, 28 Apr 1984, R and J Camenisch and spatulate at apex (®g. 156); and ventral (1, PECK); Queen Charlotte Islands, Moresby Is- operculum deeply divided, each lobe apically land, 2.5 mi N Mosquito Lake Campground, 5 Jun truncate (®g. 156). 1984, Spruce forest litter, RS Anderson (1, DESCRIPTION: Body very broadly elongate PECK); Charlotte Islands, Graham Island 1 mi oval, partially contractile, moderately convex NW Tlell, 27 Jun 1984, Sitka spruce, hemlock (®gs. 153, 154); TBL ϭ 3.2 mm. Color of forest, RS Anderson (1, PECK); Stickeen R. Can- head and pronotum reddish-brown; elytra yon, Liebeck (1, MCZC). darker, except apex; metasternum conspicu- DISCUSSION: There have been no reported ously darker; mouthparts and appendages hosts. The type was collected from ``sifting reddish-brown; antenna uniformly yellowish 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 75

Figs. 149±154. Agathidium virile Fall: Figs. 149±151, punctation detail: 149, head; 150, pronotum; 151, elytron. 152. Male metasternal fovea. Figs. 153, 154, habitus: 153, dorsal; 154, lateral. Bars ϭ 0.5 mm. red-brown; coxae, femora, mesosternum, and Pronotum broad, convex; PNW/PNL ϭ 1.6; prosternum paler, somewhat testaceous. PNL/PNH ϭ 1.4; PNW/PNH ϭ 2.2; punc- Head large, broad, subquadrate in shape; tation similar to that of head, a little more head narrowed behind eye, temporum short faintly impressed (®g. 150). Elytra very (®gs. 153, 154) (about three-®fths length of wide; length little more than combined eye), de®ned only by supraocular carina ex- width; only slightly attenuate at apex; humeri tending from posterior end of temporum to not angulate, broadly obtuse; punctures much frontoclypeal suture; OHW/MDL ϭ 1.5; larger than those of head and pronotum, OHW/PHW ϭ about 1.0; dorsal surface with more coarsely impressed, irregularly distrib- sparse, irregularly distributed, weakly im- uted and moderately dense, distance between pressed, small setigerous punctures, surface punctures about 0.5±1.5 diameter of single between with sparse, small, irregularly dis- puncture, surface between smooth, shiny, tributed punctules (®g. 149); eye large; fron- with microscopic, nearly invisible and very toclypeal suture very ®nely impressed, near- sparse micropunctules and random faint ly absent, clypeal region made distinct only curving lines (®g. 151). Mesosternum divid- by subcuticular darkened lines; surface of ed into anterior and posterior parts; anterior clypeus as on frons; without larger setae at part with longitudinal median carina and ®ne posterolateral corners of clypeal region; la- alutaceous sculpturing; with oblique carinae brum small, transverse, very shallowly emar- and lateral, longitudinal carinae. Metaster- ginate medially; antennomere III about as num ®nely alutaceous, with laterally curved, long as IV ϩ V ϩ VI; length II:III ϭ 1:2.2; concentric, rugose ridges, surface covered by VII very slightly wider than VIII, nearly ®ne setae. MSL/MTL ϭ 0.7; MTW/MTL ϭ equal in size; width VII:VIII:IXϭ 1:1:2.4. 0.3. 76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

ventral view, with pair of subapical setae and irregularly distributed pores (®gs. 155, 156); endophallic armature not extensive. Female tarsi 5±4±4. DISTRIBUTION: This species is nearly ubiq- uitous in California and occurs into western Oregon, though it does not appear to occur extensively outside California (®g. 101).

SPECIMENS EXAMINED: UNITED STATES: Cal- ifornia: state only (3, MCZC); Berkeley, 7 Dec 1919 (3, MCZC); Poway (1, CASC); Carmel, 5 Jan 1924 (1, CASC); Willows, 13 Jun 1966, RP Allen (1, FGAC); Santa Barbara, 20 Dec 1980, pan trap, S Miller (4, CUIC); Descanso Jct, 31 Mar 1969 (1, CNCI); Poway, FC Bowdin (1, MCZC); Sequoia Natl Park, 12 May 1979, R Bar- anowski (7, LUND); Yosemite Valley, 8 Jun 1930 (1, CASC); San Diego (1, CASC); Sacramento, 7 Apr 1931, HH Keifer (1, FGAC); Pasadena (8, MCZC); Pomona (1, MCZC); Pasadena, Mar, A Fenyes (1, CMNH); Santa Cruz Mts (2, MCZC); Castro Valley, 27 Mar 1938, CT Sierra (1, FGAC); Stanford Univ., 30 Apr 1950, P Barthol- omew (1, CASC); Pasadena (2, CASC); Alameda Co.: Livermore, 11 Apr 1958, JT Doyen (1, EMEC); Strawberry Can., 11 May 1984, J Whi- te®eld (1, CASC); Corral Hollow, 10 mi E Liv- ermore, 10 Mar 1976, J Doyen (1, CASC); 4 mi SE Livermore, Arroyo Mocho, 24 Mar 1976, FG Andrews (1, FGAC); Oakland, 22 Mar 1952, Quercus agrifolia, H Schuster (1, EMEC); Alpine Figs. 155, 156. Agathidium virile Fall aedea- Co.: 1.2 mi E Monitor Base, 25 Jun 1983, 8000Ј, gus: 155, lateral; 156, ventral. FG Andrews (1, FGAC); Woodfords, 18 Oct 1962 (1, FGAC); Amado Co.: Pine Grove, 5 Mar 1971, FG Andrews, RF Wilkey (1, FGAC); Butte Co.: Male tarsi 5±5±4; with pro- and mesoba- 4.4 mi SW Rackerby, 24 Mar 1983, A Hardy, F sotarsomeres moderately laterally expanded; Andrews (1, FGAC); 4.4 mi SW Rackerby, 6 Feb left mandible not modi®ed; metafemur with 1980, pitfall, AR Hardy (5, FGAC); 3 mi E Chico, conspicuous, broad, nearly apical tooth with 17 Jan 1979, Bermuda grass, DS Chandler (1, QDWC); Colusa Co.: Rumsey, 1 May 1970, oak small median protruding point; metasternal litter, FL Blano (1, FGAC); Contra Costa Co.: Mt fovea in large anteromedial oval depression, Diablo, 24 Nov 1953 (2, EMEC); Antioch, 1 mi with long ®ne setae (®g. 152). Aedeagus with E, 24 Jan 1975, J Doyen (1, CASC); Vine Hill, median lobe elongate, robust, ventrally 12 Apr 1908, FE Blaisdell (1, CASC); Antioch curved, recurved dorsally subapically, apical Natl Wildlife Refuge, 13 Mar 1982, pitfall traps, one-®fth dorsoventrally compressed and lat- Powell (1, CASC); El Dorado Co.: Peavine Ridge erally narrowed before apex; apex of median Rd 7 mi SW Ice House, 16 Apr 1992, Quercus lobe narrowed, slender to more or less arrow- kellogi litter, FG Andrews (1, FGAC); 1 mi E Pa- head-shaped, with fanlike arrangement of ci®c House, 10 Jun 1994, under bark of dead oak, ducts apically (®g. 156); operculum deeply CB Barr (11, CASC); 1.3 mi S Paci®c House, 15 Feb 1991, F Andrews, T Eichlin (3, FGAC); bisected into two pieces, each nearly parallel- Blodgett Exp. For. 14 mi E Georgetown, 8 May sided or slightly expanded apically and trun- 1976, J Doyen (1, CASC); 2 mi SE Latrobe, 18 cate or very broadly rounded at apex, with Apr 1978, oak duff, Stanley, C Kuba (3, FGAC); apical concentration of pores (®g. 156); lat- Blodgett Forest 13 mi E Georgetown, 28 Apr eral lobes long, gradually narrowed before 1976, Quercus kellogi litter, J Doyen (2, CASC); apex that is slightly enlarged, especially in 3 mi W Grizzly Flat, 24 Mar 1982, under oak 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 77 bark, F Andrews (1, FGAC); 7.7 mi SE Virner 1976, oak duff, F Andrews, KS Corwin (1, Blodgett Forest, 29 Mar 1984, under bark Pseu- FGAC); McClure's Beach, 17 Dec 1976, rock dotsuga trifolia, FG Andrews (1, FGAC); 7.7 mi crevices, upper intertidal, J Doyen (1, CASC); SE Virner Blodgett Forest, 29 Mar 1984, under Taylor State Park, 20 Apr 1947, HB Ross (1, bark Ponderosa pine, FG Andrews (2, FGAC); 3 CASC); Samuel P Taylor State Park, 31 Mar mi SW Somerset, 26 Oct 1971, R Wharton (11, 1976, FG Andrews (1, FGAC); Samuel P Tayler FGAC); 3.5 mi NW Rescue, 30 Nov 1976, Quer- State Park, 31 Mar 1976, FG Andrews, TD Ei- cus litter, FG Andrews (1, FGAC); Fresno Co.: 8 chlin (2, FGAC); Novato, 22 Nov 1969, swim- mi W Coalinga Wartham Canyon, 16 Apr 1980, ming pool, T Haig (2, FGAC); Pt Reys Mt Vin- Andrews, Paddock (27, FGAC); Cierro Hills, 16 son, 25 Feb 1982, on myxomycete, FG Andrews Apr 1980, CY Kitayama (17, CASC); 8 mi W (1, FGAC); Taylor State Park, 20 Jun 1947, ES Coalinga Warthan Canyon, 16 Apr 1980, ¯ying at Ross (1, CASC); Mariposa Co.: Tioga Rd, 1.9 mi dusk, Andrews (12, FGAC); 2 mi N Tollhouse, 28 E Crane Flat, 18 May 1976, 6600 Ј, conifer forest Nov 1977, Ceanothus duff, A Gilbert (3, FGAC); litter, A Newton, M Thayer (1, MCZC); Mendo- 2 mi NE Auberry, 13 Feb 1981, AJ Gilbert, HN cino Co.: 18 Mar 1938, J Helfer (1, CASC); 2 Jul Smith (1, FGAC); 9 mi E Coalinga, 20 Feb 1981, 1926 (1, CASC); 29 Jul 1952 (1, EMEC); Hop- Andrews, Gilbert (5, FGAC); Dinky Crk, 5 Apr land Field Sta. HQ Lake, 15 May 1977, J Powell 1970, pine duff, TR Haig (1, FGAC); 8 mi W (1, CASC); Hopland Filed Sta. HQ Lake, 15 May Coalinga, Warthan Canyon, 16 Apr 1980, An- 1979, J Powell (1, CASC); Merced Co.: Los Ba- drews, Paddock (1, FGAC); 9 mi E Coalinga, 20 nos Creek, 1 Apr 1987, 400Ј, pitfall, D Giuliani Feb 1981, pitfall, Andrews, Gilbert (4, FGAC); (2, FGAC); Modoc Co.: 5 mi NE Altura, 20 Mar 15 mi E Fresno, 16 Mar 1984, oak litter, AJ Gil- 1980, oak duff, TR Haig (3, FGAC); 5 mi NE bert (2, FGAC); 11 mi W Coalinga, 28 Oct 1971 Altura, 20 May 1980, Juniperus occidentalis duff, (11, FGAC); Humboldt Co.: Eureka, 27 May TR Haig (4, FGAC); Mono Co.: 6 mi SW Toms 1981, salix litter, TR Haig (1, FGAC); Kern Co.: Place, 8 Aug 1969, 9000 Ј, A Smetana (1, CNCI); 1 mi N McKittrick, 24 Mar 1981, pitfall, RL Aal- Monterey Co.: 19.4 mi E San Lucas, 16 Nov bu (2, FGAC); Kern River, 6 mi E Tupman, 21 1971, Heteromeles arbutifolia, FG Andrews (1, Mar 1975, Atriplex litter, J Doyen (1, CASC); Fra- FGAC); 19.4 mi E San Lucas, 16 Nov 1971, oak zier Park, 13 Jan 1961, EL Sleeper (1, FGAC); duff, FG Andrews (1, FGAC); Preist Valley, 20 Arvin Hills, 9 Mar 1986, under rock, J Schuh (1, Mar 1981, pitfall, Gilbert, Andrews (1, FGAC); MCZC); 12 mi NW Taft, 9 Apr, DB Wahl (1, Arroyo Seco, 9 Feb 1982, under oak bark, F An- AMNH); 1.4 mi S Valley Acres, 30 Jan 1979, drews (1, FGAC); 17 mi E San Lucas, 20 Mar Andrews, Hardy (1, FGAC); Kings Co.: 1 mi SW 1981, pitfall, Gilbert, Andrews (2, FGAC); Mt Kettleman City, 4 Mar 1988, pitfall trap in sand, Palo Exctio, Jun 1972, oak duff, D Giuliani (1, RL Aalbu (1, CNCI); 8 mi SSW Avenal, 29 Apr FGAC); Carmel, 1915 (1, MCZC); 6.7 mi N Hwy 1980, Andrews, Kuba, Paddock (1, FGAC); 8 mi 1, 23 Feb 1984, oak litter, berlese, AJ Gilbert (1, SSW Avenal, 17 Sep 1980, litter at base of oak, FGAC); Cast Rd along Sierra Ck 2.4 mi SE Bixby Andrews, Kuba, Paddock (4, FGAC); 1 mi SW landing, 21 Feb 1989, F Andrews, T Eichlin (1, Kettleman City, 4 Mar 1988, pitfall, RL Aalbu (1, FGAC); Napa Co.: 2 mi N St Helena White's FGAC); 1 mi SW Kettleman City, 4 Mar 1988, Cave entrance, 26 Apr 1981, pitfall, R Aalbu (1, pitfall trap, RL Aalbu (2, FGAC); Tar Canyon SE FGAC); 3 mi W Cordelia, 1 Oct 1971, oak duff, Avenal, 8 Mar 1973, pitfall, Gilbert, Bookout (1, WW Wiard (7, FGAC); 2 mi ENE St Helena, 24 FGAC); 10 mi S Avenal, Sun¯ower Velley, 9 Jan Jan 1981, RL Aalbu (2, FGAC); Nevada Co.: 4 1980, oak, pitfall, A Gilbert (2, FGAC); Tar Can- mi S Rough and Ready, 5 May 1980, under bark yon SE Avenal, 8 Mar 1975, pitfall trap, Gilbert of Quercus kellogi, JT Doyen (38, CASC); and Bookout (1, FGAC); 10 mi SW Avenal, 6 Feb Orange Co.: Buena Park, 16 Feb 1967 (1, FGAC); 1980, oak juniper duff, A Gilbert (2, FGAC); 8 Riverside Co.: 0.5 mi SE Aguanga Chaparral mi SSW Avenal, 17 Apr 1980, base of oak, An- Scrub, 7 Jan 1979, pitfall, RL Aalbu (2, FGAC); drews, Kuba, Paddock (1, FGAC); Lake Co.: Menifee Valley, 28 May 1980, JD Pinto (1, Butte Canyon 10 mi SE Middletown, 3 Apr 1981, FGAC); Sacramento Co.: Sacramento, 5 Apr Hardy, Andrews (1, FGAC); Lassen Co.: Raven- 1977, TR Haig (1, FGAC); Nimbus, Alder Creek dale, 16 Apr 1975, FG Andrews (1, FGAC); Los at US Hwy 50, 2 Mar 1990, under bark, CB Barr, Angeles Co.: San Gabriel Canyon, 9 Feb 1961, WD Shepard (1, CASC); 9 Sep 1930, blue oak FG Andrews (1, FGAC); Pine Canyon 6 mi W (1, FGAC); 1 mi E Folsom Humberg Gr, 8 Sep Lake Hughs, 27 Feb 1978, A Hardy, F Andrews 1978, pitfall, S Kuba (1, FGAC); Folsom, 1 Jan (1, FGAC); Madera Co.: 32 mi NE Fresno USDA 1938, Q Tomich (1, EMEC); San Benito Co.: 17 Exp. Station, 25 Jan 1980, woodrat nest, N Smith mi N New Idris, 9 Apr 1982, ptifall, A Gilbert, N (1, FGAC); Marin Co.: Pt Reyes Station, 11 Mar Smith (5, FGAC); 14 mi S San Benito, 16 Nov 78 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

1971 (2, FGAC); 1.8 mi SW Idria, 25 Mar 1981, ter, DS Chandler (3, CASC); 2 mi NE Dales, 13 pitfall, AJ Gilbert, N Smith (5, FGAC); 1.4 mi N Feb 1971, DS Chandler (2, EMEC); Trinity Co.: New Idris, 8 Jun 1981, manzanita, AJ Gilbert N Junction City, 12 Mar 1981, TR Haig (1, FGAC); Smith (1, FGAC); 18.4 mi NE Idris, 25 Apr 1981, Junction City, 04 Jan 1990, madrone duff, TR AJ Gilbert, HN Smith (1, FGAC); 8.2 mi W on Haig (1, FGAC); Weaverville, 15 Apr 1981, litter, Panochee Rd, 1 Jan 1979, AJ Gilbert (1, FGAC); TR Haig (1, FGAC); Del Loma, 5 Jan 1985, Al- San Bernardino Co.: 3 mi E Summit, 2 May 1978, nus duff, TR Haig (10, FGAC); Del Loma, 23 Jan KW Cooper (2, FGAC); 2 mi E Mentone, 15 May 1980, TR Haig (10, FGAC); Big Bar, 3 Aug 1976, 1978, KW Cooper (1, FGAC); 6 mi W Wright- leaf litter, FG Andrews (1, FGAC); 5 mi W Cof- wood, 14 Apr 1986, under bark, F Andrews, TD fee Ck Ranch, 19 Jun 1972, pine duff, DS Chan- Eichlin, A Hardy (2, FGAC); 1.4 mi N New Idris, dler, DP Levin (2, CASC); Tulare Co.: Ash Mt nr 8 Jun 1981, manzanita, pitfall, AJ Gilbert, N Three Rivers, 11 Feb 1983, ¯ume debris trap, R Smith (1, FGAC); 3 mi E Summit, 2 May 1978, Haines (1, FGAC); Ash Mt, 17 Mar 1984, at Neotoma nest, KW Cooper (1, FGAC); Seven ¯ume forebay, R Haines (22, FGAC); Wood L., Oaks, 26 Jan 1963, oak duff, FG Andrews (2, 19 Dec 1947, rotary trap, NW Frazier (4, EMEC); FGAC); 3 mi E Summit, 2 May 1978, Neotoma Kaweah, 16 Nov 1971, oak duff, FG Andrews (1, nest, KW Cooper (9, FGAC); Lake Fulmore, Nov FGAC); 1 Jun 1933, FT Scott (5, FGAC); Ashe 1967, 6000Ј, tullgren funnel, D Hagstrum (1, Mt HQ, 28 Apr 1979, Douglas ®r litter, JT Doyen FGAC); San Diego Co.: Pine Hills, William Heise (9, CASC); Camp Wishon 10 mi NE Springville, Co. Park, 19 Mar 1979, poison oak, berlese, KW 1 May 1979, J Doyen (1, CASC); S FK Camp 13 Cooper (1, FGAC); 4 mi E Campo, 26 Mar 1961, mi SE Three Rivers, 29 Apr 1979, Arctostaphylos EE Lindquist (2, EMEC); 7.5 mi E Julian, Banner litter, JT Doyen (1, CASC); 10 mi SE Three Riv- Crk., 2 May 1981, 2700Ј, L Herman (1, AMNH); ers, S FK Kaweah R., 29 Apr 1979, Quercus litter, Borego, 28 Apr 1955 (1, EMEC); San Luis Obis- JT Doyen (1, CASC); Sequoia Nat. Pk. Huckle- po Co.: Cambria, 29 Oct 1971, oak duff, FG An- berry Meadow, 28 May 1984, 6000Ј, R Bara- drews (1, FGAC); Arroyo Grande, 7 Aug 1987, nowski (1, LUND); Tuolumne Co.: Strawberry, pitfall, J McLaughlin (1, FGAC); 7.5 mi W Si- 23 Jun 1953, JG Rozen (1, EMEC); Ventura Co.: mmler, 30 Jan 1979, pitfall, Andrews, Hardy (12, Moorpark, 23 Mar 1971, oats and barley, Ander- FGAC); Santa Barbara Co.: Carpinteria, 19 Jan son, Johnson (1, FGAC); Yolo Co.: 2 mi W Rum- 1989, J McLaughlin (1, FGAC); Refugio, 7 Apr sey, Cache Crk, 8 Nov 1971, Bermuda grass, 1972, D Teiman (1, FGAC); Santa Clara Co.: Sar- sweeping, DS Chandler (2, CASC); Davis, 15 atoga Gap, 19 Apr 1976, FG Andrews (1, FGAC); Nov 1956, Pyrecantha litter, RO Shuster (4, 24 Apr 1923, ER Leach (1, CMNH); Sveadal, 27 CASC); Yuba Co.: Sierra Foothill Feild Sta 6 mi Apr 1968, A and A Gillogly (1, FGAC); Santa N Smartville, 4 May 1980, under bark Pinus sa- Cruz Co.: 7.5 mi S Los Gatos, Santa Cruz Mts, biniana, JA Powell (4, CASC); Dry Crk 4 mi NW 23 Nov 1963, Irwin, Westcott (1, FGAC); Shasta Smartville, 3 May 1980, under bark Pinus sabi- Co.: Redding, 1 Mar 1980, TR Haig (5, FGAC); niana, JT Doyen (3, CASC); 15 mi E Marysville, 4 mi NE Montgomery Ck, 19 Apr 1984, oak duff, 6 Feb 1980, pitfall, AR Hardy (16, FGAC); 19 berlese, TR Haig (4, FGAC); Redding, 1 Apr Apr 1913 (1, MCZC); Spencerville Wildlife Area, 1979, oak duff, TR Haig (4, FGAC); Redding, 1 9 Apr 1978, F Andrews (7, FGAC). Oregon: Clat- Apr 1980, TR Haig (2, FGAC); 4 mi NE Mont- sop Co.: 5 mi N 7 mi W Elsie, 13 Aug 1973, gomery Ck, 9 Feb 1986, oak duff, berlese, TR 700Ј, duff, E Benedict (1, QDWC); Douglas Co.: Haig (1, FGAC); Sierra Co.: Gold Lake, 14 Jul 16 km E Steamboat, 16 Oct 1972, 3200 Ј, duff, E 1921 (1, CASC); Slano Co.: Travis AFB, 6 Apr Benedict (1, QDWC); 5.5 mi NE Idleyld Pk., 1 1971 (3, FGAC); Sonoma Co.: Trinity, 30 Nov Apr 1972, 1200 Ј, duff, E Benedict (1, QDWC); 1933 (1, CASC); Armstrong Park, 14 Mar 1954, Jackson Co.: 4 mi E Eagle Point, Kanutcham Ck, J Helfer (1, EMEC); Sta Clara Co.: Guadalupe 22 Jan 1972, 1400Ј, madrone oak duff, E Benedict Res, 1 Apr 1973, under bark blue oak, CY Kitay- (2, QDWC); 9 mi N Central Point, 22 Jan 1972, ama (5, CASC); Sta Cruz Co.: S end Sunset Bch, 1400Ј, oak duff, E Benedict (4, QDWC); Jose- 4 May 1973, J Doyen (1, CASC); Stanislaus Co.: phine Co.: 1 mi S, 5 mi W O'Brien, Hwy 99, 18 Knights Ferry, 25 Mar 1987, meadow, Andrews, Dec 1971, 1400Ј, debris, E Benedict (1, PECK); Eichlin, Hard (1, FGAC); La Grange, Oct 1975, Linn Co.: Swamp Mt Rd 2 mi E Cascadia, 29 Apr RP Allen (1, FGAC); La Grange, 5 Oct 1970, at 1972, maple duff and wood, EM Benedict (1, light, RP Allen (11, FGAC); Tehama Co.: 1 mi PECK). SW Dales, 13 Feb 1971, DS Chandler (5, CASC); 15 mi W Red Bluff, 14 Feb 1971, DS Chandler DISCUSSION: The holotype of A. hebetatum (3, CASC); Red Bluff, 16 Feb 1973, TR Haig (1, Hatch (USNM) was examined and found to FGAC); S Fk Battle Creek, 25 Dec 1970, oak lit- be a synonym of A. virile based on similarity 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 79 in punctation and shape of the median lobe Agathidium conjunctum Brown of the aedeagus. This specimen is well within Figures 102, 157±165 the variation of typical members of A. virile. Agathidium conjunctum Brown, 1933: 46; Fall, Agathidium virile is a common and mor- 1934. phologically variable species in the far west- Agathidium obtusum Hatch, 1957: 36. NEW SYN- ern region of the United States. Specimens ONYM. assigned to this species are diverse in punc- tation and, to a certain extent, in the form of TYPE MATERIAL: Agathidium conjunctum: the aedeagus. The elytra are densely punc- holotype, ( in CNCI, labeled ``Langley B.C. tate, but the punctures vary from rather fee- 7.III 1931/ K.Graham [handwritten]/ HO- bly impressed to very deep and coarsely im- LOTYPE Agathidium conjunctum Brown, pressed. Among those with coarse elytral 1933 [red label with black line border]''. punctation, punctures of the head and espe- Agathidium obtusum: holotype not exam- ined. cially the pronotum may be somewhat small- TYPE LOCALITY: Agathidium conjunctum: er but similarly strikingly coarse and deep Canada, British Columbia, Langley. ranging to rather small and decidedly weakly Agathidium obtusum: Canada, British Co- impressed. The apex of the median lobe lumbia, Creston. varies from slender and gradually narrowed DIAGNOSIS: This species is distinguished to more broadly expanded. The branches of from others in the group by the elongate oval the operculum vary from very slender and body, the elytra dorsoventrally compressed elongate to distinctly broader, but generally (®gs. 161, 162), the punctures of the head the branches are narrowed medially with the similar in density to those of pronotum (®gs. apices broadly expanded and truncate. 157, 158), the elytron without serial series of It is possible that additional species exists punctures, punctation dense, confused (®g. within what we regard as A. virile. However, 159), the male metasternal fovea small, cir- no correlations were found among these var- cular (®g. 160), and the aedeagus with the iable traits that suggest a reasonable infer- apex of the median lobe arrowhead-shaped ence of species status for any of the included with a median ventral ridge (®gs. 164, 165). variable specimens, and we regard the spe- DESCRIPTION: Body broadly oval, contrac- cies as simply polymorphic. A male from tile, moderately convex (®gs. 161, 162); TBL Stanford, California (CASC), has a relatively ϭ 2.0±2.5 mm. Color dark reddish-brown, broad median lobe apex, gradually broad- nearly black; clypeal region, perimeter of ened and apically spatulate ventral opercula, pronotum, mouthparts, and appendages in combination with rather weakly impressed slightly paler. pronotal and elytral punctation. In this sense Head broad, subquadrate, dorsoventrally it is somewhat intermediate in condition with ¯attened (®gs. 161, 162); OHW/MDL ϭ 1.6; A. angustoperculum. Considering the known OHW/PHW ϭ 1.1; dorsal surface with mod- variation within A. virile, however, this spec- erately dense, shallow punctures (®g. 157), imen is most reasonably assigned here for the surface between shiny but with microscopic, sparse punctules and slightly alutaceous; present. Another specimen from Santa Clara clypeal suture absent; eyes large, slightly County, California (CNMH), and one from protruding; postocular temporum distinct, Linn County, Oregon (PECK), are tentatively short; antennomere III long, about as long as assigned to this species and have the apical IV ϩ V ϩ VI; length II:III ϭ 1:3; VII slight- portion of the median lobe much more slen- ly broader than VIII; width VII:VIII:IX ϭ 1: der than in most specimens, with the lateral 1:2.1. Pronotum broad, convex; PNW/PNL margins nearly parallel. ϭ 1.6; PNL/PNH ϭ 1.4; PNW/PNH ϭ 2.2; A specimen from McClure's Beach, Ma- without de®ned posterior angles, slightly rine County, California (CASC), was col- emarginate posteromedially; surface with lected from ``rock crevices, upper intertidal'', punctation similar to head (®g. 158). Elytra though this must certainly be an accidental broad, but longer than wide, moderately dor- occurrence for this species. soventrally ¯attened; dorsal surface with 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 157±162. Agathidium conjunctum Brown: Figs. 157±159, punctation detail: 157, head; 158, pronotum; 159, elytron. 160. M male metasternal fovea. Figs. 161, 162, habitus: 161, dorsal; 162, lateral. Bars ϭ 0.5 mm. dense, irregular punctures only slightly larger apex pointed, with ducts obliquely fanning to than those of head and pronotum (®g. 159), each side of midline (®gs. 164, 165); ventral surface between as on head and pronotum, operculum deeply emarginate, each branch including subalutaceous microscopic pattern; long, parallel-sided, and truncate, pores sutural stria present, strongly impressed, sparse and irregular except moderately dense complete. Mesosternum horizontal, with me- in apical transverse line (®g. 165); lateral dian carina; surface densely alutaceous. Me- lobes nearly as long as median lobe, enlarged tasternum surface alutaceous, pattern dense in apical ®fth in ventral aspect, with two laterally. MSL/MTL ϭ 0.7; MTL/MTW ϭ large subapical setae (®gs. 163, 164). 0.4. Female not observed. Male tarsi 5±5±4; with pro- and mesoba- DISTRIBUTION: This species was recorded sotarsomeres moderately expanded; left man- by Fall (1934) from British Columbia, Wash- dible unmodi®ed; metasternal fovea small, ington, ``W.T.'' (Washington Territory?), and round, with ®ne setae, in smooth impunctate Nevada. In this study, numerous specimens area anteromedially (®g. 160); metafemur were examined from southern British Co- gradually widened into blunt, subapical, pos- lumbia south to extreme southwestern Cali- terior tooth de®ned on apical side only. Ae- deagus long, narrow, curved, apical one-®fth fornia and east to southwestern Montana (®g. abruptly narrowed, ventrally curved but 102). arched slightly dorsally (®g. 163); median SPECIMENS EXAMINED: ``W.T.'' [probably Wash- lobe narrowed, distinctly spatulate near apex, ington Territory] (2, MCZC). 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 81

1968, malaise, WJ Turner (2, WSUC); Nevada Co.: Sagehen Crk, 15 Jul 1970, DS Chandler (1, CASC); Sagehen Crk, 24 Jun 1970, DS Chandler (1, CASC); Sierra Co.: 9 mi W Alleghany, 4 Apr 1985, under logs, Pinus sabiniana, F Andrews, A Hardy (2, FGAC); Trinity Co.: 5 mi W Coffee Crk Ranch, 19 Jun 1972, pine duff, DS Chandler, DP Levin (1, CUIC); Yuba Co.: Sierra Foothill Field Station 3 mi N Smartville, 3 May 1980, Quercus litter, JT Doyen (9, CASC). Idaho: Latah Co.: Moscow Mt, 28 Apr 1973, WJ Turner (2, WSUC). Montana: Gallatin Co.: Bozeman Crk (1, MTEC). Nevada: state only (1, MCZC). Oregon: state only (1, MCZC); Jackson Co.: Soda Mt Rd, 7 mi S 13 mi E Ashland, 15 Oct 1972, under willow bark, E Benedict (1, PECK); Umatilla Co.: 25 km E Ukiah, Frazier Camp, 25 Aug 1982, M Sorenson (1, LUND); Wallowa Co.: Wallow Pk St Prk, 8 Jun 1921, under bark, D Ferro (1, WSUC). Wash- ington: Seattle, OB Johnson (1, WSUC); Seattle, 4 May 1912 (2, MCZC); Chelan Co.: Squillchuck St Pk, 8 mi SSW Wenachee, 14 May 1983, 2900Ј, WJ Turner (7, WSUC); Columbia Co.: Tucannon Riv. 22 mi S Mareugo Umatilla NF, 18 May 1974, under bark of Pinus ponderosa, WJ Turner (2, WSUC); Whitman Co.: 8 mi SW Pullman, Lyle Grove, 14 Apr 1974, JA Logan (1, WSUC).

Figs. 163±165. Agathidium conjunctum DISCUSSION: Agathidium obtusum Hatch is Brown, aedeagus: 163, lateral; 164, ventral; 165, synonymized here with this species based on apex of median lobe, ventral. Hatch's (1957) description and illustration of the ventral view of the aedeagus. His speci- mens appear to ®t well within the range of CANADA: British Columbia: Trinity Valley, variation observed for A. conjunctum. 14 May 1929, JR Howell (1, CASC); Midday Val- Hatch's (1957: pl. IV, ®g. 20) concept of A. ley, Merritt, 14 May 1925, J Stanley (1, CASC); conjunctum appears to actually refer to A. Lorna, 10 Jul 1925, Pinus contorta, H Richmond omissum based on the key and his illustration (1, MCZC); Trinity Valley, 1 Jun 1928 (1, CASC). of the male genitalia. UNITED STATES: California: Calistoga, 12 Agathidium conjunctum has been collected Jul 1934, Bryant (1, CASC); Poway, FC Bowdin frequently under bark of various trees, in- (1, MCZC); Butte Co.: Feather Falls, 16 May cluding pine and oak. It has also been col- 1971, DS Chandler (1, CASC); Feather Falls, 16 Feb 1971, RF Lagler (1, WSUC); Feather Falls, lected from various litter types, including oak 16 May 1971, DS Chandler (1, QDWC); El Do- leaf litter and pine duff. Elevation records are rado Co.: 3 mi W Grizzly ¯at, 24 Mar 1982, on from 2900 to 4200 ft. polypore slimemold under oak bark, F Andrews (5, FGAC); 2 mi NE Riverton, 30 Nov 1976, un- Agathidium angustoperculum Wheeler and der bark of Pinus, F Andrews (1, FGAC); 0.5 mi Miller, new species N Stumpy Meadows Lake, 20 Apr 1989, 4200Ј, Figures 103, 166±168 litter from Quercus kellogi, F Andrews, T Eichlin (6, FGAC); 1 mi W Grizzly Flat, 24 Mar 1982, TYPE MATERIAL: Holotype, ( in AMNH under oak bark, F Andrews (10, FGAC); Lake (from California Department of Food and Co.: Adams Springs, 24 Apr 1976, under bark, JF Lawrence (3, MCZC); Marin Co.: 2 mi N Taver- Agriculture collection) labeled ``Sveadal ness, 23 Mar 1965, WJ Turner (1, WSUC); Pt Santa Clara Co., Calif IV-27±1968 [date Reyes Natl Seashore, Mt Vinson overlook, 29 handwritten]/ A. & A. Gillogly collectors/ Nov 1977, F Andrews (2, FGAC); Mendocino Agathidium angustoperculum Wheeler det. Co.: UC Hopland Field Station nr HQ, 13 Apr Q.D.Wheeler ``95/ HOLOTYPE Agathidium 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

nearly black, antennal stem reddish-brown, club darker, legs somewhat paler. Head large, broad, subquadrate in shape; head narrowed behind eye, temporum short (about three-®fths length eye), de®ned only by supraocular carina extending from poste- rior end of temporum to frontoclypeal suture; OHW/MDL ϭ 1.6; OHW/PHW ϭ about 1.0; dorsal surface with irregularly distributed, weakly impressed, small setigerous punc- tures; surface between with very sparse, mi- croscopic, irregularly distributed, nearly in- visible punctules; eye large; frontoclypeal su- ture very ®nely impressed, clypeal region made distinct by subcuticular darkened lines; surface of clypeus as on frons; without larger setae at posterolateral corners of clypeal re- gion; labrum small, transverse, very shallow- ly emarginate medially; antennomere III about as long as IV ϩ V ϩ VI; length II:III ϭ 1:1.8; VII very slightly wider than VIII, nearly equal in size; width VII:VIII:IX ϭ 1: 1:1.8. Pronotum broad, convex; PNW/PNL ϭ 1.7; PNL/PNH ϭ 1.2; PNW/PNH ϭ 2.2; punctation similar to that of head, a little more faintly impressed. Elytra very wide; length about equal to combined width; only slightly attenuate at apex; humeri not angu- Figs. 166±168. Agathidium angustoperculum, late, broadly obtuse; punctures much larger n.sp., aedeagus: 166, lateral; 167, ventral; 168, than those of head and pronotum, more apex of median lobe, ventral. coarsely impressed, irregularly distributed and moderately dense; distance between punctures about 0.5±1.5 diameter of single puncture; surface between smooth, shiny, angustoperculum Wheeler and Miller, 2002 with microscopic, nearly invisible and very [red label with black line border]''. sparse micropunctules and random faint TYPE LOCALITY: United States, California, curving lines. Mesosternum divided into an- Santa Clara Co., Sveadal. terior and posterior parts; anterior part with DIAGNOSIS: Agathidium angustoperculum median longitudinal carina and ®ne aluta- closely resembles A. virile and A. conjunc- ceous sculpturing; with oblique carinae and tum from which it may be distinguished by lateral, longitudinal carinae. Metasternum the following combination of characters: pro- ®nely alutaceous, with laterally curved, con- notal punctures coarse, only infrequently as centric, rugose ridges, surface covered by coarse as in A. virile; male metasternal fovea ®ne setae. MSL/MTL ϭ 0.7; MTL/MTW ϭ small, transversely oval, anteromedial; male 0.4. metafemoral tooth protruding, pointed; apex Male tarsi 5±5±4; with pro- and mesoba- of median lobe of aedeagus spatulate (®gs. sotarsomeres moderately expanded beneath; 167, 168); and rami of ventral operculum left mandible not modi®ed; metafemur with long, narrow, expanded and truncate at apex conspicuous, subapical tooth, broad with (®g. 167). small median protruding point; metasternal DESCRIPTION: Body broadly elongate oval, fovea in large transverse anteromedial oval partially contractile, moderately convex; depression, with long ®ne setae clumped in TBL ϭ 3.0 mm. Color dark reddish-brown, two tufts. Aedeagus with median lobe elon- 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 83 gate, robust, ventrally curved, recurved dor- 8 mi E Gaviota, 7 May 1952, HS Dybas (2, sally subapically, apical one-®fth dorsoven- CASC); Sonoma Co.: 9 Dec 1910, EP van Dyke trally compressed and laterally narrowed be- (1, CASC); 24 Feb 1911, Van Dyke (1, CASC); fore apex (®g. 168); apex of median lobe Tulare Co.: 10 mi SE Three Rivers, S Fk Kaweah spatulate, with fanlike arrangement of ducts R., 3 May 1979, dead Pseudostuga, JT Doyen (2, CASC); Yuba Co.: Sierra Foothill Field Sta, 6 mi apically around margin (®gs. 167, 168); N Smartville, 4 May 1980, 1500Ј, JA Powell (1, operculum deeply bisected into long, narrow CASC); Willow Glen Creek 5 mi SW Browns- pieces, each expanded and truncate apically, ville, 6 May 1980, on Polyporus sulfureus,JT with row of apical pores (®g. 168); lateral Doyen (1, CASC). Oregon: Douglas Co.: 5.5 mi lobes long, gradually narrowed before apex NE Idlewild Pk. Rock Crk Rd, 1 Apr 1972, 1200Ј, which is enlarged, bulbous in lateral view, chinkapin duff, E Benedict (1, PECK). bluntly rounded in ventral view, with pair of DISCUSSION: This species has been collect- subapical setae, with irregularly distributed ed from a variety of litter types including pores (®gs. 166, 167); endophallic armature chinkapin, oak, pine, and ®r. Elevation re- not extensive. cords are from 200 to 2250 ft. Host records Female 5±4±4. include Polyporus anceps and P. sulfureus. ETYMOLOGY: This species is named from the Latin angustus, ``narrow'', and opercu- Agathidium falcatoperculum Wheeler and lum, ``lid'', in reference to the paired, elon- Miller, new species gate, narrow ventral opercula of the median Figures 100, 169±177 lobe of the male genitalia. DISTRIBUTION: This species is found from TYPE MATERIAL: Holotype, ( in FMNH south-central California north to southern labeled ``WASH:Jefferson Co Olympic NP, British Columbia (®g. 103). 0±3.4 mi SW Hoh 400±500Ј VII.16.1975 A.Newton, M.Thayer/ leaf litter mixed hard- PARATYPES: CANADA: British Columbia: nr Mabel Lake, Squaw Valley, 4 Aug 1982, M So- wood-conifer forest/ HOLOTYPE Agathi- renson (1, LUND); 20 mi NE Creston, 16 Aug dium falcatoperculum Wheeler and Miller, 1958, M Sorenson (1, LUND). 2002 [red label with black line border]''. UNITED STATES: California: state only (1, TYPE LOCALITY: United States, Washing- AMNH); Berkeley, 4 Mar 1915, EP van Dyke (2, ton, Jefferson Co., Olympic National Park, CASC); Berkeley, 13 Apr 1961, J Lawrence (1, 3.4 mi SW Hoh. MCZC); Santa Cruz, Jun 1896 (1, MCZC); Ala- DIAGNOSIS: This species very closely re- meda Co.: Laundry Farm, 25 Apr 1907, FE Blais- sembles A. virile and A. angustoperculum, dell (1, CASC); Amador Co.: 1 mi W Pine Grove, from which it may be distinguished by the 24 Jun 1975, mixed hardwood conifer forest, A curved, convergent, and sometimes apically Newton, M Thayer (2, MCZC); Calaveras Co.: 3 overlapping rami of the ventral operculum of mi NW West Point, 20 May 1976, 2250Ј, leaf lit- ter, mixed hardwood conifer forest, A Newton, M the median lobe (®g. 177). Thayer (2, MCZC); El Dorado Co.: 1 mi W Griz- DESCRIPTION: Body very broadly elongate zly Flat, 24 Mar 1982, oak duff, F Andrews (1, oval, partially contractile, moderately convex FGAC); Marin Co.: 3.1 mi NW Inverness, 8 May (®gs. 173, 174); TBL ϭ 3.0±3.1 mm. Color 1976, 200Ј, berlese, litter, old ¯ood debri Alnus of head, pronotum, and elytra red; ventral forest, A Newton, M Thayer (2, MCZC); Taylor- surface red; mouthparts and appendages red- ville (1, MCZC); Taylorville, 28 Dec 1919, JO dish-brown; antenna uniformly reddish- Martin (1, CASC); Tocaloma, 18 May 1952, tree brown; legs reddish-brown. litter, HS Dybas (1, FMNH); Samuel P. Taylor Head large, broad, subquadrate in shape, State Park, 31 Mar 1976, F Andrews (1, FGAC); narrowed behind eye, temporum short (®gs. Nevada Co.: 4 mi S Rough and Ready, 5 May 173, 174) (about three-®fths length of eye), 1980, Quercus kellogi litter, JT Doyen (5, CASC); Wolf Mt 5 mi SW Grass Valley, 6 May 1980, present only as supraocular carina extending Quercus kellogi litter, JT Doyen (27, CASC); Sac- from posterior end of temporum to fronto- ramento Co.: Sacramento, 1 Feb 1971, F Andrews clypeal suture; OHW/MDL ϭ 1.5; OHW/ (1, FGAC); San Luis Obispo Co.: Creston, 2.5 mi PHW ϭ 1.0; dorsal surface with sparse, ir- S, 10 Apr 1961, on Polyporus anceps, J Lawrence regularly distributed, weakly impressed, (1, MCZC); Santa Barbara Co.: Refugio Canyon small setigerous punctures, surface between 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290

Figs. 169±174. Agathidium falcatoperculum, n.sp.: Figs. 169±171, punctation detail: 169, head; 170, pronotum; 171, elytron. 172. Male metasternal fovea. Figs. 173, 174, habitus: 173, dorsal; 174, lateral. bars ϭ 0.5 mm. with few, very small, irregularly distributed and moderately dense, distance between punctules (®g. 169); eye large; frontoclypeal punctures about 0.5±2.0 diameter of single suture very ®nely impressed, nearly obsolete, puncture, surface between smooth, shiny, clypeal region made distinct by subcuticular with microscopic, nearly invisible and very darkened lines; surface of clypeus as on sparse micropunctules and random faint frons; without larger setae at posterolateral curving lines (®g. 171). Mesosternum divid- corners of clypeal region; labrum small, ed into anterior and posterior parts; anterior transverse, very shallowly emarginate medi- part with longitudinal median carina and ®ne ally; antennomere III about as long as IV ϩ alutaceous sculpturing; with oblique carinae V ϩ VI; length II:III ϭ 1:2.2; VII very and lateral, longitudinal carinae. Metaster- slightly wider than VIII, nearly equal in size; num ®nely alutaceous, with laterally curved, width VII:VIII:IXϭ 1:1:2.4. Pronotum concentric, rugose ridges, surface covered by broad, convex; PNW/PNL ϭ 1.6; PNL/PNH ®ne setae. MSL/MTL ϭ 0.8; MTL/MTW ϭ ϭ 1.4; PNW/PNH ϭ 2.2; punctation similar 0.4. to that of head (®g. 170). Elytra very wide; Male tarsi 5±5±4; with pro- and mesoba- length slightly more than greatest width; sotarsomeres moderately expanded laterally; slightly attenuate at apex; humeri not angu- left mandible not modi®ed; metafemur with late, broadly obtuse; punctures much larger conspicuous, broad, subapical tooth with and more coarsely impressed than those of small median protruding point; metasternal head and pronotum, irregularly distributed fovea in large anteromedial oval depression, 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 85

operculum Latin for ``lid'', signifying the curved condition of the rami of the opercu- lum of the median lobe. DISTRIBUTION: This species is distributed in the Paci®c Northwest (®g. 100). PARATYPES UNITED STATES: California: Fresno Co.: Sierra NF Tamarack Ridge, 34 m SE Cal. 168, 16 May 1976, under conifer bark, A Newton, M Thayer (1, FMNH); Mendocino Co.: MacKerricher State Park, 1 Dec 1978, FG An- drews (1, CASC). Washington: Pierce Co.: Mt Rainier NP 4.7 mi W Longmire, 20 Jul 1975, 2200Ј, A Newton, M Thayer (1, MCZC). DISCUSSION: This species has been collect- ed from mixed hardwood-conifer leaf litter and from under bark. Elevation records are from 400 to 2200 ft.

CHECKLIST OF AGATHIDIUM SPECIES: PART 1 Agathidium sexstriatum species group A. bistriatum Horn, 1880 A. sexstriatum Horn, 1880 A. estriatum Horn, 1880 Agathidium brevisternum species group A. brevisternum Fall, 1934 A. rhinocerellum Wheeler and Miller, new species Figs. 175±177. Agathidium falcatoperculum, A. dioperculum Wheeler and Miller, new species n.sp., aedeagus: 175, lateral; 176, ventral; 177, operculum. Agathidium revolvens species group A. revolvens LeConte, 1850 A. jasperanum Fall, 1934 transverse, with long ®ne setae (®g. 172). A. depressum Fall, 1934 Aedeagus with median lobe elongate, robust, A. dubitans Fall, 1934 A. dubitanoides Wheeler and Miller, new species ventrally curved, in lateral view slightly re- A. omissum Fall, 1934 curved dorsally subapically, apical 1/5 dor- A. cavisternum Fall, 1934 soventrally compressed and laterally nar- A. virile Fall, 1934 rowed before apex (®g. 176); in ventral view ϭ A. hebetatum Hatch, 1957, new synonym apex of median lobe narrowed, more-or-less A. conjunctum Brown, 1933 arrowhead-shaped, with fanlike arrangement ϭ A. obtusum Hatch, 1957, new synonym of ducts apically (®g. 176); operculum deep- A. angustoperculum Wheeler and Miller, new spe- ly bisected into pieces, each elongate, broad, cies with medial margin concave, curved, apex A. falcatoperculum Wheeler and Miller, new spe- pointed and directed medially (®g. 177); lat- cies eral lobes long, gradually narrowed before apex that is slightly expanded subapically in ACKNOWLEDGMENTS ventral view, with pair of subapical setae and This study has pro®ted from the advice, irregularly distributed pores (®gs. 175, 176); counsel, and encouragement of many col- endophallic armature not extensive. leagues, only some of which are singled out Female tarsi 5±4±4. for acknowledgment here. To those who have ETYMOLOGY: This species is named from contributed to the project in one or more the words falcatus, Latin for ``curved'', and ways and who are inadvertently omitted go 86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 290 thanks and apologies. The senior author frontispiece illustrations. Many individuals, thanks C.A. Triplehorn (The Ohio State Uni- particularly student assistants, have made im- versity), his advisor, mentor, and friend, for portant contributions to the database MY- taking an aspiring undergraduate coleopterist COL over the past decades. Last, but fore- on an expedition to Mexico that forged a most, F. Fawcett deserves considerable commitment to the study of beetles (includ- thanks for her superb scienti®c illustrations ing Leiodidae), systematics, and biological that combine artistic beauty with scienti®c diversity. L.E. Watrous accompanied him on precision and communicate as much or more several early collecting trips that were instru- than is possible through the text. Her contri- mental in elucidating the complex of ¯ight- butions to this project are incalculable in less species of the Appalachians, and was a quality and quantity. constant source of inspiration, sound advice, and stimulating theoretical arguments. K.C. REFERENCES Nixon (L. H. Bailey Hortorium, Cornell Uni- versity) collaborated in the development of Allen, A.A. 1954. Rhynchites cupreus L. (Col., the phylogenetic species concept used here, Attelabidae) etc., in West Sussex. Entomolo- is an eternal fountainhead of innovation and gists Monthly Magazine 90: 87. insights, and has made several versions of his Ammann, J., and H. Knabl. 1922. Die KaÈferfauna des nordwestlichen Tirol. Entomologische programs ClaDOS, Dada, MDP, and Win- Blaetter fuer Biologie und Systematik der Kae- Clada available. The following students, fer 18: 28±36, 49±64, 97±112, 145±160. friends, colleagues, wife, and children assist- Angelini, F. 1984. Reports of Agathidiini from ed on one or more collecting trips: B. Lind- Mongolia (Coleoptera, Leiodidae). Folia Ento- say, J.V. McHugh, J. Pakaluk, D. Peters, mologica Hungarica 45: 9±13. K.A. Wheeler, K.J. Wheeler, and M.A. Angelini, F. 1986. XXX contributo allo studio de- Wheeler. gli Anisotomini. Note sinonimiche (Coleoptera: Without the generous lending of material Leiodidae). Bollettino della SocietaÁ Entomolo- from collections listed in the appendix this gica Italiana 118: 147±160. work would have not been possible. Curators Angelini, F. 1988. Gli Anisotomini del Museo Civico di Storia Naturale di Milano (Coleop- enumerated therein are heartily thanked. tera, Leiodidae). Atti della Societa Italiana di Long-term loans by the Canadian National Scienze Naturali e del Museo Civico di Storia Collection, Field Museum of Natural Histo- Naturale in Milano 129: 305±366. ry, and Museum of Comparative Zoology Angelini, F. 1990. Anisotomini from Caucasus were especially valuable. (Coleoptera, Leiodidae). Annales Historico-Na- This research was supported by grants turales Musei Nationalis Hungarici 82: 91±122. from the U.S. National Science Foundation Angelini, F. 1993. Studi sugli Agathidium. Desig- (BSR-8315457, BSR-8717401), the High- nazione di un nuovo genere, un nuovo sotto- lands Biological Research Station Founda- genere e gruppi di specie (Coletopera Leiodi- tion, the American Philosophical Society, dae). Bollettino della SocietaÁ Entomologica It- aliana 125: 29±44. and the College of Agriculture and Life Sci- Angelini, F. 1995. Revisione tassonomica delle ences in Cornell University, especially Hatch specie paleartiche del genere Agathidium Pan- Project NY(C)139426 to the senior author. zer (Coleoptera: Leiodidae: Agathidiini). Mu- Two individuals who completed an incred- seo Regionale di Scienze Naturali Monogra®e ible amount of quality work, including dis- (Turin) 18: 1±485. sections of specimens, library research, spec- Angelini, F., and L. de Marzo. 1980. Utilita di imen measurement, working with databases, nuovi caratteri nella sistematica del genere editing materials, and such, deserve special Agathidium Panzer (Coleoptera, Leiodidae) e thanks: P.R. Fraissinet and J.V. McHugh. loro impiego nella designazione di due sinoni- McHugh also provided a very useful and mi. Entomologica (Bari) 16: 47±76. Angelini, F., and L. de Marzo. 1983. Anisotomini careful review of an earlier version of the nuovi o poco conosciuti reperti in Nepal e manuscript. D.A. Grimaldi, while a graduate Kashmir dal Prof. H. Franz (Coleopter, Leiod- student at Cornell, developed strategies for idae). Entomologica (Bari) 18: 5±16. measuring Agathidium. The late B. Alexan- Angelini, F., and L. de Marzo. 1984. Note siste- der, also while a grad student, completed the matiche sugli Agathidium dell'Africa centrale 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 87

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dium associated with an epimycetic slime mold Wheeler, Q.D., and N.I. Platnick. 2000a. A cri- plasmodium on Pleurotus fungi (Coleoptera: tique from the Wheeler and Platnick phyloge- LeiodidaeÐMyxomycetes: PhysaralesÐBasid- netic species concept perspective: Problems iomycetes: Tricholomataceae). Coleopterists with alternate concepts of species. In Q.D. Bulletin 41: 395±403. Wheeler and R. Meier (editors), Species con- Wheeler, Q.D. 1990a. Morphology and ontogeny cepts and phylogenetic theory: a debate: 133± of postembryonic larval Agathidium and Ani- 145.New York: Columbia University Press. sotoma (Coleoptera: Leiodidae). American Mu- Wheeler, Q.D., and N.I. Platnick. 2000b. The phy- seum Novitates 2986: 1±41. logenetic species concept (sensu Wheeler and Wheeler, Q.D. 1990b. Ontogeny and character Platnick). In Q.D. Wheeler and R. Meier (edi- phylogeny. Cladistics 6: 225±264. tors), Species concepts and phylogenetic theo- Wheeler, Q.D., and J.V. McHugh. 1987. Watrous ry: a debate: 55±69.New York: Columbia Uni- trays: A storage system for disarticulated and versity Press. Wiepken, C.F. 1886. Nachtrag zu dem Systema- dissected insects. Curator 30: 73±76. tischen Verzeichnisse der bis jetzt im Herzog- Wheeler, Q.D., and J.V. McHugh. 1994. A new thum. Oldenburg gefundenen KaÈferarten. Na- southern Appalachian species, Dasycerus bi- turwissenschaftlicher Verein zu Bremen 9: color (Coleoptera: Staphylinidae: Dasycerinae), 339±354. from declining endemic ®r forests. Coleopter- Williams, B.S. 1924. Coleoptera collected in the ists Bulletin 48: 265±271. Harpenden District. Entomologists Monthly Wheeler, Q.D., and R. Meier. 2000. Preface. In Magazine 60: 76±80. Q.D. Wheeler and R. Meier (editors), Species Woerndle, A. 1950. KaÈfer von Nordtirol. Inns- concepts and phylogenetic theory: a debate: ix± bruck: UniversitaÈtsverlag Wagner, 388 pp. xii.New York: Columbia University Press. Woodroffe, G.E. 1968. Some rare Coleoptera Wheeler, Q.D., and K.C. Nixon. 1990. Another from Wychwood Forest National Nature Re- way of looking at the species problem: a reply serve, Oxon., with records of six species of He- to de Queiroz and Donoghue. Cladistics 6: 77± miptera-Heteroptera new to Oxfordshire. Ento- 81. mologists Monthly Magazine 104: 22.

APPENDIX

The following is a list of collections from which CMNH Carnegie Museum of Natural History, specimens were borrowed for this project. Section of Invertebrate Zoology, 4400 Forbes Ave., Pittsburgh, PA 15213 (R. AMNH American Museum of Natural History, Davidson). Division of Invertebrate Zoology, Cen- CNCI Canadian National Collection of In- tral Park West at 79th St., New York, sects, Canadian Museum of Nature, NY 10024 (L. Herman). EntomologyÐCollections Division, ANSP Academy of Natural Sciences, Depart- P.O. Box 3443, Station ``D'', Ottawa, ment of Entomology, 1900 Benjamin ON, K1P 6P4, Canada (Bousquet, Y.). Franklin Parkway, Philadelphia, PA CUIC Cornell University Insect Collection, 19103 (D. Otte). BMNH The Natural History Museum, Crom- Department of Entomology, Comstock well Road, London, SW7 5BD, United Hall, Cornell University, Ithaca, NY Kingdom (P. Hammond). 14853 (J. Liebherr). CASC California Academy of Sciences, De- DENH Entomological Museum, Department of partment of Entomology, Golden Gate Zoology, University of New Hamp- Park, San Francisco, CA 94118 (D.H. shire, Nesmith Hall, Durham, NH Kavanaugh). 03824 (D.S. Chandler). CMNC Canadian Museum of Nature Collec- EMEC Essig Museum of Entomology, Univer- tion, 240 McLeod Street, Ottawa, ON sity of California, Berkeley, CA 94720 K1P 6P4, Canada (R.S. Anderson). (C. Barr). 2005 WHEELER AND MILLER: SLIME-MOLD BEETLES. PART I 95

FAIC F. Angelini Collection, SS7 Latiano, OSUC Museum of Biological Diversity, De- Km 0±500, I-72021 Francavilla Fon- partment of Entomology, Ohio State tana, Brindisi, Italy. University, 1315 Kinnear Road, Co- FGAC F.G. Andrews Collection, CDFA Plant lumbus, OH 43212 (N. Johnson). Pest Diagnostics Center, 3294 Mea- PECK Stewart B. Peck Collection, Carleton dowview Rd., Sacramento, CA 95832. University, Department of Biology, Ot- FMNH Field Museum of Natural History, De- tawa, ON, K1S 5B6, Canada. partment of Zoology, Roosevelt Road PSUC Frost Entomological Museum, Depart- at Lake Shore Dr., Chicago, IL 60605 ment of Entomology, Pennsylvania (A.F. Newton). State University, University Park, PA FSCA Florida State Collection of Arthropods, 16802 (K.C. Kim). Museum of Entomology, 1911 SW QDWC Quentin D. Wheeler Collection, De- 34th Street, Gainesville, FL 32608 partment of Entomology, Comstock (M.C. Thomas). Hall, Cornell University, Ithaca, NY JRAC J.R.A. Baranowski Collection, Univer- 14853. sity of Lund, Department of Systematic SEMC Snow Entomological Museum, Univer- Zoology, Helgonav, S-22362 Lund, sity of Kansas Natural History Muse- Sweden. um, University of Kansas, Lawrence, KSIC K. Stephan Collection, Route 1, Box KS 66045 (J.S. Ashe). 913, Red Oak, OK 74563. TAMU Texas A&M University Insect Collec- LACM Natural History Museum of Los An- tion, Department of Entomology, Texas geles County, Entomology Section, 900 A&M University, College Station, TX Exposition Boulevard, Los Angeles, 77843 (J.D. Oswald). CA 90007 (C. Bellamy). UASM E.H. Strickland Entomological Muse- MCMC Museo de Historia Natural de al Ciudad um, University of Alberta, Edmonton, de Mexico, Nuevo Bosque de Chapul- AB, Canada T6G 2M7 (F.A.H. Sper- tepec, Apartado Postal 18±845 Dele- ling). gacion Miguel Hidalgo 11800 Mexico, UGCA University of Georgia Museum of Nat- (G. Quinters) . ural History Collection of Arthropods, MCZC Museum of Comparative Zoology, Har- University of Georgia, Athens, GA vard University, Cambridge MA 02138 30602 (J.V. McHugh). (P. Perkins). UMMZ University of Michigan Museum of Zo- MNHN Entomologie, Museum National ology, Insect Division, University of d'Histoire Naturelle, 45 bis, Rue de Buf- Michigan, Ann Arbor, MI 48109 (B.M. fon, Paris, 75005, France (J.J. Menier). O'Connor). MTEC Montana Entomological Collection, USNM National Museum of Natural History, Department of Entomology, Montana Smithsonian Institution, Washington, State University, Bozeman, MT 59717 DC 20560 (D.G. Furth). (M.A. Ivie). WSUC M.T. James Entomological Collection, MZLU Museum of Zoology and Entomology, Washington State University, Depart- Lund University, HelgonavaÈgen 3 S- ment of Entomology, Pullman, WA 223 62 Lund, Sweden (R. Danielsson). 99164 (R.S Zack). NAUF The Colorado Plateau Museum of Ar- ZMHB Museum fuÈr Naturkunde der Hum- thopod Biodiversity, Northern Arizona boldt-UniversitaÈt, Humboldt-Universi- University, Flagstaff, AZ 86001 (N. taÈt zu Berlin, D-10099 Berlin, Germa- Cobb). ny (M. Uhlig). OSAC Oregon State University Entomological Museum, Department of Entomology, Oregon State University, Corvallis, OR 97331 (D.D. Judd).