Taxonomic review of the North American dung beetle genus Boreocanthon Halffter, 1958 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini)

Center for Systematic Entomology, Inc., Gainesville, FL Date of issue: October 14, 2022 0952 Taxonomic review of the North American dung beetle genus Boreocanthon Halffter, 1958 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini) W. D. Edmonds 2625 SW Brae Mar Ct. Portland, OR 97201 Insecta MundI A journal of world insect systematics Page Count: 65 Edmonds Edmonds WD. 2022. Taxonomic review of the North American dung beetle genus Boreocanthon Halffter, 1958 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini). Insecta Mundi 0952: 1–65. Published on October 14, 2022 by Center for Systematic Entomology, Inc. P.O. Box 141874 Gainesville, FL 32614-1874 USA http://centerforsystematicentomology.org/ Insecta Mundi is a journal primarily devoted to insect systematics, but articles can be published on any nonmarine arthropod. Topics considered for publication include systematics, taxonomy, nomenclature, checklists, faunal works, and natural history. 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This is an open access article distributed under the terms of the Creative Commons, Attribution Non-Commercial License, which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author(s) and source are credited. http://creativecommons.org/licenses/by-nc/3.0/ Insecta MundI0952: 1–65 2022 Taxonomic review of the North American dung beetle genus Boreocanthon Halffter, 1958 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini) W. D. Edmonds 2625 SW Brae Mar Ct. Portland, OR 97201 wdedmon[email protected]m Abstract. This paper presents a taxonomic review of the genus Boreocanthon Halffter, restored generic status, a group of ball-rolling (telocoprid) dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) endemic to North America. The genus, heretofore treated as a subgenus of Canthon, comprises 13 species, each keyed, diagnosed, illustrated and presented with information on distribution, relationships, and biology (*signifies restored generic combination): *Boreocanthon ateuchiceps (Bates, 1887); B. coahuilensis (Howden, 1966); *B. depressipennis (LeConte, 1859c); *B. ebenus (Say, 1823); *B. forreri (Bates, 1887); *B. integricollis (Schaeffer, 1915); *B. lecontei (Harold, 1868); *B. melanus (Robinson, 1948); *B. praticola (LeConte, 1868); *B. probus (Germar, 1823); *B. puncticollis (LeConte, 1866); *B. simplex (LeConte, 1857); as well as Boreocanthon halffteri Edmonds here described as a new species. Other actions taken are a) Canthon mixtus Robinson declared junior subjective new synonym of Boreocanthon puncticollis; b) Canthon bisignatus Balthasar, 1939, declared junior subjective new synonym of Boreocanthon simplex; c) Boreocanthon coahuilensis (Howden) new generic combination; and d) neotype designated for the type species of the genus Boreocanthon, Ateuchus ebenus Say, 1823, here Boreocanthon ebenus (Say). Key words. Canthon, new species, new synonymies, neotype, key to species. ZooBank registration. urn:lsid:zoobank.org:pub:74CC7BA2-4E8D-4780-BB1B-E47370CBB19D Introduction The genus Canthon Hoffmannsegg, 1817, is a large (~200 species), very diverse New World group with a long, somewhat complicated taxonomic history (Cupello and Vaz-de-Mello 2018). As defined by Halffter and Martínez (1977) and adopted by most workers since (e.g., Vaz-de-Mello et al. 2011), Canthon comprises nine subgenera (including Boreocanthon) and several residual species groups that lie outside current subgeneric limits. The species-level taxonomic picture of the deltochiline fauna of North America has remained relatively stable for the last 150 years or so, during which time it has been treated in numerous comprehensive works (LeConte 1859c, 1863; Harold 1868; Blanchard 1885; Bates 1887; Horn 1870; Leng 1920; Schmidt 1922; Balthasar 1939; Robinson 1948; Halffter 1958, 1961; Halffter and Martínez 1977). All of these works include species now assigned to Boreocanthon. In the mid-20th century, Canthon became the focus of intense taxonomic splitting as various workers sought to separate distinctive groups of species into new genera (Martínez 1948a,b; Pereira and Martínez 1956; Martínez and Pereira 1956; Halffter 1958). The emergent generic level reorganization was summarized by Vulcano and Pereira (1964) and Halffter and Martínez (1977). Halffter (1958) established Boreocanthon to accommodate a distinctive group of North American Canthon characterized primarily by their smaller size (4–10 mm), often strongly quadridentate clypeus (sometimes accompanied by dentate paraocular areas so as to produce a sexdentate head margin), and lack of a carinate line along the anterior margin of the metafemora. He initially moved ten species from Canthon to Boreocanthon: B. simplex (LeConte), B. melanus (Robinson), B. lecontei (Harold), B. probus (Germar), B. puncticollis (LeConte), B. praticola (LeConte), B. depressipennis (LeConte), B. ebenus (Say), B. integricollis (Schaeffer), and B. mixtus (Robinson) (here considered a synonym of B. puncticollis). Ateuchus ebenus Say was designated type species of the new genus, as Canthon ebenus (Say). He later (Halffter 1961) added Canthon ateuchiceps Bates and C. forreri Bates to the genus, considering them distinct enough to perhaps represent a new subgenus of Boreocanthon. Howden 2 · October 14, 2022 Edmonds (1966) added a new species to the group that he named Canthon coahuilensis, and assigned it with doubt to Boreocanthon, which he regarded as a subgenus (see comments for B. coahuilensis). Halffter and Martínez (1977) later confirmed membership of C. coahuilensis in Boreocanthon, which, in step with Howden (1966), they reduced in rank to subgenus; they also added to the group C. bisignatus Balthasar, here regarded as a synonym of B. simplex. With the addition here of Boreocanthon halffteri and the deletions resulting from two new synonymies (B. mixtus and B. bisignatus) the total number of valid species in the genus becomes thirteen. To the limited extent member species have figured in cladistic studies (Kohlmann and Halffter 1990; Medina et al. 2003; Halffter et al. 2022), there is evidence that the genus is a monophyletic group nested within the loose assemblage of North American representatives of the deltochiline Scarabaeinae. Kohlmann and Halffter (1990) is the most comprehensive of the three studies in the context of Boreocanthon (regarded a subgenus by them). In their study, the genus (minus B. coahuilensis) is clearly resolved as a monophyletic group closely related to Melanocanthon, the two taxa forming the sister group of B. coahuilensis. Within Boreocanthon there is little resolution: B. simplex is an outlier; B. depressipennis-ebenus-praticola resolve as a clade within a large polytomy uniting it with the remaining seven species. The cladistic analysis presented by Medina et al. (2003), while it clearly united Melanocanthon and Boreocanthon, includes too few species of the latter to provide further insight. The phylogenetic analysis by Halffter et al. (2022) placed the Boreocanthon – Melanocanthon duo as sister to Bajacanthon, a monobasic subgenus for Canthon obliquus Horn, endemic to the southern tip of Baja California Sur. Materials and Methods This study is based almost exclusively on museum specimens where species limits and definitions are morphology-based concepts whose practical purpose is to sort observable variation. While the basic unit of the resulting classification is the species, I cannot claim that those recognized here are “biological” in the sense that they represent anything other than a hypothesis of the current manifestation (morphological, molecular, behavioral, ecological, geographical and phylogenetic) of several putative historical lineages (in the present case, 13). The classification proposed here is conservative (see Zachos (2014) for context), provisional and subject to future changes as additional information becomes available and existing information is re-analyzed. It is but a snapshot of what one taxonomist now knows about a defined group of beetles named Boreocanthon. This review is based on examination of approximately 9550 specimens, all of which were graciously provided by the caretakers of the institutional and private collections cited in the Acknowledgements. Species and supraspecific diagnoses employ the terminology used by Edmonds (1972 and elsewhere) unless noted otherwise. In addition, presented below is a summary of morphological variation observed in Boreocanthon focused primarily on its taxonomic context as well as notations on the use of certain terms. Specified body lengths include the head; these measurements are approximate and intended only for comparing relative size. Collection localities were gleaned mainly from specimens seen by me. Localities are arranged alphabetically by country followed by first-order political subdivisions (state, department, province or equivalent) and secondorder divisions (county, municipality, etc.). Individual collection points follow their second-order division, each preceded by a solid black dot (● or, in the case of certain B. simplex, a red diamond ♦) and followed in brackets by month(s) of collection, if specified. The symbols are used to clearly punctuate locality lists, otherwise visually difficult to separate using more conventional marks. If a label cites only a U.S. county location, the name of the county is entered followed by the notation “(no data).” In order to minimize errors, no attempt is made to convert distance and elevation, which are presented as transcribed (miles or kilometers, or, feet or meters). Similarly, I have not standardized the format of geographical coordinate data; these are presented as they appear on labels, but each was checked for consistency with accompanying information (name of town, or distance from point, elevation, etc.) using Google Earth. The geographical distribution maps (base map modified from original source: https://www.gifex.com/fullsize-en/2009-11-08-10949/North_America_Political_Outline_Map.html) represent my current inferences about species geographic distributions based on the collection data presented in the species treatments; they are not intended to be static representations of range. On the contrary, the margins of the shaded areas should be regarded as tenuous, but I do regard them as reliable enough for supporting generalizations and comparisons about geographical distributions. In a very general way, ecogeographic characteristics of Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 3 each species are indicated by reference to the vegetational mapping in Figure 2, and in the section “Ecogeographic environment” at the end of each species diagnosis. This section consists of a designated biome followed parenthetically, when appropriate, by ecoregion(s) inhabited by a given species. Figure 2 is modified from Ecoregions 2017© Resolve (https://ecoregions.appspot.com/), an interactive website where the limits of specified ecoregions (subunits of biomes) for each species, as well as other biogeographical information can be viewed (Olson et al. 2001; Dinerstein et al 2017). Figures 1–2. 1) Approximate corporate distribution of the genus Boreocanthon. 2) Generalized depiction of vegetation biomes of the United States (Biome key in box; See Materials and Methods for explanation). 4 · October 14, 2022 Edmonds A weakness of this study is the fact that I was unable to examine directly type material. In most cases, however, photographs of putative type material was available, and these have been critical to achieving an acceptable degree of nomenclatorial confidence. For each species treatment, the section “Type Material” will provide information I have about the species-group name(s) in question. Location of type material, where known, is indicated by name of museum/collection and city. In using the term “holotype” I have been mindful of ICZN Recommendation 73F, “Avoidance of assumption of holotype”. Consequently, I have reserved reference to “holotype” only to cases I regard as highly reliable and have used “syntype” in other cases. To my knowledge, no lectotypes have been designated for species in the genus, and I hesitate to do so here because I have not been able to examine available type material directly. Type localities are determined in accordance with ICZN Article 76. In all cases, I rely upon information provided in the original description of the taxon corroborated by label data accompanying specimens and reliable information from published sources. Morphology of Boreocanthon. The following summary is offered as an aid to understanding and assessing the taxonomic features cited in the key and various diagnoses for Boreocanthon. It is neither exhaustive nor meant to apply directly to other groups, although none of the morphology described here is unique to the genus. Color: Viewed from reading distance, almost all Boreocanthon appear dull and sombre-colored, usually black or nearly so; B. forreri (Fig. 102–103) while dark, presents a subdued, metallic green or coppery green dorsum, and B. ateuchiceps (Fig. 38–39) is bright, shiny green or yellow green. Many of the “black” species at least occasionally show dull green or blue highlights on the head and pronotum revealed under magnification (×5) under broad spectrum light. Boreocanthon depressipennis (Fig. 11) almost always presents conspicuous green highlights. Such highlights, while usually much subdued on casual inspection, are sometimes much enhanced on specimens submerged in liquid (for example, blue in B. puncticollis and usually green in B. melanus). Pigmented color (as opposed to the structural colors just mentioned) is found in but one species, B. simplex, certain populations of which include individuals with orange/red-orange elytral umbones (Fig. 96–97). Mario Cupello (pers. comm.) reports that individuals with similar orange umbones occur as intraspecies variants in several distantly related South American dung beetles, including Ateuchus globulus (Boucomont 1928) and Canthidium bituberculatus (Fabricius 1801). Size: Boreocanthon are generally small, robust dung beetles with lengths often below 6 mm. The lengths of all species vary by at least 50%, sometimes by almost 100%. The average length of the smaller species ranges from about 4–6 mm, that of the larger species (B. ebenus, B. depressipennis, B. praticola and B. coahuilensis) from about 5–8 mm, with exceptional individuals reaching 10 mm. Sculpturing: Dorsal (head+pronotum+elytra+pygidium) sculpturing is highly variable in Boreocanthon and is a primary consideration in diagnosing species. Terminology applied to sculpturing of the integument has a complicated, sometimes inconsistent lexicon (see Harris 1979); that adopted below is for convenience and clarity in the context of this review. Much of the sculpturing is subtle and highly subject to distortion caused by the wear and tear exacted by a fossorial lifestyle. Assessment of sculpturing for taxonomic purposes, therefore, is best based on “fresh” specimens. Sculpturing in the genus consists of a suite of four features best assessed under moderate (>10× magnification). 1) Shagreen (primary granulation; microgranulation) (Fig. 13, 48, 71, 75)—The upper surface of all Boreocanthon species is more or less evenly covered by very small, densely packed (contiguous), rounded granules giving the magnified (×25) surface an evenly roughened, or “alutaceous” or shagreen texture resembling that of untanned leather, cobblestone or calade ground cover. The shagreen commonly also extends to the ventral surfaces. This sculpture feature is almost always distinct and coexistent as a background to other surface features (punctures, etc.). In B. ateuchiceps (Fig. 38) and less so in B. forreri (Fig.105), the shagreen is largely effaced to produce a polished, shiny surface; in B. lecontei (Fig. 31), it is largely obscured on the pronotum by the very fine, densely compacted secondary granulation characteristic of this species. 2) Granulation (secondary granulation; macrogranulation)—In three species, B. ebenus (Fig. 27), B. depressipennis (Fig. 11, 13), and B. praticola (Fig.8), the upper and some ventral surfaces are uniformly covered by a field of coarse, rounded, elevated granules. These are usually discrete (not contiguous) but regularly coalesce on the pronotum of B. ebenus. A fourth species, B. lecontei (Fig. 31) is unique in that the head and pronotum are covered Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 5 by a fine, dense, secondary granulation that largely obscures the shagreen background; unlike the preceding three species, the granulation does not extend to the elytral interstriae. Macrogranulation is only very rarely accompanied by distinct puncturing, and then only in limited areas (such as the pygidium). The other North American groups of deltochilines, Melanocanthon and the Canthon pilularius species group (sensu Halffter 1961, Halffter and Martínez 1977), share the macrogranulate condition (see Medina et al. 2003). 3) Microspots (maculae)—In all species lacking distinct macrogranulation, the dorsum presents a field of small, shiny spots, or maculae, usually accompanied by distinct puncturing (Fig. 61, 88, 115). The size and distribution of microspots strongly suggest that they are vestigial granules, with which they never coexist, and that a granulated surface is the likely plesiomorphic condition of Boreocanthon. Circumstantial evidence of plesiomorphy resides in a single, unusual specimen of B. simplex observed for this study that conserves limited elytral granulation accompanied by maculae (see Comments for B. simplex). 4) Puncturing—As for microspots, the upper surface of species lacking secondary granulation almost always presents puncturing. Puncturing is generally weak, very fine and usually sparse and difficult to assess without at least low (5×) magnification (Fig. 59, 75, 109). Puncturing usually extends to ventral surfaces and is usually absent from the elytral interstriae. Punctures on the pygidium as well as elsewhere over the dorsal surface sometimes bear minute setae visible only at higher magnification (50×) from an oblique angle. While never dense and always simple, pronotal puncturing varies mainly in the degree of conspicuousness, or “crispness,” especially on the pronotum, from clearly expressed (B. puncticollis, Fig. 61) to very weak and nearly effaced (B. coahuilensis [Fig. 48] as well as some B. probus [Fig. 75] and B. halffteri). Head: A conspicuously dentate outer head margin is a hallmark feature of Boreocanthon. The clypeus bears four, usually prominent, upturned teeth, of which the median pair is always larger than the lateral pair. The head margin behind the lateral teeth (at the junction of the fronto-clypeal suture) is almost always interrupted by an indentation of variable size and shape, here called the paraocular notch, that sets off the anterior angle of the paraocular area (gena auct.). The size of the paraocular notch and corresponding shape of the paraocular angle provide important diagnostic characters. If the notch is small and inconspicuous (as in B. simplex [Fig. 92] and B. lecontei [Fig 32], for example), setting off only a small angular portion of the paraocular area, the head will appear decidedly quadridentate or only weakly sexdentate. If the Uor V-shaped notch is large, setting off a strongly developed, acute angle (as in B. probus [Fig. 74] and B. ebenus [Fig. 11], for example), the head will appear decidedly sexdentate. Some species (B. melanus [Fig. 112] and B. simplex [Fig. 92], for example) present intermediate shapes. In B. coahuilensis (Fig. 52) certain individuals have very poorly developed lateral clypeal teeth and obsolete paraocular notches such that the head is virtually bidentate, a condition that prompted Howden (1966) to question its placement in Boreocanthon. The dentition of the head (as well as protibiae) is highly prone to distortion/obliteration resulting from wear produced by digging in the rocky, sandy soils common to the habitats of these beetles, which, at beetle scale (Fig. 78), present formidable obstacles to digging. Figure 62 compares the clypeal margin in “fresh” and highly abraded individuals of B. puncticollis. Reliance on well-worn specimens for taxonomic assessment can, unsurprisingly, lead to avoidable errors (e.g., the case of Canthon bisignatus Balthasar; see B. simplex.) Sculpturing of the upper surface of the head, especially in non-granulate species, consists of two zones (Fig. 47, 55, 92): Immediately following the marginal teeth, the surface is usually roughened to some degree for about one-half the length of the clypeus and thereafter that of the remainder of the clypeus, the frons and the paraocular areas approximates the surface of the pronotum. The reflectivity of the head surface is usually uniform but occasionally somewhat brighter anteriorly. In B. probus (Fig. 79), the clypeus often presents a conspicuous shiny, transverse band, a condition often also seen in B. integricollis. The labio-gular fimbria marks the putative boundary between the mentum and gular sclerites. It consists of an array of long, recumbent setae of variable shape mixed with median field of coarse punctures. The fimbria may consist of a simple, posteriorly bowed row of setae (Fig. 69), or, more commonly, display a posteriorly directed, narrowed, V-shaped configuration (Fig. 7, 15, 60) extending up to one-half the length of the gula. In either case, it often includes a group of coarse punctures arranged along the gular midline. Prothorax: Sculpturing of the pronotum is usually evenly expressed over its entire surface. Among Boreocanthon species, there are two principal patterns: granules on a shagreen background (Fig. 13), and punctures mixed with microspots on a shagreen background (Fig. 48, 75). As the clarity of puncturing varies from hardly visible to bold, so, too, does the expression of microspots. (See puncturing, above) 6 · October 14, 2022 Edmonds The anterolateral angles of the pronotum are usually little, if at all, upturned (explanate). In B. probus (Fig. 73) they can be narrowly elevated, especially in western populations, while in the closely related B. halffteri (Fig. 87) they are broadly elevated between the lateral and anterior angles of the pronotal margin. The curving of the dorsal profile of the pronotum (viewed laterally, Fig. 53) usually reaches only slightly above the level of the elytral surface. In B. coahuilensis, as well as B. ebenus, the curvature is much stronger, producing a humpbacked profile. At times the underside of the pronotal margin, near the anterior end, bears a small, rounded tubercle or small group of tubercles that has been regarded as diagnostic by Robinson (1948) for certain species (B. melanus, B. lecontei, B. mixtus [=puncticollis]). This feature is variable in these species, to the extent that I have not regarded it as diagnostic. Only in B. coahuilensis does the presence of “Robinson’s tubercle” appear to be a reliable species character. The posterior margin of the procoxal cavity is usually thickened only to a point near its lateral limit (Fig. 24). In some Boreocanthon (B. ateuchiceps [Fig. 40], B. forreri [Fig. 108], B. melanus, B. praticola and B. simplex) it “escapes” the coxal margin as a fine carina making a broad turn laterally to a point approximately one-fifth to onefourth of the distance to the lateral pronotal margin. I here refer to this line as the hypomeral carina, which, in the case of these species, is incomplete and, as pointed out by Blanchard (1885), sometimes variable in expression within a single species, such as B. simplex. In some North American Canthon (e.g., C. humectus and C. viridis), it is complete, continuing uninterrupted to the lateral margin. Similar hypomeral carinae (complete and incomplete) are found in numerous other scarabaeines. Pterothorax: The mesoventrite (mesosternum of Edmonds (1972)) is very short and broad and usually smooth, or with any puncturing confined to its lateral extremities (Fig. 12). In some Boreocanthon, notably B. integricollis (Fig.70), it is uniformly punctured. The pleura and metaventrite (metasternum of Edmonds (1972)) usually present sculpturing similar to the elytra (except for microspotting), although the metaventrite is often smooth, shiny and sparsely, but sharply punctate (Fig. 70). Legs: The outer protibial margin is strongly tridentate in all Boreocanthon, and the margin between the teeth and along its basal one-half are conspicuously serrate. The inner margin varies from gently curved from base to apex (Fig. 104) to strongly offset (stepped) near its midpoint (Fig. 16, 51), with some species showing an intermediate shape. In the granulate species (B. ebenus, B. depressipennis, and B. praticola), the anterior end of the inner margin is prolonged as a conical tooth (Fig. 16, arrow; weakest in B. praticola). In most Boreocanthon, the protibial spurs are sexually dimorphic: strongly curved laterad and tapering to an acute point in the female (Fig. 116a); curved laterad, apically slightly expanded and asymmetrically, acutely bidentate in the male (Fig. 116b); in two species (B. ebenus and B. depressipennis) this spur is curved inward and tapering to an acute point in both sexes. Boreocanthon lack any trace of the fine ridge along the anterior face of the metafemora common in other North American deltochilines. All leg features mentioned above are highly susceptible to abrasive wear distorting or obliterating the pristine condition. Elytra: Elytral sculpture is always similar to that of the pronotum except for distinct puncturing, which can extend to the humeral angle (Fig. 71). Interstriae 1–6 are generally flat (For convenience, the space between the elytral suture and first stria is here considered the first interstria). The anterior ends of the first (sutural) interstriae are usually depressed to some degree. In a few species (most notably B. puncticollis, Fig. 59), the anterior ends of interstriae 2 and 3 are conspicuously raised and tumid. Not uncommonly, the anterior edges of various interstriae (most commonly the 3rd and 5th, Fig. 8) each bear a minute, round, often acute tubercle, sometimes obscured by the posterior margin of the pronotum; but the variability of these tubercles renders them only minimally useful taxonomically. Boreocanthon possess nine elytral striae. The discal striae (1–6) are superficial, not perceptibly impressed, and usually extend to the elytral apex; the lateral striae (7–9) vary in definition. In a few species (especially B. puncticollis, Fig. 59, 61), the strial margins are carinulate and the striae thus sharply marked; in most, the strial margins are not sharply defined and the striae appear somewhat effaced (“fuzzier”). The 7th (humeral) stria is effaced basally where it would cross the umbone, and extends from the umbone to the elytral apex. In several species the 8th (subhumeral) stria can be marked by a fine carina extending from near the base to a point mid-way or beyond the length of the elytron (Fig. 81, 88, arrow). The 9th (epipleural) stria, if not effaced, is expressed only posteriorly in the narrow space between the 8th stria and upper edge of the elytral epipleuron. Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 7 Abdomen: The pygidium in Boreocanthon is completely margined and always separated from the propygidium by a transverse ridge; its apex is commonly raised and more convex than the rest of the surface (Fig. 6). In two species, B. ebenus (Fig. 27) and B. depressipennis (Fig. 14), the basal area of the pygidium is strongly impressed (concave) on either side of the midline. Sculpturing of the pygidium mimics that of the pronotum, although it is usually more weakly expressed; punctures, if present, often bear very short, minute setae visible only at a steep angle under higher magnification (25×). The last (6th) ventrite (visible abdominal sternum of Edmonds (1972)) is usually sculptured like the pronotum; its posterior margin is broadly, weakly emarginate in the male and more-or-less straight in the female, although the difference is often subtle and difficult to assess. Ventrites 1–5 always exhibit weaker sculpturing than the 6th, especially medially. Male genitalia: I here use the term genital capsule (sensu Edmonds (1972)) to refer to the sclerotized, tubular structure enclosing the penis/endophallus and consisting of a tubular base and pair of apical appendages, the parameres; it is very often referred to (including by me) as the aedeagus (but see Lawrence et al. 2010). In all Boreocanthon, the parameres are laterally flattened, obliquely truncated and closely appressed apically; their lateral profile and lower angles of the apical margin (viewed caudally) can assume species-characteristic shapes (cf. Fig. 50, Fig. 56–57 and Fig. 77). The form of the parameres in Boreocanthon is shared by Melanocanthon (Robinson 1948) and Bajacanthon (Halffter et al. 2022); that of the pilularius group of Canthon is quite distinctive, the lower apical angle being extended as a finger-like appendage (Fig. 43). The endophallus and its components were not examined for this study. Female genitalia: The paraprocts (genital plates of Martínez-Morales and Morón (2015); see also Cristóvão and Vaz-de-Mello (2020)) of the female are elongate, apically rounded and slightly twisted appendages arising below the genital opening. Their cupped apical margins (Fig. 80, arrow a) appear to form a curved shelf that combines with the semicircular emargination of the marginal flange of the pygidium (Fig. 80, arrow, and Fig. 21, asterisk), presumably forming a guide for engaging the parameres. The paraprocts of Boreocanthon were not examined systematically for this study, but they appear to vary in ways that could later prove to be taxonomically useful. Systematics Genus Boreocanthon Halffter, 1958, restored generic status Boreocanthon Halffter 1958: 208 (as genus) Canthon (Boreocanthon Halffter) (downgraded to subgenus of Canthon Hoffmannsegg per Halffter and Martínez 1977: 81) Type species. Ateuchus ebenus Say, 1823, by original designation (as Canthon ebenus [Say]). Summary diagnosis. North American dung beetles, ball-rolling species attracted to a wide range of animal feces; usually dark or black in color, small-sized (length usually 4–7 mm); head margin conspicuously quadrior sexdentate; parameres compressed laterally, truncate apically, lacking elongate projection of lower apical angle; hind femora lacking fine carina along anterior edge; metatibia with single apical spur. Diagnosis. Dorsum (head + pronotum + elytra + pygidium) shagreened (alutaceous), with either punctures (Fig. 38, 48, 75, 105) or raised granules (Fig. 8. 19), punctures often accompanied by field of faintly lustrous microspotting (Fig. 61, 88); pterothoracic and abdominal sclerites shagreened, usually with accompanying flattened granules or punctures. Head: Clypeus quadridentate (Fig. 20, 66, 79, 112) (sometimes virtually bidentate in B. coahuilensis, Fig. 47); paraocular areas often also angulate such that head appears sexdentate (Fig. 66, 79). Clypeal teeth reflexed, surface behind concave, finely rugose and often somewhat shinier than rest of head surface, which is usually punctate on shagreened background. Clypeal process a low, transverse ridge. Labiogular fimbria always setose, often V-shaped, usually with associated field of coarse punctures at midline (Fig. 60, 69). Prothorax: Subapical tubercle beneath anterior angle of pronotum usually absent. Hypomeral carina usually absent (Fig. 24), sometimes partially developed but never long, extending at most only one-fifth to one-quarter of distance to edge of prothorax (Fig. 40, 108, arrow). Legs: Protibia strongly tridentate, usually serrate along 14 · October 14, 2022 Edmonds Figures 3–9. Boreocanthon praticola (LeConte). 3) Dorsal habitus. 4) Dorsal view of head. 5) Genital capsule, a: lateral view, b: apical view. 6) Posterior view of pygidium. 7) Ventral view of mentum and gula. 8) Dorsal view of union of pronotum and elytra. 9) Oblique ventral view of mesoand metaventrites. Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 15 Collection localities. CANADA—ALBERTA: Cypress County (Mpal. Dist.) ●Medicine Hat [Apr, Aug] ●Cypress Hills [Jun]; Wheatland County (Mpal. Dist.) ●Cluny [Jun]; Ranchland Mpal. Dist. ●36 km SW Longview, 50.264° -114.264°, 1400 m [May]. MANITOBA: Mpal. Two Borders ●Lyleton. MEXICO—CHIHUAHUA: Mpio. Cuauhtémoc ●11 mi W Cuauhtémoc [Oct]; Mpio. Huejotitán ●Valle de Olivos, 5500 ft [Jul]; Mpio. Janos ●20 km W Janos, 30°50ʹ33.8ʺ N 108°23ʹ25.1ʺ W [Oct] ●80 mi NNW Casas Grandes [Aug]; ●45 mi NW Casas Grandes [Aug]; Mpio. Juarez ●Samalayuca [Jun] ●Samalayuca, Laguna de Riataso [Aug]; Mpio. López ●Salaices [Jul]; Mpio. Matamoros ●Mariano Matamoros, 6000 ft [Aug]; Mpio. Metachí ●Metachí [Aug]. DURANGO: Mpio. Diez de Octubre ●San Lucas, 6700 ft [Aug]; Mpio. Lerdo ●Cd. Lerdo; Mpio. Ocampo ●173 mi N Durango [Aug]. SONORA: Mpio. Alamos ●Alamos [Jul] ●San Bernardo [Jul]; Mpio. Cananea ●Cananea, 1525 m [Aug]; Mpio. Hermosillo ●Hermosillo [Jul]; Mpio. Naco ●Naco, 5000 ft [Aug]. UNITED STATES—ARIZONA: Apache Co. ●16–19 mi N St. Johns [Sep] ●Lyman Lake [Aug] ●6 mi N St. Johns, 34.5858° N 109.3288° W, 1720 m [Aug]; Cochise Co. ●2 mi SE Apache, 4400 ft [Aug] ●8.5 mi S Apache [Aug] ●Douglas [Jun-Sep] ●12 mi NE Douglas [Aug] ●1 mi E Douglas [Aug] ●McNeal [Sep] ●Sycamore Canyon, Peloncillo Mountains [Jul] ●8 mi E Portal [Jul, Sep] ●1 mi NE Portal [Sep] ●5 mi SW Portal, Southwest Research Station [Aug] ●Guadalupe Canyon [Jul] ●2.3 mi WNW Chiricahua [Jul]; ●22.5 and 36 mi SE Willcox [Jul]; ●5.8 km SE Willcox, 32°13ʹ10ʺ N 109°46ʹ51ʺ W [Aug] ●Willcox Playa [Jul-Aug] ●Chiricahua Mountains, mouth of Pinery Canyon, 5300 ft [Jun-Aug] ●Palominas [Aug] ●Texas Canyon [Aug] ●Dos Cabezas [Aug] ●Kansas Settlement [Jul]; ●Hereford [Jul] ●Huachuca ●Skeleton Canyon [Aug] ●10 mi NW Bisbee [Jul] ●Rustler Park [Aug]; Coconino Co. ●4.5 mi E Moenkopi [Jun] ●Tuba City [Aug] ●5 mi NE Cow Springs [Jul] ●Williams [Aug]; Gila Co. ●Pine ●Globe [Aug]; Graham Co. ●65 mi N Willcox, Deer Creek Ranch, Galiuro Mts. [Jul]; Greenlee Co. ●16 mi S Clifton [Aug] ●Franklin [Aug]; Maricopa Co. ●Webb [Jul] ●El Dorado [Jun]; Pima Co. ●Tucson [May, Aug] ●Boboquivari Mt. [Jul]; Santa Cruz Co. ●Patagonia [Aug-Sep] ●Sonoita [Jul] ●4 mi S Sonoita [Jul]; ●12 mi NE Sonoita [Jul, Sep]; ●10 mi SE Patagonia; ●3 and10 mi E Patagonia [Aug]; ●Santa Cruz River, 2.5 mi E Lochiel, 4500 ft [Sep]; ●Madera Canyon ●Coronado National Forest, Parker Canyon at FM 61, 4800 ft [Sep] ●Elgin [Sep]; Yavapai Co. ●17 mi W Jerome [Aug]; ●Seligman [Aug] ●Prescott Valley [Jul-Sep]; ●2 mi W Dewey [Oct] ●5 mi N Dewey [May] ● Hwy I-40, exit 103, Jolly Rd., 35°15.412ʹ N 113°10.090ʹ W, 5300 ft [May] ●Mayer [Aug]. COLORADO: Conejos Co. ●Antonito [Jun]; Denver Co. ●Denver [Jul]; El Paso Co. ●Colorado Springs [Jun, Oct]; Huerfano Co. ●Gardner [Jun]; Jefferson Co. ●Leyden [Aug]; La Plata Co. ●Durango [Jul]; Larimer Co. ●Ft. Collins [May]; Mesa Co. ●Mesa [Jul] ●10 mi NW Mack [Apr]; Moffat Co. ●Craig, 6200 ft [Jun]; Morgan Co. ●Hwy 52, 6.3 mi N Fort Morgan [May]; Prowers Co. ●Lamar [Aug]; Weld Co. ●Greeley. KANSAS: Barber Co. ●10.5 km W Hardtner, 37.02047 -98.76700 [Sep] ●2mi W Hardtner [Nov]; Clark Co. ●(no data), 1962 ft [Jun]; Gove Co. (no data); Gray Co. (no Data); Hamilton Co. ●Syracuse [Jun]; Kiowa Co. ●10 mi NW Mullinville [Jun] ●13 mi S Greensburg [Jan-Feb, Apr, Jul]; Logan Co. (no data); Meade Co. ●Meade [Jul]; Morton Co. (no data); Norton Co. ●Prairie Dog State Park [May]; Reno Co. ●Medora [May]; Republic Co. ●Kakley [Jul]; Sheridan Co. (no data) Thomas Co. Colby [May]; Wallace Co. (no data) [Jul]. MONTANA: Beaverhead Co. ●Dillon [Apr-May] ●Bond [Aug] ●P&O Ranch [Aug] ●Blacktail Creek [Aug]; Big Horn Co. ●Lodge Grass [Jun]; Blaine Co. ●Chinook [Jul]; Cascade Co. ●Great Falls [May]; Chouteau Co. ●15 mi S Loma [Jun]; Custer Co. ●4 mi S Miles City [Aug]; Fergus Co. ●Suffolk [Jun]; Gallatin Co. ●Bozeman [May-Jul] ●20 mi N Bozeman [Jun] ●Three Forks [Jul]; Glacier Co. ●Glacier National Park [Jul]; Jefferson Co. (no data) [Mar]; Judith Basin Co. ●Hobson [Aug]; Madison Co. (no data) [Jun]; Petroleum Co. 3 mi NW Winnett [Sep]; Teton Co. ●Bynum [Jul]; Yellowstone Co. ●Broadview [Jul]. NEBRASKA: Buffalo Co. ●10 mi W Amherst, 40.859° N 99.380° W, 715 m Chase Co. ●Imperial [Jul]; Cheyenne Co. ●3mi SE Lodgepole [Aug]; Dundy Co. ●Sanborn, 5 mi W Haigler, 40.06°W 102.02° W, 1010 m ●4 mi E Haigler, 40°01ʹ25ʺ N 101°51ʹ36ʺ W, 285 m [Aug]; Frontier Co. ●Red Willow Reservoir State Wildlife Management Area, 40.40° N 100.68° W, 800 m [May]; Greeley Co. ●7.3 mi S Greeley [Apr]; Keith Co. ●Lake McConaughy [May, Jul, Sep] ●5mi S, 2 mi W Paxton [Jul]; Morrill Co. ●8 mi W Bridgeport, 41.68° N 103.25° W, 1130 m [Jul]; 2.7 mi W Bridgeport [May]; Red Willow Co. ●McCook [May]; Sioux Co. ●42°54ʹ59.88ʺ N 103°45ʹ35.82ʺ W, 1125 m Thomas Co. ●Nebraska National Forest, Whitetail Campground, 41.79413 -100.27381 [Sep]. NEW MEXICO: Bernalillo Co. ●Albuquerque [Aug]; Catron Co. ●Luna; Chaves Co. ●Bottomless Lakes State Park [Jun]; Colfax Co. ●Maxwell [Jul] ●Koehler [Jul]; Doña Ana Co. ●Jornada Range [Jul] ●Isaack Lake [Jul]; Hidalgo Co. ●Lordsburg [Jul] ●40 mi S Lordsburg [Jul] ●1.5 mi NE Rodeo [Jul]; ●39 mi SW Animas [Sep] ●Rodeo, 4000 ft [Jul-Aug] 16 · October 14, 2022 Edmonds ●34 mi S, 3 mi W Animas [Sep] ●8 mi S Animas [Jul] ●32 mi S Animas, Double Adobe Ranch, 5500 ft [Jul-Aug]; Luna Co. ●7.5 mi W Akela Flats [Jul] ●Deming [Aug]; Roosevelt Co. ●8 mi NE Portales, ENMU Natural History Preserve [Apr, Jul]; Sandoval Co. ●La Ventana, Jemez Mountains, 7200 ft [Jul]; San Juan Co. ●Aztec; Santa Fe Co. ●Santa Fe [May, Jul]; Socorro Co. ●20 mi E Bernardo, 34°02.4ʹ N 106° 36.49ʹ W, 1607 m [Sep] ●Sevilleta National Wildlife Reserve [Aug]; Union Co. ●8 mi N, 2.5 mi W Sofia (Kimble Ranch) [Sep]●10 mi S, 0.5 mi E Sofia (Britt Ranch) [Sep]. NORTH DAKOTA: Billings Co. ●South Unit, Theodore Roosevelt National Park [Apr]; Morton Co. (no data); Richland Co. (no data). OKLAHOMA: Comanche Co. ●Wichita Refuge [Aug]; Noble Co. ●(no data); Payne Co. ●(no data) [Aug]. SOUTH DAKOTA: Custer Co. ●Custer State Park [Jun]; Jackson Co. ●vicinity Figure 10. Boreocanthon praticola (LeConte). Upper diagrams depict consolidation of ranges of the prairie dog species, Cynomys gunnisoni and C. ludovicianus. Inset depicts approximate distribution of B. praticola. Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 17 Badlands National Park [Jun]; Pennington Co. ●Hill City [Sep]; Perkins Co. ●Grand River National Grasslands [Aug]. TEXAS: Brewster Co. ●Alpine [Jun] ●7 mi NW Alpine [Jun] ●Marathon [Jun]; Callahan Co. ●Baird [Jun]; Culberson Co. (no data); Dallam Co. ●Perico [Aug]; Dallas Co. ●Dallas; Dawson Co. ●Lamesa [May]; Jeff Davis Co. ●Ft. Davis [Jul-Aug] ●Valentine [Jun]; ●8 mi SE Valentine [Jul] ●16 km S Fort Davis (along TX 17), 30°27ʹ48ʺN 103°58ʹ59ʺW, 1600 m (Aug); ● ~16 km NE Valentine, Muerto Springs Ranch (Muerto Springs), 30°40ʹ50ʺN 104°20ʹ22ʺW, 1555 m (Jul); ● ~16 km NE Valentine, Muerto Springs Ranch (Muerto Springs, near headquarters), 30°40ʹ28ʺN 104°24ʹ07ʺW, 1475 m (Jul); Lubbock Co. ●Lubbock [Aug] ●Ransom Canyon [Sep]; Presidio Co. ●Marfa [Jun] ●37 km SSW Marfa (along FM 2810, Petan Ranch – Cherry Hills sector), 30°07ʹ35ʺN 104°19ʹ24ʺW, 1630 m (Jun); ●20–26 km SSE Marfa (along FM 169), 1355–1415 m (Jun); ●8 km W La Viuda Peak, 29°42ʹ30ʺN 103°54ʹ30ʺ, 1200 m (Sep) ● ~16 km W Valentine (Miller Ranch, near headquarters), 30°33°30ʺN 104°38ʹ44ʺW, 1350 m (Jul–Aug); ● ~6.5 km W Marfa (Hip-O Ranch), 30°21ʹ54ʺN 104°07ʹ12ʺW, 1530 m (Aug– Sep) ●4 mi SE Presidio [Jul]; Reeves Co. ●Toyah [Jun]; Wichita Co. ●Foster Ranch [Aug]; Wilbarger Co. ●Vernon [Aug]. UTAH: Duchesne Co. ●Monument Butte, near Myton [Jul]; San Juan Co. ●Blanding [Jul] ●9.7 mi ENE Monticello, 2063 m [May]. WYOMING: Albany Co. ●Laramie [Jun]; Crook Co. ●4 mi N Sundance [Jun] ●Devil’s Tower National Monument [Jun]; Laramie Co. ●Cheyenne [Jul]; Sheridan Co. ●25mi E Sheridan [Jun]. Comments. Boreocanthon praticola is a very common, widespread species often confused with B. ebenus, from which it is easily distinguished by its convex (not impressed basally) pygidium and sexually dimorphic protibial spurs. The two are widely sympatric where cool (elevations >1200 m south, >800 m north) grasslands are punctuated by enclaves of vegetated sand habitat. Because it conserves a macrogranulated sculpture as well as dimorphic protibial spurs, I regard it a taxonomic isolate and annectant between B. ebenus-B. depressipennis (granulate dorsum) and rest of genus (sexually dimorphic protibial spurs). Boreocanthon praticola is subject to a considerable amount of variation, especially in size and surface texture — smaller individuals are routinely more finely granulose and require close inspection (10×). Populations from northern portions of its range (Nebraska north and northwestward) tend to be smaller and with finer dorsal granulation. I have examined specimens labeled Brazoria Co., Texas, and Benton Co, Arkansas; I consider them records that require corroboration. Pierce’s (1946) fossil taxon, Canthon praticola vetustus (along with B. simplex, q.v.) was collected from substrate drawn from Pit 81 of the Rancho La Brea Tar Pits in Los Angeles, California, excavated in 1915 (Miller et al. 1981; Miller 1986); co-occurring Equus remains were later dated ca. 11,000 years BP (Miller 1986). Pierce’s decision to place it near B. praticola is clearly reasonable inasmuch as the holotype (an elytron) is of the proper size (4.5 mm) and exhibits the dual granulation one would expect of this species. Moreover, while it does not presently occur west of Arizona, the presence of B. praticola at La Brea from the Pleistocene/Holocene boundary (ca. 11,000 years BP) is a reasonable assumption. The scarabaeine fauna at that time included at least Canthon, Copris, Onthophagus and Phanaeus (Pierce 1946, Miller 1983, Edmonds 1994). At the time, when the biotic landscape was quite different than now, the scarabaeine fauna of southern California was evidently very similar to the modern fauna of southern Arizona. Ample evidence suggests that B. praticola maintains a strongly preferential association with prairie dogs. Indeed, its US distribution corresponds rather closely to the combined ranges of Gunnison’s Prairie Dog, Cynomys gunnisoni (Baird, 1855) and the more widespread Black-Tailed Prairie Dog, C. ludovicianus (Ord, 1815) (Fig. 10). Field studies by Gordon and Cartwright (1974) in North Dakota and Colorado revealed that these beetles are commonly found in mounds surrounding burrow entrances where dung pellets were plentiful; they observed no activity in nearby bison droppings. James Saulnier (pers. comm.) has collected specimens in large numbers from active prairie dog colonies in northern Yavapai Co., Arizona. Floate and Gill (1998) found B. praticola to be a rare member of the cattle dung beetle community in southern Alberta, Canada, a coprofauna overwhelmingly dominated by aphodiines. Other than prairie dog droppings, B. praticola has been recorded from cattle feces as well as that of pig, bison, humans and rabbits. Edmonds (2018) reported B. praticola from open grasslands of the Big Bend region of west Texas, where it is commonly collected on cattle and human feces. McNamara (2013) recorded B. praticola from four Canadian provinces: Alberta, Saskatchewan, Manitoba and British Columbia; the latter, while not out of the question, needs confirmation. In addition to the syntype cited above, there is another probable syntype also present in the LeConte Collection at the Museum of Comparative Zoology, Harvard University (Crystal Maier, pers. comm). 18 · October 14, 2022 Edmonds Boreocanthon depressipennis (LeConte, 1859), restored generic combination Fig. 11–18, 28 Canthon depressipennis LeConte 1859c: 11. Boreocanthon depressipennis (LeConte) (new combination per Halffter 1958: 209). Canthon (Boreocanthon) depressipennis LeConte (new combination per Howden 1966: 727). Boreocanthon depressipennis LeConte, restored generic combination. Type material. Syntype (sex undetermined), Museum of Comparative Zoology, Cambridge (MCZ-ENT00003696). Examined by photograph (https://mczbase.mcz.harvard.edu/guid/MCZ:Ent:3696). Type locality. Georgia. Diagnosis. Head: Anterior portion of clypeus finely roughened, only slightly shinier than rest of head; posterior portion evenly granulate on shagreen background. Sexdentate (Fig. 11); clypeus with four strong teeth, paraocular notch deep, setting off dentate anterior angle of paraocular area. Labio-gular fimbria (Fig. 15) widely V-shaped, strongly narrowed medially, length exceeding one-half that of gula. Prothorax: Evenly, rather densely covered by smaller, rounded granules, granules isolated, only rarely coalescent (Fig. 11, 13b). Anterior pronotal angles not upturned; circumnotal ridge serrate posterolaterally, ridge extending beyond posterior angle to point even with elytral umbone (Fig. 13b, arrows). Hypomeral carina obsolete or only feebly indicated. Pterothorax: Mesoventrite densely, finely punctate medially, smooth laterally. Metaventrite (Fig. 12) with obsolete granules and scattered, minute punctures on shagreen background. Elytra: Interstriae (Fig. 11) evenly, rather densely granulate on shagreen background; 2nd and 3rd weakly raised. Striae superficial, edges ill-defined, course determined by lack of granulation (tracks through field of granules). Suhumeral (8th) stria not carinulate; epipleural (9th) clearly indicated. Legs: Inner margin of protibia (Fig. 16) clearly offset at level of basal (3rd) tooth, prolonged apically as conical tooth (Fig. 16 arrow). Protibial spur (Fig. 16) acute, curved mesally in both sexes. Hind femur lacking row of conspicuous setae along anterior margin. Abdomen: Pygidium (Fig. 14) depressed basally on each side of median swelling; evenly granulate. Sixth ventrite completely granulate; granules on remaining ventrites confined to lateral (upper) areas adjacent to elytral epipleuron. Genital capsule: Compressed, distal portion of parameres elongate, triangular (Fig.18), lower apical angles in form of elongate knobs (Fig. 17). General: Dorsum dark, often appearing black to unaided eye, almost always displaying dull green or yellow-green highlights under magnification. Length: 4.5 – 9.0 mm. Geographic distribution (Fig. 28): Southeastern United States, from eastern Louisiana to southern North Carolina, southward into the Florida peninsula. Ecogeographic environment (Fig. 2): Temperate Grasslands, Savannas and Shrublands biome (Southeast U.S. conifer savannas ecoregion). Specimens examined: 567. Collection localities. UNITED STATES—ALABAMA: Dale Co. ●Fort Rucker Military Reservation [May]. FLORIDA: Alachua Co. ●0.75 mi W Archer [Jun] ●4 mi N High Springs [Mar] ●10 mi SW Gainesville [Aug]; Baker Co. ●Glen St. Mary [Sep]; Calhoun Co. ●Clarksville [Mar]; Citrus Co. ●Inverness [May]; Clay Co. ●Camp Crystal (Mar-Apr]; DeSoto Co. ●27°19.356ʹ N 81°48.640ʹ W, 8 mi N Arcadia; Dixie Co. ●29°34.6ʹ N 82°58.3ʹ W, near Old Town ●3.7 and 5 mi N Old Town [May]; Duval Co. ●Jacksonville [Apr]; Escambia Co. ●Pensacola; Flagler Co. ●Hwy 1, 2 mi S Hwy I-95 [Oct]; Gilchrist Co. ●5 mi W Newberry (Alachua Co.) [Jun]; Hernando Co. ●Powell [Jul]; Highlands Co. ●Archbold Biological Station [Mar, Jun] ●Dover; Hillsborough Co. ●Tampa, University South Florida campus [May-Jun]; Lafayette Co. ●Picket Lake [Mar]; Lee Co. ●Hwy 41, 3.4 mi NW Koreshan State Park [Apr]; Lake Co. (no data); Levy Co. ●3.8 mi SW Archer [Mar]; ●2 mi W Archer [Jun]; ●Route 24, 0.2 mi S Alachua county line [Mar] ●Hwy 241, 1 km N Santa Fe River ●11 mi W Otter Creek [Nov]; Liberty Co. ●Apalachicola Bluffs & Ravines Preserve, Alum Bluff [Jul]; Madison Co. ●Pinetta [Aug]; Okaloosa Co. ●Valparaiso [Jul]; Orange Co. ●Walt Disney World [Aug]; Putnam Co. ●Red Water Lake [Jul]; Saint Johns ●St. Augustine; Sarasota Co. ●Myakka River State Park, 27°17.541ʹ N 82°17.420ʹ W [Sep]; Seminole Co. ●Econ River Wilderness Area [Jul]; Suwanee Co. ●7 mi N Live Oak [Mar]; Taylor Co. ●County Road 150, 5 mi W Hwy 19 [May]. (See also Woodruff for additional Florida records.) GEORGIA: Baker Co. ●Newton [Jun] ●Ichauway Ecological Reserve [Mar]; Burke Co. ●10.2 km NW Girard, Yuchi Wildlife Management Area, 33°05ʹ36ʺ N 81°48ʹ04ʺ W [Apr]; Charlton Co. ●Billys Island, Okefenokee Swamp [Oct] ●Camp Cornelia [May]; Chatham Co. ●12 km NNW Richmond Hill, 32°02ʹ41ʺ N 81°20ʹ13ʺ W [May]; Cook Co. ●Reed Bingham State Park [Jul]; Decatur Co. ●Spring Creek [Jul-Aug]; Dodge Co. Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 19 Figures 11–18. Boreocanthon depressipennis (LeConte). 11) Dorsal habitus (inset magnifies elytral sculpturing). 12) Oblique ventral view of mesoand metaventrites. 13) Comparison of posterolateral margin of pronotum; asterisks mark posterolateral angle of pronotum; arrows mark posterior extension of circumnotal ridge in B. depressipennis. a) Boreocanthon ebenus. b) Boreocanthon depressipennis. 14) Posterior view of pygidium. 15) Ventral view of mentum and gula. 16) Anterior view of protibia (arrow indicates apical tooth). 17) Apical view of parameres. 18) Lateral view of genital capsule. 20 · October 14, 2022 Edmonds ●1.2 mi NE Gresston [Mar]; Dougherty Co. ●(no data) [Sep]; Emanuel Co. ●Swainsboro [Jul]; Glynn Co. ●Brunswick [Jul]; Jenkins Co. ●Millen [Jul]; Johnson Co. ●0.5 mi E Kite [Oct]; Lowndes Co. ●Vidalia [Jul-Aug]; McIntosh Co. ●Sapelo Island, Nanny Goat Beach, 31.39065° -81.26533° [May]; Telfair Co. ● McRae (McRae-Helena) [Jul]; Thomas Co. ●Thomasville; Tift Co. ●Tifton [May, Jul]; Toombs Co. ●Vidalia [Aug]; Ware Co. ●Waycross [Sep]; Wheeler Co. ●Bells Ferry Road, along Ocmulgee River [Apr, Sep]. LOUISIANA: West Feliciana Par. ●Feliciana Natural Area, 30°47.7ʹ N 91°15.2ʹ W [Sep]. MISSISSIPPI: Jones Co. ●Laurel [Aug]. NORTH CAROLINA: Moore Co., ●Eastwood [Jul]. SOUTH CAROLINA: Aiken Co. ●Aiken Gopher Tortoise Preserve, 10.7 km NE White Pond, 33°30ʹ15ʺ N 81°24ʹ22ʺ W, 259 ft [Apr]; Beaufort Co. ●Beaufort [Apr]; Colleton Co. ●Islandton [Jun]; Florence Co. ●Florence [Sep]; Georgetown Co. ●10 km E Georgetown [May]; Horry Co. ●Myrtle Beach [Apr, Jul]. Comments. I regard this species as the sister of B. ebenus, the two having arisen from Pleistocene glaciation splitting the founding ancestral population into two enclaves, one on the Florida Peninsula and the other in a south-central refuge, probably in present day Texas. Holocene expansion may have brought the two into closer proximity than suggested by their presumed distributions (Fig. 28). I have seen two specimens (Louisiana State Arthropod Museum database 0243611 and 0305000) here regarded as B. depressipennis from eastern Louisiana (West Feliciana Parish) very near the Mississippi River. These specimens have the expected weak greenish cast and prolonged pronotal ridge but exhibit the coarser, more coalesced pronotal granulation of B. ebenus. On the other hand, I have also recorded B. ebenus adjacent to the Mississippi River in Bolivar Co., Mississippi (see Comments for that species). This area, along the southern portion of the Mississippi River, would be a good venue to test the notion that it is a zone of hybridization, whether by secondary contact or nascent divergence. On the type locality of B. depressipennis Woodruff (1973) stated, “Originally described from Kansas, although Robinson [1948] … erroneously listed the type locality as Georgia.” Woodruff in fact erroneously corrected Robinson as LeConte explicitly states that it is “… a species found in Georgia …” The distribution of B. depressipennis follows the coastal lowlands of the southeastern United States, including the Florida peninsula. Woodruff’s (1973) distribution map extends it also to the west and northward into a large portion of the Great Plains, which I assume is based upon erroneous records attributable to B. ebenus and B. praticola, which are very common in the region. This species very often presents a dark green color, but it can be black like its close relative, B. ebenus, within a single population. Fincher et al. (1970) reported that this species was commonly collected on chicken droppings, as well as opossum, rabbit, swine, fox and rat feces, but not those of human, sheep or dog, in Tift County, southern Georgia. Woodruff (1973) found it common in cow dung in open, sandy Florida pastures. In addition to the syntype cited above, there is another probable syntype also present in the LeConte Collection at the Museum of Comparative Zoology, Harvard University (Crystal Maier, pers. comm). Boreocanthon ebenus (Say, 1823). restored generic combination Fig. 13, 19–28 Ateuchus ebenus Say 1823: 208 Canthon ebenus (Say) (new combination per Melsheimer 1853: 53) Boreocanthon ebenus (Say) (new combination per Halffter 1958: 209) Canthon (Boreocanthon) ebenus (Say) (new combination per Howden 1966: 727) Boreocanthon ebenus (Say) restored generic combination Type. Neotype, here designated. Snow Entomological Collections, Biodiversity Institute, University of Kansas (see Comments below). Type locality. Kansas, 10 mi NW Mullinville, here restricted (see Comments below). Diagnosis. Head: Anterior portion of clypeus very finely roughened, weakly shiny, remainder of head densely covered by coarse granules on shagreen background. Strongly sexdentate, median clypeal teeth acute, separated by V-shaped emargination (Fig. 20); lateral teeth acute. Paraocular notch deep, setting off strong, dentate anterior angle of paraocular area. Labio-gular fimbria broadly V-shaped, reaching mid-length of gula (Fig. 25). Prothorax: Pronotum densely, coarsely granulate (Fig. 13a, 19–20); granules on disk frequently joined into short, irregular chains, those on sides usually isolated. Lateral margin not at all explanate anteriorly; circumnotal ridge serrate posteriorly, not extending beyond posterior angle (Fig. 13a). Hypomeral carina absent (Fig. 24) or rudimentary, Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 21 Figures 19–27. Boreocanthon ebenus (Say). 19) Habitus, anterolateral view; 20) same, dorsal view. 21) Apical view of female abdomen (pyg = pygidium; arrow indicates apices of paraprocts; asterisk indicates extended marginal flange of pygidial margin). 22) Lateral view of genital capsule. 23) Apical view of parameres. 24) Ventral view left hypomeron. 25) Ventral view of gula and submentum. 26) Oblique ventral view of mesoand metaventrites. 27) Posterior view of pygidium. 22 · October 14, 2022 Edmonds indicated only by angular projection of procoxal margin. Pterothorax: Pleurites granulate, granulation extending to sides of metaventrite (Fig. 26); mesoventrite shagreened, weakly punctate laterally; middle of metaventrite with weakened granulation and scattered punctures on shagreen background. Elytra: Interstriae bearing strong field of isolated granules, anterior ends of 2nd and 3rd interstriae weakly raised, that of 1st interstria depressed. Striae superficial, lacking discrete margins other than bordering granules; subhumeral (8th) stria never carinulate, epipleural (9th) stria sometimes indicated. Legs: Protibia (as in Fig. 16) abruptly widened, inner margin stepped at level of third (basal) tooth; inner apical angle prolonged as conical tooth. Protibial spur strongly curved and acute in both sexes. Hind femora with conspicuous row of long setae along anterior margin. Abdomen: Pygidium (Fig. 27) sculptured as elytra, impressed basally on each side of raised midline, or as continuous transverse concavity. Ventrites usually more or less evenly granulate, granules becoming weaker medially on ventrites 1–5, uniform across ventrite 6. Genital capsule: Compressed apical portion of genital capsule triangular (Fig. 22), lower apical angles in form of elongate, rounded lobes (Fig. 23). General: Black, very rarely showing strong green or weak blue undertones. Length: 6.0–11.0 mm. Geographic distribution (Fig. 28): Central United States generally west of the Mississippi River to the Rocky Mountains south of the 44th parallel, with southern extension through Texas and New Mexico west to southern Arizona. Ecogeographic environment (Fig. 2): Temperate Grasslands, Savannas and Shrublands, and Deserts and Xeric Shrubland biomes. Specimens examined: 1278. Notes on the neotype. The male neotype conforms with the diagnosis above annotated as follows: Specimen exhibits moderate wear, especially notably rounded marginal teeth of head and protibiae. Left protarsus missing. Hypomeral carina suggested by angular projection of raised procoxal margin. First ventrite with scant, weak granulation, almost smooth; ventrites 2–5 with weak lateral granulation; 6th ventrite completely granulate, posterior margin strongly emarginate medially. Genital capsule not extracted. Completely black at low magnification (< ×5), very weak green undertones emerge under strong light and moderate magnification (×15). Length 10 mm. Neotype was chosen because of its appropriate collection locality (see discussion below). Label information is given below. Collection localities. MEXICO — CHIHUAHUA: Mpio. Juarez ●Samalayuca [Jun] ●Samalayuca, Laguna de Riataso [Aug]. UNITED STATES — ARIZONA: Cochise Co. ●Chiricahua Mountains, Horseshoe Canyon, 1.6 mi W Hwy 80 [Sep]●5.8 km SE Willcox, 32°13ʹ10ʺ N 109°46ʹ51 Wʺ [Aug]. COLORADO: Denver Co. ●Denver [May-Aug]; El Paso Co. ●Peyton, 5675 ft [Jul] ●Cheyenne Mt., 6300 ft [Sep]; Logan Co. ●Meringo [Aug]; Morgan Co. ●3mi N Orchard [Jun]; Prowers Co. ●Lamar [Jun]; Weld Co. ●Nunn [Sep] ●Roggen [Aug] ●9 mi NE Roggen [May]. KANSAS: Barber Co. (no data); Cheyenne Co. ●9 mi NNE Wheeler [May]; Clark Co. ●5 mi N Englewood [Jun]; Clay Co. ●(no data) [Jun]; Comanche Co. ●4 mi NW Coldwater [Sep]; Cowley Co. ●10 mi SW Winfield; Douglas Co. ●Lawrence, 900 ft; Hamilton Co. ●Syracuse, 3500 ft [May]; Jefferson Co. ●Medina; Kiowa Co. ●10 mi NW Mullinville [Jun] ●Belvidere [Jul] 5 mi N Greensburg [Apr, Jun]; Meade Co. ●13 mi S Meade [Jul]; Morton Co. (no data); Reno Co. ●Medora [May-Jun]; Sedgwick Co. ●(no data); Trego Co. (no data); Wallace Co. ●(no data). MISSISSIPPI: Bolivar Co. ●2 mi W Rosedale [May-Jun]. MISSOURI: Jasper Co. (no data) [Aug.]. NEBRASKA: Arthur Co. ●4.5 mi N Arthur [May]●Arapaho Prairie [Jun-Jul]; Blaine Co. ●10 mi SW Brewster [Jul] ●3 mi N Brewster, 41.9721 N 99.8515 W; Brown Co. ●Koshopah [Jun]; Chase Co. ●Imperial; Cherry Co. ●Merritt Dam [Aug] ●Ft. Niobrara [Aug] ●2 mi W, 1 mi S Sparks [Jul] ●42.1643 N 100.5299 W [Jun]; Dakota Co. ●Sioux City [Jun]; Dundy Co. ●Benkleman [May] ●Sanborn, 5 mi W Haigler, 40.06° N 102.02° W, 1010 m [Jun]; Garfield Co. ●21 km E Taylor, 41.8179° N 99.1075° W; Hooker Co. ●41.9815° N 101.1139° W [Jun]; Keith Co. ●Cedar Point Biological Station [Jun]; Loup Co. ●41.8658 N 99.3773 W [Jul]; McPherson Co. ●Tryon [Jun] ●Gudmundsen Sandhills Laboratory [Jul]; Sheridan Co. ●10-13 mi E Alliance [Jun]; Thomas Co. ●Halsey [Aug] ●Nebraska National Forest, 1.3 km SSW Bessey Campground, 41.88937° N 100.30676° W [Sep] ●Nebraska National Forest, Whitetail Campground, 41.79413° N 100.27381° W [Sep]. NEW MEXICO: Bernalillo Co. ●Albuquerque [Apr, Jun]; Catron Co. ●Luna; Chaves Co. ●5.5 km SW Elkins [Jul] ●38 mi E Roswell, Mescalero Sand Dunes [Jul]; Cibola Co. ●Laguna [Jul]; Colfax Co. ●6.3 mi E Springer, Canadian River [Jun]; Doña Ana Co. ●Jornada Experimental Range, 15 mi N Las Cruces [Jul, Sep] ●Las Cruces [Jul] ●Rincon ●8 mi W Las Cruces [Jul]; Eddy Co. ●16 mi E Loving, Los Medanos [Aug] ●near Waste Isolation Pilot Plant, 32°23.6ʹN 103°46.1°ʹW [Jun]; Harding Co. ●33.5 km SE Gallegos, 1200 m [Apr]; Lea Co. ●3 mi E Jal [Jun] ●32°22.8ʹN 103°43.3ʹW [Jul]; Otero Co. Newman, Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 23 4000 ft [May]; Quay Co. ●Tucumcari [Sep] ●7 mi SW Logan [Jun] ●Glenrio [Jul] ●3 mi N San Jon; Roosevelt Co. ●Oasis State Park [Jul] ●Portales, 34.12371° -103.58029° [Jul] ●8 mi E Portales, ENMU Natural History Preserve [May-Jul, Oct] ●14 mi SW Portales; Sandoval Co. ●Jemez Pueblo [Jul] ●San Ysidro [Jun]; San Miguel Co. ●Las Vegas [Jul]; Sierra Co. ●Elephant Butte Lake State Park [Jul]; Socorro Co. ●19km W Bernardo, 34.401° N 107.028° W ●18 km SE Bernardo ●Sevilleta National Wildlife Refuge, Saltbush [Jun] ●Sevilleta National Wildlife Refuge, San Lorenzo [Jun] ●Sevilleta National Wildlife Refuge, Black Butte [Jun] ●Sevilleta National Wildlife Refuge, 34.311° N 106.678° W [Jun, Aug]; Union Co. ●18 mi W and 4 mi S Clayton (Snyder Ranch) [Aug]. OKLAHOMA: Cimarron Co. ●Boise City [Jul]; Cleveland Co. ●Norman [Apr] ●Hwy 62 at South Canadian River [Apr]; Cotton Co. ●Hwy 281 at Red River [Apr]; Jefferson Co. ●Hwy 79 at Red River [Apr] ●Ryan [Jul]; Kingfisher Co. ●Dover [Jul]; Love Co. Hwy I-35 at Red River [Apr]; Marshall Co. ●5 mi N Willis [Jun]; McCurtain Co. ●Red River sand bars [Jul]; Payne Co. ●(no data) [May]; Woods Co. ●Little Sahara State Park, 2 mi S Waynoka [May-Jun]; Woodward Co. ●Woodward [Jul]. SOUTH DAKOTA: Bennett Co. ●10 mi E Martin. Lacreek National Wildlife Refuge [May]; Perkins Co. ●Grand River National Grassland [Jul]. TEXAS: Anderson Co. ●Engeling Wildlife Management Area [Jul-Aug] ●25 mi NW Palestine [Apr]; Angelina Co. ●10 mi N Zavalla [May]; Atascosa Co. ●8 mi NW Poteet [Apr-May] ●5 mi SW Somerset (Bexar Co.) [Mar]; Bastrop Co. ●8.6 km N Bastrop, Camp Swift ●10 mi N Bastrop [Sep]; Bexar Co. ●San Antonio [Oct] ●10 mi W San Antonio [Oct] ●Hwy 281, 6 mi S Loop 1604 [Oct] ●Applewhite Rd. at Loop 1604 [Nov]; Brazos Co. ●College Station [Apr, Sep]; Bosque Co. ●2 mi SE Clifton; Briscoe Co. ●Quitaque [Jun]; Brooks Co. ●9 mi S Falfurrias [Jul]; Burleson Co. ●2 mi N junction Hwy 21 and FM 908 [Feb-Mar];●4 mi W Cooks Point [Mar]; Callahan Co. ●Baird [Apr]; Cameron Co. ●San Benito [Jul] ●14 mi E Rio Hondo [Oct]; Colorado Co. ●Columbus; Comal Co. ●Bulverde [Apr]; Comanche Co. ●Comanche [Jun-Jul]; Crane Co. ●Junction routes 1053 and 1233, 31°29ʹN 102°39ʹW [Jul]; Crosby Co. ●White River Lake [May]; Dallas Co. ●Dallas [May]; Dickens Co. ●2 mi SW Dickens [Oct] ●10 mi NE Dickens [Aug-Sep]; Dimmit Co. ●4 mi W Carrizo Springs [Sep] ●8 mi W Artesia Wells (LaSalle Co.), Chaparral State Wildlife Management Area [Mar]; El Paso Co. ●El Paso [Jun]; Freestone Co. ●Old Spring Seat Church, near Donie [May]; Gaines Co. ●10 mi SW Seagraves; Gillespie Co. ●1 mi S Fredricksburg [Oct]; Grimes Co. ●Lamb Spring [Mar]; Hall Co. ●6 mi SE Turkey [May-Jul]; Hardin Co. ●3 mi W Silsbee, Sandylands Preserve, 30°20ʹ57ʺ N 94°14ʹ16ʺ W [Jun]; Hardeman Co. ●20 mi N Goodlett, Farm Road 680 at Red River [Aug]; Henderson Co. ●Athens, Sand Flats Cemetery [Apr]; Hidalgo Co. ●Santa Ana Wildlife Refuge [Apr] ●7-10 mi N Linn [Apr]; Karnes Co. Ecleto Metz Ranch [Mar]; Kenedy Co. ●Turcotte Ranch, 2 mi S Sarita [May] ●Armstrong [Mar-Apr] ●3 and 5 mi S Sarita [Jun, Sep-Oct]; ●20 mi N Raymondville [Apr] ●Kenedy Ranch, San Pedro Camp, 26°57ʹ03.9ʺN 97°39ʹ22.2ʺW [Apr]; ●20 mi SW Riviera [Nov] ●Norias [Oct]; Leon Co. ●1.5 mi N Centerville [Jun]; Lipscomb Co. ●Higgins [Jun]; Live Oak Co. ●Mestena Ranch, 12 mi S George West [May] ●10 mi S George West [Aug]; Lubbock Co. ●Lubbock [Apr]; Mason Co. ●Double Helix Ranch, 30.87053 -99.05167; McColloch Co. ●2 mi SE Brady [Sep]; Mills Co. (no data); Montague Co. ●3 mi N Forestburg [Apr] ●Hwy 81 at Red River [Apr] Parker Co. ● Dennis [Jul]; Randall Co. ●Canyon [Jul]; Refugio Co. ●Austwell [May]; Robertson Co. ●Hearne [Apr]; Starr Co. ●2 mi NE La Gloria [May]; Terry Co. ●Brownfield [Jul]; Travis Co. ●Austin [Oct]; Val Verde Co. ●Del Rio, 955 ft [Jun]; Ward Co. ●Wickett [Jun] ●9.4 km ENE Monahans, 31°37ʹ11ʺ N 102°47ʹ52ʺ W [May] ●Monahans [May, Jul] ●Monahans Sandhills State Park, 31.639357° -102.829678° , 825 m [Apr -Jul]; Washington Co. ●Hwy 105, 8 mi SW Brazos River [Apr]; Wichita Co. ●2 mi ENE Burkburnett, US Hwy 281/I-44 at Red River [Jun]; Webb Co. ●Laredo [Jul]; Willacy Co. Lyford [Jul] ●Point Mansfield (May); Wilson Co. ●Floresville [Jul]; Winkler Co. ●Co. Rd. 404, 17 mi E Hwy 18, 31.792° N 102.791° W●11 mi NE Kermit, 3200 ft [Sep]. WYOMING: Goshen Co. ●Lingle [Jul]; Laramie Co. ●Cheyenne; Platte Co. ●Uva [Jul]. Comments. Boreocanthon ebenus is regarded here as sister to B. depressipennis (q.v.). It is a common, large species easily confused with B. praticola, from which it differs in a number of ways, most obviously by its basally impressed pygidium. The two species are broadly sympatric and sometimes collected together, in prairie dog colonies, for example. However, B. ebenus is clearly closely associated with sand dunes and other habitats with exposed, very sandy substrate (Brown 1927; Nealis 1977; Fincher et al. 1986). Collection locations are very often places where vegetated sand dune habitat is common; indeed, casual searching using Google Earth with precise GPS data often pinpoints places clearly having sand dune or similar habitat. Schoenly (1983) collected B. ebenus 30 · October 14, 2022 Edmonds fimbria broadly, shallowly V-shaped, apex followed by loose field of coarse, setose punctures. Prothorax: Pronotum evenly covered by variable-sized, sharp, well-spaced punctures on highly shiny background, background shagreen largely effaced (Fig. 38); anterior angles only weakly upturned, posterior portion of circumnotal ridge entire, not serrate, posteromedian angle not depressed. Hypomeral carina present, reaching one-fifth distance to lateral prothoracic margin (Fig. 40). Pterothorax: Mesoventrite coarsely punctured medially, smooth laterally. Metaventrite evenly, sparsely covered by multi-sized, sharp, fine punctures. Elytra: Interstriae (Fig. 39) very shiny, smooth, bright, 2nd and 3rd not swollen anteriorly; discal interstriae with ill-defined, scattered, minute punctures, shagreen background largely effaced. Striae superficial, lateral margins weakly defined; subhumeral (8th) stria finely carinulate; epipleural (9th) stria usually weakly evident at least anteriorly. Legs: Inner margin of protibia Figure 45. Approximate distributions of Boreocanthon melanus (Robinson), B. forreri (Bates) and B. ateuchiceps (Bates). Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 31 not offset; spur sexually dimorphic, acute apically in female, bifurcate in male. Hind femur lacking anterior row of conspicuous setae. Abdomen: Pygidium with sparse, shallow punctures on shagreen background. Last (6th) ventrite punctate medially; ventrites 1–5 smooth. Genital capsule: Distal portion of parameres triangular (Fig. 41), lower apical angles in form of elongate, rounded knobs (Fig. 42). General: Dorsum bright metallic green or yellow-green; venter colored similarly except ventrites 1–5, which are black. Length: 4.5–7.0 mm. Geographic distribution: Sierra Madre del Sur, Mexico? (Fig. 45, see Comments below). Ecogeographic environment: Tropical Dry Broadleaf Forest biome? Specimens examined: 4. Collection localities. See Comments below. Comments. Howden (1966) likened this species to B. forreri, while Bates (1887) placed it near Canthon deyrollei Harold. I do not consider it particularly close to B. forreri; it is an outlier species both morphologically and geographically, but, given especially the form of the clypeus and genital capsule, definitely a Boreocanthon. Canthon deyrollei is a Central American species only remotely related to Boreocanthon. Boreocanthon ateuchiceps is the only brightly metallic-colored member of the genus; the color of B. forreri (q.v.) is much darker, approaching black in some specimens. My knowledge of this species is very limited. I have been able to directly examine only four specimens labeled “Matamoros Pue. Tlacotli 8-ix-43 F. Islas” collected almost 80 years ago by Federico Islas. I have not been able to identify “Tlacotli” nor confirm that “Pue” means Puebla, and I can only guess that Islas’ reference to “Matamoros” is to the Pueblan city or municipality of Izúcar de Matamoros. Howden (1966) mentioned seeing specimens from the states of Guerrero and Mexico in The Natural History Museum (London); I have confirmed their presence there thanks to photographs provided by Max Barclay. Most are labeled “Tejupilco, Mex., Temascaltepec” and “Bejucos, Mex., Temascaltepec”. Each of these two towns is the seat of its own eponymous municipality in the state of Mexico, and both are located on the same highway (No. 134) descending southwest from the city of Temascaltepec into the Balsas River basin in southern Mexico. The type, also in London, bears the label “Acapulco, Guerrero Höge.” Others there bear the same Islas label as those accompanying specimens I have seen courtesy of Gonzalo Halffter. Islas (1942) himself cited it from “tierra caliente de Guerrero,” with no mention of any Puebla locality or Acapulco. Given what little I know, my best guess is that this species is a scarce denizen of the Sierra Madre del Sur, perhaps descending into the Río Balsas valley. Boreocanthon coahuilensis (Howden, 1966), new generic combination Fig. 46–53, 100 Canthon (Boreocanthon?) coahuilensis Howden 1966: 732. Canthon (Boreocanthon) coahuilensis Howden (subgeneric confirmation per Halffter and Martínez 1977: 82). Boreocanthon coahuilensis (Howden), new generic combination. Type material. Holotype male, Canadian National Collection, Ottawa (CNC312445 = CNC Type No. 9143). Examined by photograph (https://www.cnc.agr.gc.ca/taxonomy/Specimen.php?id=3684). Type locality. 21 mi SE Saltillo, Coahuila, Mexico, 6000 ft. Diagnosis. Head: Apical one-third of clypeus very finely rugose, slightly shinier than rest of head; posterior portion of clypeus and frons with dense shagreen and widely spaced, small punctures (Fig. 47). Clypeus usually distinctly quadridentate, if only weakly so; middle teeth moderately developed, rounded apically, separated by broad, rounded emargination; lateral teeth small, weakly angular, sometimes obsolete such that head appears bidentate (Fig. 46–47). Paraocular notch virtually absent, lateral margin of head only very slightly curved inward. Labio-gular fimbria V-shaped, apex extending about one-third length of gula. Prothorax: Pronotum convex, elevated above elytral surface (viewed laterally, Fig. 53b), not depressed posteromedially; surface weakly lustrous, silky, with dense shagreen and widely spaced, inconspicuous punctures and numerous weak microspots (Fig. 46, 48); lateral margin very weakly upturned apically. Underside of lateral margin bearing a small rounded tubercle near anterior end, which can be very small or obsolete. Hypomeral carina absent. Pterothorax: Mesoventrite minutely punctured laterally. Metaventrite evenly, widely punctate on shagreen background. Elytra: Interstriae sculptured like pronotum, 2nd and 3rd not swollen apically, 2nd, 3rd and 5th usually bearing minute tubercle on anterior end (sometimes obscured by pronotal margin). Discal striae (1st–6th) weak, lacking well defined margins, 32 · October 14, 2022 Edmonds Figures 46–53. Boreocanthon coahuilensis (Howden). 46) Habitus, dorsal view (inset magnifies elytral sculpturing). 47) Frontal view of head and pronotal margin. 48) Dorsal view of portion of pronotum. 49) Lateral view of genital capsule. 50) Apical view of parameres. 51) Anterior view of protibia. 52) Dorsal view of head. 53) Comparison of body shape in profile. a) Boreocanthon simplex. b) Boreocanthon coahuilensis. Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 33 lateral striae (7th–9th) weaker, effaced anteriorly, subhumeral (8th) never carinulate, humeral area with scattered, fine punctures. Legs: Inner margin of protibia abruptly widened (offset) at level of basal tooth (Fig. 51); protibial spur sexually dimorphic, apically acute in female, bifurcate in male. Hind femur lacking conspicuous anterior row of long setae. Abdomen: Pygidium sculptured like pronotum. Last (6th) ventrite punctate, more strongly so than pygidium; other ventrites shagreened, with few lateral punctures. Genital capsule: Compressed apical portion of parameres narrowly triangular (Fig. 49); lower apical angles in form of elongate, rounded knobs (Fig. 50). General: Black, pronotum and venter weakly shiny. Length: 5.5–10.0 mm. Geographic distribution (Fig. 100): Sierra Madre Oriental in extreme SE Coahuila and neighboring Nuevo Leon, Mexico. Ecogeographic environment (Fig. 2): Desert Shrublands biome (Meseta Central matorral ecoregion). Specimens examined: 40. Collection localities. MEXICO — COAHUILA: Mpio. Saltillo ●21 mi SE Saltillo, Coahuila, 6000 ft [Jul] ●Chapinque Mesa, 20 km SE Saltillo, 2050 m [Jul]. NUEVO LEON: Mpio. Galeana ●4 mi E San Roberto [Jul]; Mpio. Montemorelos ●Las Raíces [Aug]. Comments. In his original description, Howden (1966), with some doubt, assigned this species to Canthon using the heading “Canthon (Boreocanthon?) coahuilensis n. sp.”. That his hesitancy applied to subgeneric rather than generic placement is certain as he expressly (p. 727) treated Boreocanthon as a subgenus: “I have treated all [generic and subgeneric] divisions [of Canthon] as subgenera, and my assignments of some species considered below are tentative.” I gather that his hesitancy to assign B. coahuilensis to Boreocanthon derived especially from the variation in clypeal dentition that departed from the usual form in other Boreocanthon. Nevertheless, he was struck by obvious similarities to B. simplex. I agree with Howden’s assessment. Indeed, its closest relative appears to be B. simplex, with which it shares several common features: moderately to weakly developed clypeal teeth; small paraocular notch; basally effaced 7th and 8th elytral striae, and truncated profile of the parameres. The abruptly offset profile of the inner protibial margin is shared only with B. depressipennis and B. ebenus. The pronotum of B. coahuilensis is rather strongly humped (Fig. 53b), a feature rarely seen in other Boreocanthon. This species is so far known from but four nearby localities in far southern Coahuila and across the state border into the southwest corner of Nuevo Leon. My guess is that it extends farther to the south along the flanks of the Sierra Madre Oriental, at least as far south as southern San Luis Potosí, that is, along the length of the Meseta Central matorral ecoregion (Fig. 2). Its geographical isolation and sharp morphological distinctness bespeaks an early separation from B. simplex. Given that it is an arid land species, which regularly exhibit population explosions during sort-lived favorable conditions, I rather doubt that it is rare, or even uncommon, in spite of what appears might be a very localized distribution. Howden’s description was based on a type series of 35 specimens collected near Saltillo, Coahuila. For this study, I was fortunate to examine two other series totaling 37 specimens collected from horse and cattle dung by Paul Lago from Chapinque Mesa, Coahuila, and San Roberto, Nuevo Leon, as well as three specimens from the Halffter collection from Las Raíces. Boreocanthon puncticollis (LeConte, 1866), restored generic combination Fig. 54–63 Canthon puncticollis LeConte 1866: 381. Boreocanthon puncticollis (LeConte) (new combination per Halffter 1958: 210). Canthon (Boreocanthon) puncticollis LeConte (new combination per Howden 1966: 727). Boreocanthon puncticollis (LeConte), restored generic combination. Canthon nyctelius Bates 1887: 31 (new synonymy per Schaeffer 1915: 50). Canthon mixtus Robinson 1948: 91, new synonymy. Type material. 1) Canthon puncticollis LeConte: syntype, female. Museum of Comparative Zoology, Harvard University, MCZ-ENT00003698. Examined by photograph (https://mczbase.mcz.harvard.edu/guid/MCZ:Ent:3698). 2) Canthon nyctelius Bates: Syntype (sex undetermined). The Natural History Museum, London. Not examined. 3) Canthon mixtus Robinson: Holotype, male. National Museum of Natural History, Washington, D.C. Not examined. 34 · October 14, 2022 Edmonds Figures 54–62. Boreocanthon puncticollis (LeConte). 54) Habitus, dorsal view. 55) Habitus, frontal view. 56–57) Apical views of parameres. 58) Lateral view of genital capsule. 59) Dorsal view of mid-body (note triangular, posteromedian depression of pronotum and swelling of interstriae). 60) Ventral view of submentum and gula. 61) Oblique dorsal view of pronotum and elytron. 62) Comparison of unworn (above) and heavily worn (below) head margin (dorsal views). Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 35 Type localities. 1) Canthon puncticollis LeConte: Cape San Lucas, Lower California [Mexico]; 2) Canthon nyctelius Bates: Santa Clara (Chihuahua), Chihuahua City, Durango City (Durango); 3) Canthon mixtus Robinson: Marfa, Texas. Diagnosis. Head: Clypeus (Fig. 55, 62) finely roughened, weakly shiny; remainder of head surface distinctly punctate on shagreen background with very weak microspotting. Sexdentate, clypeal teeth prominent, rounded apically, median pair separated by broadly rounded emargination, paraocular notch large, nearly right-angled, anterior angle of paraocular area obtusely dentate. Labio-gular fimbria broadly V-shaped, constricted apically and reaching about one-third length of gula. Prothorax: Pronotum (Fig. 59, 61) boldly, conspicuously punctate, punctures shallow, evenly distributed over entire surface shagreen, microspotting conspicuous. Anterior angles of pronotum only very slightly reflexed; postero-median angle depressed as dark, triangular declivity adjacent anterior ends of first elytral interstriae (Fig. 55, 59). Hypomeral carina absent. Pterothorax: Mesoventrite smooth, sometimes with few coarse, median punctures. Metaventrite shiny, evenly covered by small, sharp, evenly dispersed punctures. Elytra: Interstriae (Fig. 59, 61) alutaceous, microspotting dense, punctures absent. Striae bold, with crisp, sharply defined edges, especially anteriorly. Anterior ends of 2nd and 3rd interstriae swollen, those of 1st depressed, forming with the adjacent depression of the pronotum a rimless median “crater” or fossa (Fig. 55, 59). Subhumeral (8th) stria carinulate, usually strongly so; epipleural (9th) stria almost always distinct apically, occasionally weakly carinulate. Legs: Inner margin of protibia gently curved, not offset. Protibial teeth sexually dimorphic, apically acute in female, bifurcate in male. Metafemora lacking conspicuous anterior row of long setae. Abdomen: Pygidium weakly punctured on shagreen background, microspotting weak, apex sometimes more convex and shinier than rest of surface. Sixth ventrite with small median field of coarse punctures, remaining ventrites smooth, impunctate. Genital capsule: Compressed distal portion of parameres (Fig. 58) triangular, lower apical angles bearing lateral, ear-like flanges (Fig. 56–57). General: Black, usually with strong blue highlights, very rarely with green highlights. Length: 4.0 – 6.5 mm. Geographic distribution (Fig. 63): Southern Baja Peninsula and Central Mexico from the Transverse Volcanic Axis to far western Texas and southern Arizona. Ecogeographic environment (Fig. 2): Desert and Xeric Shrublands biome. Specimens examined: 833. Collection localities. MEXICO — AGUASCALIENTES: Mpio. Aguascalientes ●15 mi E Aguascalientes. BAJA CALIFORNIA: Mpio. Ensenada ●4.3 mi W El Rosario [Sep]. BAJA CALIFORNIA SUR: Mpio. Los Cabos ●5 mi N Cabo San Lucas [Oct] ● Santiago ●San José del Cabo ●11 km N Santa Anita [Aug] ●20 km NW Cabo San Lucas, 120 m [Sep] ●22 km NW Cabo San Lucas, 23°01.538ʹ N 110°04.895ʹ W, 30 m [Jul] ●4.3 mi SW Miraflores ●Miraflores [Jul]; Mpio. Mulejé ●San Venancio [Oct] ●Santa Rosalia; Mpio. La Paz ●2.5 mi SE La Huerta, 2200 ft [Oct] ●9.5 mi S San Pedro [Sep] ●3 mi N San Pedro [Jul] ●La Paz [Oct] ●8 mi SE La Paz, 1000 ft [Oct] ●17 mi S La Paz [Sep] ●15 mi W La Paz ●11 mi S Todos Santos, Playa Los Cerritos [Sep] ●2 mi W El Triunfo, 1900 ft [Oct] ●3 mi S El Triunfo [Aug-Sep] ●11.8 mi N El Triunfo [Sep] ●0.5 mi NW El Triunfo, 1800 ft [Oct] ●7 mi W El Triunfo, 1500 ft [Oct] ● Sierra de la Laguna, 3 mi W Cuadaño [Sep] ● Sierra de la Laguna, 5 mi S San Antonio [Sep] ●Sierra de la Laguna, 27 km W Hwy 1 on Ramal a Las Naranjas, 23°14ʹ12ʺ N 109°57ʹ15ʺ W, 2450 ft [Jul-Aug] ●3.5 mi W Los Barriles, 500 ft [Dec]; Mpio. Loreto ●Canipolé [Oct]. CHIHUAHUA: Mpio. Chihuahua ●12 km N Chihuahua [Aug]; Mpio. Janos ●45 mi NW Casas Grandes [Aug]. COAHUILA: Mpio. Castaños ●1 mi NE Junction Hwy 53 and Baján Rd.; Mpio. Ocampo ●35.5 mi W Ocampo, Laguna La Leche Flats, 27°17.0ʹ N 102°53.6ʹ W ●Sierra El Carmen (Maderas del Carmen), 28°59ʹ54ʺ N 102°36ʹ 42ʺ W, 2310 m. [Jul] ●Boquillas del Carmen [Jul]; Mpio. Saltillo ●6 km E Saltillo [Jul]. DISTRITO FEDERAL: ●Tepeyac [Aug]. DURANGO: Mpio. Cuencamé ●Hwy 49, 20 km SE Cuencamé; Mpio. Durango ●Tapías, 6400 ft [Jun]; Mpio. Lerdo ●La Loma, 4100 ft [Aug]; Mpio. Mapimí ●Reserva de la Biósfera Mapimí [Jun, Sep]; Mpio. Ocampo ●173 mi N Durango [Aug]; Mpio. Saucillo ●Estación Conchos [Jun]. HIDALGO: Mpio. Jacala de Ledezma ●7 mi N Jacala [Jul]; Mpio. Zimapán ●Zimapán [Jun]. OAXACA: Mpio. Santiago Chazumba ●San Sebastián Frontera [Jul]. PUEBLA: Mpio. Tepeyahaulco ●Alchichica [Jun]. SAN LUIS POTOSÍ: Mpio. Guadalcázar ●19.6 mi N Huizache [Jul]; Mpio. Matehuala ●Matehuala [Jul]. SONORA: Mpio. Naco ●Naco, 5000 ft [Aug]. ZACATECAS: Mpio. Villa de Cos ●Tropic of Cancer marker, Hwy 54 (Zacatecas – Saltillo), ~ 23°26ʹ12ʺ N 102°12ʹ05ʺ W, 1958 m [Jul]. UNITED STATES — ARIZONA: Apache Co. ●White Mountains, Diamond Creek [Aug]; Cochise Co. ●10 mi N Portal [Sep] ●2 mi NE Portal [Jul] ●4 mi NNW Portal, 31°58ʹ N 109°09ʹ W, 4600 ft ●13 mi W Gleeson 36 · October 14, 2022 Edmonds [Aug] ●Dragoon Mountains, Cochise Stronghold [Jul] ●Dos Cabezas [Aug] ●Palominas [Aug] ●Guadalupe Canyon [Jul] ●Douglas [Jul] ●15 mi E Douglas [Aug] ●Turkey Creek Canyon, Chiricahua Mountains, 31.8657° N 109.4125° W, 1642 m [Jul] ●San Simon [Aug]; Graham Co. ●12 mi S Safford [May]; Mojave Co. ●17 mi S Wikieup [Aug]; Pima Co. ●Tucson [Sep] ●Saguaro National Park (West) [Aug] ●Green Valley [Sep] ●18 mi S Tucson [Jul]; Santa Cruz Co. ●Nogales [Aug] ●Santa Rita Mts [Aug]; Yavapai Co. ●4 mi N Clarkdale [Jul]. CALIFORNIA: San Bernadino Co., New York Mountains [Aug]. NEW MEXICO: Catron Co. ●5 mi NW Reserve [Jul]; Doña Ana Co. ●La Jornada Experimental Range, 15 mi N Las Cruces [Jul-Sep] ●6 km NW Mesilla Dam ●Las Cruces [Jul-Aug] ●1 mi SE Afton Pump Station [Jul] ●10 mi NE Las Cruces, 32.490206 -106.781367, 4523 ft; Grant Co. ●Silver City ●8 mi E Silver City [Jul]; Hidalgo Co. ●1 mi N Rodeo [Jul]●5 mi N Rodeo [Aug] ●13 mi NE Rodeo [Jul] ●1 mi S Rodeo; Luna Co. ●Deming [Jul-Aug] ●6 mi NW Columbus [Jul]; Otero Co. White Sands Figure 63. Approximate distributions of Boreocanthon puncticollis (LeConte) and B. integricollis (Schaeffer). Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 37 National Park, Lake Lucero [Aug] ●Lake Holloman Camp, 32.811° N 106.121° W, 1230 m; Quay Co. ●1.1 mi N Tucumcari, 4100 ft [Aug]; Sandoval Co. ●4 km NE Sandia, Sandia Pueblo, 35.224547 -106.529917 [Jul]; Sierra Co. ●Elephant Butte [Oct] ●Eagles Point, Elephant Butte Reservoir [Aug]. TEXAS: Brewster Co. ●~17 km W Alpine (Paisano Baptist Encampment), 30°17ʹ37ʺN 103°47ʹ35ʺW, 1550 m (Jul) ●Chisos Mountains [Jul] ●15 mi S Marathon [Jun] ●20 mi E Marathon, 30°12ʹ04ʺ N 102°54ʹ47ʺ W [Jun] ●Boquillas [Jul] ●Tornillo Creek at Hwy 227, Big Bend National Park [Jul] ●Elephant Butte Wildlife Management Area, 26 mi S Alpine [Apr] ●Big Bend National Park, Buttrill Springs [Jul]; Culberson Co. ●17 mi E Van Horn [Jul, Sep] ●4 mi E Kent [Jun]; Dickens Co. ●2 mi E Dickens [May]; Ector Co. ●Odessa, 31.884°N 102.32°W [Jul]; El Paso Co. ●18.5 mi E El Paso, Hueco Mts. [Sep]; Howard Co. ●Big Spring; Hudspeth Co. ●8 mi W Sierra Blanca, 31.21° N 105.49° W, 1390 m [Aug]; Jeff Davis Co. ●Ft. Davis [Jul] ●16 km S Fort Davis (along TX 17), 30°27ʹ48ʺN 103°58ʹ59ʺW, 1600 m [Aug] ●Valentine [Jul] ●13 mi E Valentine [Aug] ●~16 km NE Valentine, Muerto Springs Ranch (Muerto Springs, near headquarters), 30°40ʹ28ʺN 104°24ʹ07ʺW, 1475 m [Jul] ●8 km SE Fort Davis (via TX 118), Chihuahuan Desert Research Institute (Visitor Center area), 30°32ʹ32ʺN 103°50ʹ11ʺW, 1555 m [Aug] ●Chihuahuan Desert Research Institute (Quarry Unit), 30°32ʹ06ʺN 103°50ʹ37ʺW, 1480 m [Aug–Sep]; Pecos Co. ●25 mi W Fort Stockton [Jul]; Motley Co. (no data) [May]; Presidio Co. ●C.E. Miller Ranch (~16 km W Valentine), 30°32ʹ50ʺN 104°39ʹ40ʺW (Camp Holland) 1410 m (Aug) ●~16 km W Valentine (Miller Ranch, near headquarters), 30°33°30ʺN 104°38ʹ44ʺW, 1350 m [Jul–Aug] ●~6.5 km W Marfa (Hip-O Ranch), 30°21ʹ54ʺN 104° 7ʹ12ʺW, 1530 m [Aug–Sep] ●Pinto Canyon Ranch (~58 km SSW Marfa on FM 2810), 30°01ʹ18ʺN 104°27ʹ42ʺW (headquarters area), 1475 m [Aug] ●Dalquest Research Site, 29°33ʹ30ʺN 103°47ʹ30ʺ, 1010 m [Sep]; ●37 km SSW Marfa (along FM 2810, Petan Ranch – Cherry Hills sector), 30°07ʹ35ʺN 104°19ʹ24ʺW, 1630 m [Jun] ●20–26 km SSE Marfa (along FM 169), 1355–1415 m [Jun] ●3 km NE Marfa (along FM 1112), Marfa Golf Course, 30°19ʹ40ʺN 103°59ʹ41ʺW, 1470 m [Jul, Sep] ●~30 km SSE Marfa (along FM 169), Humphreys Ranch, 30°02ʹ30ʺN 104°01ʹ00ʺW, 1285 m [Jul] ●~8km W La Viuda Peak, 29°42ʹ30ʺN 103°54ʹ30ʺ, 1200 m [Sep] ●~19 km E Marfa (via US 90/67), 30°16ʹ07ʺN 103°48ʹ44ʺW, 1565 m [Jul] ●~3 km N Marfa (along TX 17), 30°20ʹ27ʺN 104°01ʹ7ʺW, 1500 m [Jul] ●near Plata (FM 169/Alamito Creek bridge) Kennedy Ranch (headquarters), 29°52ʹ16ʺ N 103°59ʹ28ʺ W, 1200 m [Jun] ●Pinto Canyon Ranch (~65 km SSW Marfa on FM 2810), 29°59ʹ54ʺN 104°30ʹ48ʺW (Pinto Canyon Creek), 1460 m [Aug] ●7 mi N Presidio [Oct] ●Big Bend Ranch State Park, La Cuesta Campground, 29.298° N 103.954° W [Aug] ●FM 170 (River Road) jct. Casa Piedra Rd., 8 mi NW Redford ●2.5 mi S Shafter [Jun]; Terrell Co. ●Sheffield [Jun] ●~15 mi W Sanderson, Hwy 90 at Longfellow Draw, 30°09.7ʹ N 102°38.2ʹ W, 3380 ft [Apr] ●Oasis Ranch, Independence Creek, 15 mi S Sheffield, 595 m, 30.4665° N 101.8008° W [May]; Tom Green Co. ●12.8 km N San Angelo [Oct]; Ward Co. ●1 mi SE Barstow [Jul]. Comments. Boreocanthon puncticollis is a common species in arid central Mexico and the southwestern United States as well the southern portion of the Baja Peninsula. It is sister species to B. integricollis (q.v.), with which it shares apically flanged parameres, strong (“crisp”) dorsal sculpturing, and tumid 2nd and 3rd elytral striae. The apical flanges of the parameres vary somewhat in shape (Fig. 56–57), but the variation is subtle and appears continuous and taxonomically insignificant. Besides its presence in the southern portion of the Baja California Peninsula, this species is widely distributed across the arid highlands (Altiplano) of Mexico between the Sierra Madre cordilleras flanking it on the east and west, from which it ranges into Arizona, New Mexico and the western portion of Texas. Whether or not the Baja California Sur distribution of B. puncticollis is truly disjunct from its distribution on the mainland remains an open question. My inclination is that it is, indeed, isolated as shown in Figure 63. The northern portion of the Sonoran Desert (i.e., the Yuma Desert portion of the Sonoran Desert) appears to produce a collective void in the distribution of the genus. (See biogeographical comments in the Introduction.) In their analysis of dung beetle ecology in Mexican arid zones, Halffter et al. (2012) pointed out that “… only few species … range across the expansive arid zones [of Mexico]. These species are widely distributed and co-occur in the dry habitats of the northern Mexican Altiplano and southwest of the United States … Canthon (Boreocanthon) puncticollis is distributed in the desert zones of Baja California, Arizona and Chihuahua, arid zones of the Mexican Altiplano (even near Barranca de Metztitlán [Hidalgo], with a southern limit in Tehuacán [Puebla-Oaxaca].” I have seen several specimens of B. puncticollis from the Tehuacán–Cuicatlán desert region of Puebla/Oaxaca, Mexico, and suspect 38 · October 14, 2022 Edmonds that this region marks the southernmost point of its distribution. I have seen a single specimen labeled Puente de Ixtla, Morelos, but its presence there needs confirmation. Blume’s (1972) reference to B. puncticollis in Kerr and Bexar counties, Texas, is likely incorrect and should probably read B. integricollis. As far as I know, B. puncticollis does not occur as far east as the Edwards Plateau of central Texas. This species is commonly collected along with B. praticola throughout the Texas–New Mexico– Arizona portion of its range. As with most other congeners, this species is poorly known biologically. Edmonds (2016) reported it to be diurnal and a generalist feeder (cattle, human, and equine feces) inhabiting grassland and open scrub habitats in the Big Bend region of west Texas; M. J. Paulsen (pers, comm.) reported collecting it from burrows (probably a species of ground squirrel) in Hudspeth County, Texas. Lobo (1996) collected specimens in low numbers from cattle dung in various desert habitats at Mapimí Biosphere Reserve in northern Durango, Mexico, where the dung beetle fauna was overwhelmingly dominated by two introduced scarabaeines, Digitonthophagus gazella (Fabricius) an Euoniticellus intermedius (Reiche) (see Edmonds 2018). LeConte’s (1866) original description of Canthon puncticollis was based on two specimens from “Cape San Lucas, Lower California.” The description is somewhat ambiguous in several respects, but the photographs of the syntype I have examined clearly show the salient features used here to diagnose this common species. Bates (1887) distinguished his species, Canthon nyctelius, from B. puncticollis by its having a “… remarkable depression at the base of the thorax and elytra, and the corresponding elevation of the second and third elytral interstices at their base.” Schaeffer (1915), in his argument supporting their synonymy, pointed out that B. puncticollis also shares this feature and noted that it was not clearly mentioned by LeConte (1866) in his description. Bates (1887) did not consider B. puncticollis in his work and probably did not have specimens on hand to compare with his species, prompting him to rely on LeConte’s description for comparison. In addition to the syntype cited above, there is another probable syntype also present in the LeConte Collection at the Museum of Comparative Zoology, Harvard University (Crystal Maier, pers. comm), and the two specimens are almost certainly LeConte’s original type series. Robinson (1948) based his description of Canthon mixtus on three specimens. I was not able to examine the holotype, which resides in the USNM collection in Washington, D.C. However, I have examined the paratype referenced by Robinson from the Casselberry Collection, now deposited in the collection of the American Museum of Natural History. It lies well within the taxonomic limits I adopt here for B. puncticollis, which I here regard as senior synonym. My references (Edmonds 2018) to Canthon mixtus in western Texas should now be taken as B. puncticollis. Boreocanthon integricollis (Schaeffer, 1915), restored generic combination Fig. 63–71 Canthon puncticollis var. integricollis Schaeffer 1915: 50. Canthon integricollis Schaeffer (new status per Robinson 1948: 90). Boreocanthon integricollis (Schaeffer) (new combination per Halffter 1958: 210). Canthon (Boreocanthon) integricollis Schaeffer (new combination per Howden 1966: 727). Boreocanthon integricollis (Schaeffer), restored generic combination. Type material. Syntype (sex undetermined). National Museum of Natural History, Washington, D.C. Not examined. Type localities. Hidalgo and Brownsville, Texas; Santa Rita Mountains, Arizona (see Comments below). Diagnosis. Head: Margin strongly sexdentate (Fig. 66), anterior one-half of clypeus rugose and moderately shiny; remainder of head surface flat, with fine, shallow punctures on dense shagreen. Paraocular notch strong, setting off angulate, anteriorly-directed, toothlike angle of paraocular area. Middle clypeal teeth narrowly rounded apically, separated by rounded, V-shaped emargination; lateral teeth strong, rounded apically. Labio-gular fimbria (Fig. 69) gently curved posteriorly, often interrupted medially by coarse punctures. Prothorax: Pronotum (Fig. 65, 71) covered by dense shagreen, punctured, punctures scattered, somewhat larger, deeper than those on head, microspots numerous; anterior angles weakly, narrowly reflexed; posteromedian angle at most only slightly impressed. Hypomeral carina vestigial. Pterothorax: Mesoventrite (Fig. 70) densely, coarsely punctured. Metaventrite (Fig. 70) with widely spaced, small punctures on dense shagreen. Elytra: Interstriae (Fig. 64, 67, 71) with Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 39 Figures 64–71. Boreocanthon integricollis (Schaeffer). 64) Habitus, dorsal view. 65) Habitus, anterodorsal view. 66) Dorsal view of head. 67) Dorsal view of mid-body. 68) Genital capsule. a) Lateral view. b) Apices of parameres. 69) Ventral view of labium and gula. 70) Ventral view mesoand metaventrites. 71) Dorsolateral view of pronotum and elytron. 46 · October 14, 2022 Edmonds Figures 83–89. Boreocanthon halffteri, new species. 83) Holotype male, dorsal view. 84) Holotype male, oblique anterolateral view. 85) Lateral view of genital capsule. 86) Apical view of parameres. 87) Dorsal view of portion of pronotum. 88) Dorsolateral view of elytron (arrow marks carinulate 8th stria). 89) Dorsal view of head margin. Notes on the holotype. The holotype male (Fig. 83–84) conforms with the diagnosis above annotated as follows: Left protibia-tarsus missing. Pronotal microspots dense, almost contiguous in some places. Edges of elytral striae 1–6 sharply defined by carinulate margins for almost entire length; 8th elytral stria with strong carinula extending from umbone to point even with first abdominal ventrite; 9th stria obsolete but vaguely indicated. Apex of pygidium distinctly convex, somewhat shinier than base, microspots virtually effaced, shagreen extremely fine (×50). Last abdominal ventrite with small, shiny bump at middle of posterior margin. Genital capsule not extracted. Length: 6.0 mm. The holotype was chosen because it conserves almost unblemished surface sculpturing. It carries Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 47 the following five labels: (1) TX: El Paso Co., red dunes 14 NE of Fabens, VII-11-96, C. Wolfe & D. Marqua. (2) ex. Charles S. Wolfe Collection, June 2006, TAMU Insect Collection. (3) TAMU-ENTO, X0294090. (4) [small light yellow circle]. (5) HOLOTYPE, Boreocanthon halffteri Edmonds, 2022. [Label notes: Distance units not specified in label (1), presumed to be 14 miles. Small yellow circle is reference to habitus photographs, Fig. 83–84. All labels on white paper printed in black, HOLOTYPE in red]. Collection localities. MEXICO — CHIHUAHUA: Mpio. Juarez ●Samalayuca [Jun]. UNITED STATES — NEW MEXICO: Bernalillo Co. ●Albuquerque [Jun-Jul, Oct] ●Volcano Cliffs, Albuquerque [Aug]; Doña Ana Co. ●2.5 mi N Mesquite [Oct] ●USDA Jornada Experimental Range, 32.619590 -106.787845, 4356 ft [Jul-Aug] ●Anthony [Jun]; Sandoval Co. ●Coronado State Park, Bernalillo [Aug]; Socorro Co. ●Bosque de Apache NWR [Jul] ●Sevilleta NWR, Rio Salado [Jul] ●17 km SW/S Bernardo, 34.292° N 106.928° W [Jul] ●Sevilleta NWR, LTER Site 3,4 [Jul, Sep]; Torrance Co. ●Town of Gran Quivara, 6500 ft [Aug]; Valencia Co. ●5 mi N Belen [Jul]. TEXAS: El Paso Co.● red dunes on Montana Ave., 31.841°N 106.131°W, 1245 m [AugSep] ●14 mi NE Fabens [Jul]. Paratype data (arranged by institutional/personal collection; all bear my determination labels printed in black on white paper with PARATYPE in blue and dated either 2021 or 2022; no allotype designated): American Museum of Natural History. New York NY – 3 females, Samalayuca Chih[uahua] Mexico, vi-24-47, Rockefeller Exp[edition], [C.D.] Michener, Accession Codes 00011194–96. Arizona State University, Hasbrouck Insect Collection, Tempe AZ – 1 male, NM Doña Ana Co., Anthony, 24-vi-70, M. Cazier, sandy hills, depressions, Atriplex ASUHIC 0145853. California Academy of Sciences, San Francisco CA – 28: 12 males, 15 females, New Mex., Torrance Co., Town of Gran Quivara, 6500 ft. alt., 29-viii-1967, Hugh B. Leech, ex vertical tunnel in sandy soil; one female, New Mex., Torrance Co., Town of Gran Quivara, 29-viii-1967, Hugh B. Leech, with humus in burrow. Carnegie Museum of Natural History, Pittsburgh PA – 4: 2 males, one female, TX: El Paso Co. Red Sands, US 62, 180, Sept. 17, 1985, Col. G.A. Coovert, GAC 1254; one male, NM: Bernalillo Co., Albuquerque, X-18-1928, H. Klages Collection, CM Acc No. 11414. Monte L. Bean Life Science Museum, Brigham Young University, Provo UT – one male, NM: Sandoval Co., Coronado State Park, Bernalillo, 21 Aug 1985, Bauman, Huish, Nelson, Wells, Whiting. Museum of Southwestern Biology, University of New Mexico, Albuquerque NM – 29: one male, Socorro Co. NM, Jul 89, C. Blanco M.// MSB Cat. No. 8226; 3 females, one male, NM: Socorro Co., Sevilleta NWR, Rio Salado, vii-17-1995, D. Holiday; 3 females, 2 males, USA NM Socorro Co., 17 km SW/S Bernardo, N 34.292 W 106.928, 17 July 1998, coll. S.L. Brantley // MSB Cat. No. 17784–17790; 2 males, one female, Bernalillo Co., NM, Volcano Cliffs, Alb., 10 Aug 1979, C. S. Crawford // MSB Cat No. 7851–7854; 2 males, Valencia Co, NM, 5 mi N Belen, 21 July 1987, K. Flies // MSB Cat No. 7855 and 7857; 2 females, one male, Socorro Co., NM, Sevilletta [sic] sand dunes, 20 April 1982, C. S. Crawford; 3 females, one male, Socorro Co. NM, Sevilleta NWR, LTER Site 3, 4 Sept. 1992; 2 males, USA New Mexico, Dona Ana Co., USDA Jornada Experimental Range, 32.619590 -106.787845, 17 August 1989, 4356’elev., mesquite dunes, pitfall trap, D. C. Lightfoot collector; one male, Socorro Co. NM, Sevilleta NWR, LTER Site 3, 31 July 91; one male, N. Mex., Dona Ana Co., 2.5 mi N Mesquite, 2 Oct 1982, D. Lightfoot; one female, Socorro Co. NM, Bosque de Apache NWR, line A, trap #6, 14 July 1996; one female, Bernalillo Co, NW Albuquerque, 16-VI-93, Dick Fagerlund // MSB Cat. No. 8225; one female, N. Mex., Dona Ana Co., Jornada Range, 15 m N of Las Cruces, 24.VII.79, pit trap // David Richmond. University of Nebraska State Museum, Lincoln NB – 8: 1 male, 7 females, TX: El Paso Co, red dunes on Montana Ave., 31.841° N 106.131° W, 1245 m, rolling AM after PM rains, 22-VIII2010, M. J. Paulsen. New Mexico State University Arthropod Collection, Las Cruces NM – 20: NMSUAC Accession Codes (yellow labels): Males (6): 0016583, 0020360, 0020364, 0020371, 0020375, 0020385. Females (14): 0016584, 0020357, 0020359, 0020361, 0020362, 0020365, 0020366, 0020368, 0020370, 0020377, 0020378, 0020384, 0020386, 0020389 Locality Data: 0016583 and 0016584: NM, Doña Ana Co., Jornada Exp. Range, pit trap, vii-14-1989, D. Richman; all others: N. Mex., Doña Ana Co., Jornada Range, 15 m N Las Cruces, 24-vii-79, D. Richman. Paul K. Lago Collection, University MS – 2: one male, one female, Red sands US [Hwy] 62,180, El Paso Co. Tex., Sept. 17, 1985, Coll. G.A. Covert//GAC 1254. Snow Entomological Collections, University of Kansas, Lawrence KS – one male, NM, Albuquerque, VII-5-41, E. L. Todd. Richard A. Cunningham Collection, Show Low AZ – 2 males, N. Mex., Doña Ana Co., Jornada Range, 15 m N of Las Cruces, pit trap, 24 vii 79. 48 · October 14, 2022 Edmonds Texas A&M University Insect Collection, College Station TX – 22: 7 males, 4 females (same data as holotype) database numbers TAMU-ENTO X 0295573, 0296014, 0296864, 0297061, 0297466, 1205379, 1212529, 1224107, 1224300, 1224163, 1224681; 2 males, 2 females [ex CAS], New Mex., Torrance Co., Town of Gran Quivara, 6500 ft. alt., 29-viii-1967, Hugh B. Leech, ex vertical tunnel in sandy soil, X 1224343, 1695095, 1699095, 1700596; one female [ex MSB], Socorro Co., NM, Sevilletta [sic] sand dunes, 20 April 1982, C. S. Crawford, X 1695695; one female [ex MSB], 17 km SW/S Bernardo, N 34.292 W 106.928, 17 July 1998, coll. S.L. Brantley, X 1698174; one male [ex MSB], Bernalillo Co., NM, Volcano Cliffs, Alb., 10 Aug 1979, C. S. Crawford, X 1570302; one female [ex MSB], NM, Bernalillo Co., Albuquerque, Aug. 2, 1974, B. Blake, X 0012317; one male, one female [ex NMSUAC], N. Mex., Doña Ana Co., Jornada Range, 15 m N Las Cruces, 24-vii-79, D. Richman, X 1699292, 1700581; one female [ex UNSM], TX: El Paso Co, red dunes on Montana Ave., 31.841° N 106.131° W, 1245 m, rolling AM after PM rains, 22-VIII-2010, M. J. Paulsen, X 1547783. Total paratypes: 120 Comments. Boreocanthon halffteri is very closely related to B. probus, from which it differs by a) distribution; b) anterolateral pronotal angles conspicuously explanate; c) more elongate parameres; d) rounded shape of paraocular areas; d) somewhat more acute, narrower and widely spaced clypeal teeth. Some specimens of the latter species from eastern New Mexico and adjacent west Texas counties are intermediate between B. halffteri and more “typical” B. probus. They could be considered reasonable grounds for regarding the two taxa as subspecies. However, what I know now suggests that B. halffteri, in addition to being morphologically distinct, is restricted to the Rio Grande drainage and geographically isolated from B. probus, which to me weaken an assumption that they are conspecific. The most proximate populations between the two species are that of B. halffteri in El Paso (El Paso County) and that of B. probus near Monahans (Ward County), Texas, separated by about 200 miles (Fig. 82, inset). Of note is the fact that red sands characterize the El Paso site while white sands predominate in those near Monahans. Viewed on Google Earth, the intervening east-west gap between about 31°N and 32° N latitude, where neither species is yet known to occur, appears largely devoid of dune habitat. Whether or not the disjunction is real will be tested by the results of future sampling in that area. Of note is the fact that B. ebenus occurs at both the El Paso and Monahans locations. Schoenly’s (1983) reference to B. melanus on the west slopes of the Hueco Mountains (El Paso County, 31°44ʹ N 106°04ʹ W) is likely assignable to this new species; however, his voucher specimens, collected along with B. ebenus, could not be located at Angelo State University, where Schoenly reported depositing them (Serina Brady, pers. comm.). Boreocanthon simplex (LeConte, 1857), restored generic combination Fig. 90–101 Canthon simplex LeConte 1857: 41. Canthon corvinus Harold 1868:129 (new synonymy per Horn 1870: 46). Canthon simplex var. militaris Horn 1870: 46 (new synonymy per Robinson 1948: 87). Canthon simplex var. humeralis Horn 1870: 46 (new synonymy per Robinson 1948: 87). Canthon simplex antiquus Pierce 1946: 120 (new synonymy per Miller et al. 1981: 626). Boreocanthon simplex (LeConte) (new combination per Halffter 1958: 209). Canthon (Boreocanthon) simplex (LeConte) (new combination per Howden 1966: 727). Boreocanthon simplex (LeConte), restored generic combination. Canthon bisignatus Balthasar 1939: 229, new synonymy. Canthon (Boreocanthon) bisignatus Balthasar 1939 (new combination per Halffter and Martínez 1977: 82). Type material. 1) Canthon simplex LeConte: syntype (male). Museum of Comparative Zoology, Cambridge, MCZ-ENT00003699. Examined by photograph (https://mczbase.mcz.harvard.edu/guid/MCZ:Ent:3699); 2) Canthon corvinus Harold: syntype (sex undermined), Museum für Naturkunde, Berlin. Examined by photograph courtesy F. V. Vaz-de-Mello; 3) Canthon simplex humeralis Horn: syntype? (female). Museum of Comparative Zoology, Cambridge, MCZ-ENT00008108. Examined by photograph (https://mczbase.mcz.harvard.edu/guid/ MCZ:Ent:8108); 4) Canthon simplex militaris Horn: syntype? (female). Museum of Comparative Zoology, Cambridge, MCZ-ENT00008109. Examined by photograph https://mczbase.mcz.harvard.edu/guid/MCZ:Ent:8109); 5) Canthon bisignatus Balthasar: syntypes (two, sex undetermined), National Museum, Prague. Examined by photograph courtesy Jiří Hájek. 6) Canthon simplex antiguus Pierce: holotype (sex undetermined). Los Angeles County Museum of Natural History, LACMIP 2594. Not examined. Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 49 Figures 90–99. Boreocanthon simplex (LeConte). 90) Habitus, anterolateral view. 91) Habitus, dorsal view. 92) Dorsal view of anterior portion of head. 93) Apical view of parameres. 94) Lateral view of genital capsule. 95) Dorsal view of base of left elytron. 96–97) Maximum (96) and minimum (97) expression of orange/red elytral humerus. 98) Anomalous orange elytral markings. 99) Lateral view of base of pronotum and elytron. 50 · October 14, 2022 Edmonds Type localities. 1) Canthon simplex LeConte: Oregon and California. 2) Canthon corvinus Harold: Sonora, California (see Comments); 3) Canthon simplex humeralis Horn: California, Coast Mountains south; 4) Canthon simplex militaris Horn: California, Tejon and Visalia; 5) Canthon bisignatus Balthasar: California; 6) Canthon simplex antiguus Pierce: Rancho La Brea, Tar Pits A and 81, Hancock Park, Los Angeles, California. Diagnosis. Head: Anterior one-third of clypeus (Fig. 92) finely rugose, weakly shiny, posterior portion and paraocular areas minutely punctured on dense shagreen background, microspots very weak. Clypeus (Fig. 92) bearing four rather weak marginal teeth, median pair short, rounded apically, separated by rounded emargination, lateral pair small, angulate apically. Paraocular notch subtle, sometimes almost effaced, setting off very small, angular corner of the paraocular area. Labio-gular fimbria broadly triangular, apex extending about onethird length of gula. Prothorax: Pronotum (Fig. 95–97) evenly covered by dense shagreen and widely spaced punctures, microspots faint, denser than punctures. Anterior pronotal angles very weakly upturned, posteromedian angle at most only very weakly depressed; circumnotal ridge smooth posterolaterally. Hypomeral carina usually absent or only feebly developed, occasionally distinct. Pterothorax: Mesoventrite with few weak punctures on shagreen background; metaventrite with evenly distributed, widely spaced, small punctures. Elytra (see also Comments below): Interstriae covered by dense shagreen, microspots small, faint, discal interstriae with few, very widely spaced micropunctures; 2nd and 3rd not usually swollen apically; anterior margin of some or all interstriae 2–5 bearing minute tubercle (sometimes obscured by pronotal margin). Humeral umbone dull black, sometime shiny (Fig. 99) or with orange/red spot (Fig. 96–97). Elytral striae weak, edges not sharply defined; lateral striae (7-9), especially anterior one-half, often effaced or nearly so, 8th stria never with any trace of carinula. Legs: Inner margin of protibia (Fig. 90) at most only gently widened at base of basal tooth. Protibial spur sexually dimorphic: strongly curved and apically acute in female, curved and apically bifurcate in male. Anterior margin of hind femora with conspicuous row of long setae, ventral surface punctate with few short setae. Abdomen: Pygidium evenly convex, uniformly dull; pygidium and 6th abdominal ventrite sculptured as elytra, remaining ventrites smooth. Genital capsule: Compressed distal portion of parameres broadly triangular (Fig. 94), ventral apical angles in form of slender, elongate knobs (Fig. 93). General: Black, rarely weakly shiny, rarely with green or blue undertone on head, pronotum; elytral umbone sometimes partially or completely orange or red. Length: 4.5 – 8.0 mm. Geographic distribution (Fig. 100–101). Widely distributed in United States west of the Rocky Mountains. Ecogeographic environment: Desert and Xeric Shrublands, Temperate Conifer Forests, and Mediterranean Forests, Woodlands and Scrub biomes. Specimens examined: 3196. Collection localities. Note. Red diamonds mark collection localities with at least one observed specimen bearing orange/red elytral umbones. CANADA — BRITISH COLUMBIA: Cariboo RD ●Chilicotin; East Kootenay RD ●Roosville [Aug] ●Kimberley [Jun] ●Cranbrook [May]; Columbia-Shuswap RD ●Salmon Arm [Apr]; Kootenay Boundary RD ●Rock Creek [Jul]; North Okanagan RD ●Vernon [Apr, Aug]; Thompson-Nicola RD ●Aspen Grove [Jul]. MEXICO — BAJA CALIFORNIA: Mpio. Ensenada ●47 km E San Telmo (road to Parque Nacional San Pedro Mártir), 950 m [Jun] ●Rancho La Bellota, 32° 09.411ʹ N 116° 27.680ʹ W, 1863 ft [May] ●Mikeʹs Sky Rancho, 4000 ft [Jun] ●San Vicente, 200 m [May] ●43 mi E Ensenada, Santa Catarina, Sierra de Juarez, 3600 ft [Aug] ●Ensenada [Jun] ●7 mi N Ensenada ●Santo Domingo [May] ●Valle de las Palmas [Nov] ●20 mi S Santo Tomás [Aug] ●Punta Banda [Jun] ●40 mi S Tecate. UNITED STATES — ARIZONA: Apache Co. ●Chuska Mountains, Wagon Wheel Forest Camp, 36.452° N 109.1704°W [Aug] ●White Mountains, Diamond Creek [Jun]; Coconino Co. ●Bear Canyon [Jun] ●Flagstaff [Sep] ●5 mi S Sedona, Red Rock Crossing Campground [Apr]; Cochise Co. ●Portal [Jul] ●Benson [Apr]; Gila Co. ●0.5 mi W Miami ●3 mi NE Gisela [Mar] ●13 mi S Globe [Aug] ●Globe [Apr, Jul] ●Lake Roosevelt ●base Pinal Mts., 4000 ft [May]; Graham Co. ●12 mi S Safford [May]; Maricopa Co. ●Tempe [Mar] ●Phoenix [Apr] ●Ft. McDowell [Apr-May] ●0.5 mi SE Sunflower [Apr] ●Rock Springs, Agua Fria River [Apr]; Mojave Co. ●Beaver Dam (near Littlefield), 36°53ʹ41ʺ N 113°55ʹ55ʺ W, 1916 ft [Apr]; Pima Co. ●Tucson [Mar-Apr, Jun] ●Santa Catalina Mountains [Jun]; Pinal Co. ●Superior [Mar] ●6 mi S Superior, 33.2162 N -111.0655 W, 990 m [Apr-Jul] ●3.5 mi W Superior, Boyce-Thompson Arboretum [Feb] ●Aravaipa Canyon [Apr]; Yavapai Co. ●Bumble Bee [May] ●12 mi S Camp Verde, 3000 ft [Jun] ●4 mi N Clarkdale ●4 mi S Sedona [Jun] ●34°13ʹ23ʺ N 112°05ʹ37ʺ W ●Hwy Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 51 Figures 100–101. Boreocanthon distribution. 100) Approximate distributions of Boreocanthon simplex (LeConte), B. lecontei (Harold) and B. coahuilensis (Howden). 101) Solid red diamonds mark California collection sites for B. simplex with red elytral humeri in one or more specimens; shaded area marks general limits of the Central Valley. 52 · October 14, 2022 Edmonds I-17, 2 mi S jct Hwy 179 [Apr] ●2.1 mi NW Moctezuma Well, Moctezuma Castle National Monument, 3984 ft [May] ●8 mi NE Yarnell [Jun] ●Prescott [Mar]. CALIFORNIA: Alameda Co. ♦Oakland [Apr, Jun]; Alpine Co. ●Monitor Pass 8300 ft [Jun, Aug] ●38.4504° N 119.5979° W, 8100 ft [Jun-Jul] ●Ebbettʹs Pass, 8950 ft [Jun] ●Bloods [Jul] ●Hope Valley, 7000ft [May]; Amador Co. ●El Dorado National Forest, Panther Creek Road [Jun]; Butte Co. ●Butte Meadows [Jun] ●Richardson Springs [May]; Calaveras Co. ●13 mi E Dorrington [Jun] ♦Mokelumne Hill [Jul] ●5 mi NE Tamarack [Jun]; El Dorado Co. ●Kyburz [Jul] ●Caldor [Jun] ♦Somerset [Jun] ♦13 mi E Georgetown, Blodgett Forest [Jun] ♦Pacific House [Sep] ●Ice House [May] ●3 mi N Wrights Lake [Jun] ●Camino [Jun] ●Twin Bridges, Pyramid Creek [Jun] ♦White Hall [Jun] ●Mt. Ralston [Aug] ♦Riverton [Jun-Jul] ●Placerville [Jun]; Fresno Co. ♦Squaw Valley [Apr, Aug] ♦6 mi S Shaver Lake [May] ♦Stevenson Creek, Sierra National Forest [May] ♦Camp Greeley, 2800 ft [Jun] ♦Huntington Lake, 7000 ft [Jul] ♦Kings Canyon National Park [Jun, Aug] ♦Lost Lake [Apr] ●Wishon Reservoir [Jul]; Humboldt Co. ♦Willow Creek [Jun]; Kern Co. ♦Tehachapi ♦Grapevine [Apr] ♦Lake Isabella [Jul] ♦Kelso Valley Road [Apr] ♦2.5 mi NW Canebrake [Apr] ●9 mi E Onyx ♦Onyx [May] ♦9 mi N Onyx ●Frazier Park [Apr] ♦Rancheria Creek, Walker Basin [Jun] ♦Ft. Tejon ♦Tejon Canyon [May] ♦Deer Creek [Apr] ●Lebec (Dutchmanʹs Camp) [Apr] ♦5 mi S Alta Sierra [Apr] ♦Weldon [Mar]; Lake Co. ♦Anderson Springs [May] ♦Bartlett Springs [Jun]; Lassen Co. ●Hallelujah Junction [Jul] ●Norval Flats, 5500 ft [May, Jul] ●Hayden Hill [Aug]; Los Angeles Co. ●Los Angeles [May] ♦Pasadena [Feb Apr-May] ●Claremont ●Big Tijunga Canyon [May] ♦Fairmont [Apr] ●Azusa [May] ●Chilao Recreational Area, 5300 ft [May]; Madera Co. ♦Oakhurst [May] ♦Coarsegold [May] ♦Bass Lake, 5000 ft [Jun-Jul] ●Sugar Pine ♦Southfork [Jul]; Marin Co. ♦Fairfax [Jul] ♦Stinson Beach [Apr] ♦Mill Valley [Aug] ♦Inverness [Aug]; Mariposa Co. ●Yosemite Valley [Jun] ●Glacier Point, 7214 ft ♦Fish Camp [May] ♦Miami [Mountain] Ranger Station, 37°25ʹ09.62ʺ N 119°44ʹ41.15ʺ W [May]; Mendocino Co. ♦Yorkville [May–Jun] ♦Jumpoff Creek [Aug] ♦2 mi NW Philo, Navarro River [Jul] ♦Newport; Modoc Co. ♦Hackamore [Apr] ●Warner Mountains, near Lost Lake, 8000 ft [May-Aug]; Mono Co. ●2 mi E Tioga Pass, 9500 ft [Jun] ●Mammoth Lake, 8000 ft [May-Jun] ●Hwy 108, Sonora Pass, 9700 ft [Jun-Jul] ●Saddleback Lake, 10000 ft [Jul] ●Bodie, 8500 ft [Aug]; Monterey Co. ●Stone Canyon [Apr] ●Paraiso Springs [Jun] ●Tassajara Hot Springs [May] ●Bradley [Apr] ●Carmel Valley [Jul] ●Bryson [Apr] ●Pacific Grove [Jun]; Napa Co. ♦Napa [JunJul] ♦Mt. St Helena [May-Jun] ●Deer Park [Jul]; Nevada Co. ♦Grass Valley [May] ●5 mi N Hobart Mills [Jun] ●Sagehen Creek [Jun] ●Truckee, 5800 ft [Jun, Aug]; Orange Co. ●Anaheim [Jun] ●Laguna Beach [Jun] ●Temple Hills [Mar] ●Laguna Beach [Jun] ●Laguna Niguel [Jun]; Placer Co. ●Tahoe City [Jul] ●Morongo Valley [May] ♦Sugar Pine [Aug] ●Cisco [Jun] ●Colfax [Jun]; Plumas Co. ●Quincy ●Lake Almanor [Jun] ♦Chester [Jul] ♦Meadow Valley, 6000 ft [Jul]; Riverside Co. ●Cabazon [Apr] ●Riverside [Mar] ●Aguanga [May] ●0.5 mi SE Aguanga [Apr] ●8 mi SE Hemet, Bautista Canyon, 33°41ʹ N 116°50ʹ W, 2300 ft [Apr] ●Coahuilula Creek [May] ●6 mi E Temecula [Mar] ●San Jacinto [Mar] ●Idyllwild [Jun-Jul] ●Palm Springs [May] ●5 mi W Sage [Jul] ●Gavilan Hills [Apr] ●Goff Flats Campground, San Jacinto Mountains [Jul]; San Benito Co. ♦Pinnacles National Monument [Apr, Jun]; San Bernadino Co. ●2.9 mi N Cedar Springs [Jul] ●0.9 mi NE Cedar Springs ●Colton [Mar] ●San Bernadino [May] ●Glenwood Springs [Aug] ●Deep Creek Campground [Jun] ●Big Bear Lake [May] ●Fawnskin [Jul]; San Diego Co. ●Campo [May] ●Scissors Crossing [Apr] ●10 mi N Scissors Crossing [Jun] ●Lake Cuyamaca [Apr] ●Pamo Valley [Jun] ●Indian Flats Campground, Warner Springs [Jun-Jul] ●Dulzura [Apr] ●Boulevard [Jul] ●Mason Valley [Jun] ●Poway, 500 ft [Apr] ●Lake Moreno [Mar] ●Bonsall, Gopher Canyon Rd. [May] ●Camp Pendleton, 5 mi E I-5, Mateo Creek ●6 mi E Banner [Jul] ●San Felipe Valley, 3000 ft [May] ●Murray Dam [May] ● Mt. Palomar [Jun] (See also McPeak et al. 2014 for additional San Diego Co. records.); San Francisco Co. ♦San Francisco; Santa Barbara Co. ●Clear Creek, Cuyama Canyon [Mar] ●Santa Maria [Mar] ●Buellton [May] ●Santa Barbara ●10 mi E Santa Maria, Cuyama River [Jul]; Santa Clara Co. ♦30 mi SE Livermore (Alameda Co.) [Jun]; Santa Cruz Co. ♦Ben Lomond [Jun] ♦Santa Cruz [Apr]; Shasta Co. ♦Cayton [Jul] ●Old Station [May-Jun] ♦Hat Creek (May-Jun] ●Shingletown [Jun]; Sierra Co. ●Sierraville [Aug] ●10 mi S Sierraville [Jun]; Siskiyou Co. ●Yreka ●Walker [Jun] ●McCloud [Jun] ●Bartle [Jul] ●Cole [Jul] ●Klamath River [May] ♦Taylor Lake, 5600 ft [Aug] ●Castle Lake, 5200 ft [Jul] ●Burney Valley [Aug] ●Mt. Hebron [Aug]; Sonoma Co. ♦Mark West Springs [May] ●Mt. St Helena [Jun] ♦Rio Nido [Aug]; Tehama Co. ●South Fork Battle Creek [Aug]; Trinity Co. ♦Carrville, 2500 ft [Jun] ♦Zenia [Jun] ●Plummer Spring [Jun] ♦Hayfork [May] ●Mountain Meadows Ranch, head Coffee Creek, 5100 ft [Jul]; Tulare Co. ♦Kaweah ♦California Hot Springs [Jun-Jul] ●Sequoia National Park, Kings River Canyon [May] ♦2 mi E Johnsondale ●Shake Camp Campground [Jun] ♦Porterville [May] ♦Springville; Tuolumne Co. ●Tuolumne Meadows ●Sonora Pass, 38.3281° -119.6371°, 2950 m ●Spring Gap [Jun] Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 53 ♦Strawberry [Aug] ♦Twain Harte [Mar] ♦17 mi S Sonora [May] ♦Pinecrest [Jun-Jul] ♦Hardin Flats [May] ♦6 mi E Groveland, 2800 ft [Jul]; Ventura Co. ●Lake Piru [Jun] ●Ventura [May] ●Sespe Canyon [Apr] ●Santa Paula [Nov]; Yolo Co. ♦Rumsey [May]; Yuba Co. ●Strawberry Valley [Sep]. COLORADO: Boulder Co. ●8mi SW Boulder, 8400 ft [Jul]; ●Ward ●Gold Hill [Jul]; El Paso Co. ●Cascade [Jul]; Fremont Co. ●Coal Creek [May]; Garfield Co. ●Glenwood Springs [Jul]; Gunnison Co. ●8 mi NE Almont, One Mile Canyon, 8200 ft [Jul] Jackson Co. ●Walden [Aug]; Larimer Co. ●Estes Park, 40°21ʹ11ʺN 105°28ʹ45ʺ W, 2500 m [Aug] ●Red Feather Lake [Jun]; Mineral Co. ●Creede [Jul]; Moffat Co. ●10 mi W Maybell [Jun]; Ouray Co. ●Ouray [Jul]; Routt Co. ●Steamboat Springs [Jun]; Teller Co. ●Florissant Canyon, 8500 ft [Jul]. IDAHO: Ada Co. ●Boise, 2690 ft [May]; Adams Co. ●Packer Johns Cabin State Park [Jun] ●6 mi E New Meadows [Jun]; Bear Lake Co. ●Montpelier ●Georgetown [May] ●Paris [May]; Bannock Co. ●Pocatello [May, Jul] ●Lava Hot Springs; Benewah Co. ●St. Maries [Aug]; Blaine Co. ●Prairie Creek [Jul] ●Frenchman Creek, Hwy 75 below Galena Summit ●10 mi S Bellevue [May] ●Clarendon Hot Springs [Jul]; Boise Co. ●Thorn Creek [Jul] ●Gardena [Jun] ●8.5 mi NE Gardena [Jun]; Bonner Co. ●Sagle [Jun] ●Sandpoint, 2085 ft [May-Jul] ●Priest River Experimental Forest [Jun] ●Granite [Jun]; Bonneville Co. ●Grayʹs Lake [Apr] ●Willow Creek [Jun]; Butte Co. ●Craters of the Moon National Monument, Little Cottonwood Creek [Jun-Jul]●6 mi S Howe [May]; Caribou Co. ●Diamond Point [Aug] ●6 mi SW Soda Springs [Apr] ●3 mi E Wayan [Jun]; Clark Co. ●Dubois [Jun] ●Camp Creek, 5 mi S Humphrey [Jul]; Clearwater Co. ●Elk River [May] ●1 mi N Elk River [May]; Custer Co. ●Double Springs Ranch [Jun]; Elmore Co. ●11 mi S Pine [Jun] ●8.5 mi E Featherville [Jun] ●Little Camas Reservoir [Jun]; Fremont Co. ●St Anthony [Jul] ●Coal Creek [May]; Gem Co. ●Sweet [Jul]; Kootenai Co. ●Worley [May] ●Chilco [May] ●Coeur dʹAlene, 2150 ft [May-Jun] ●6 mi W Athol [Jun]; Latah Co. ●Moscow, 2560 ft [Apr, Jun-Jul] ●Bovill [Apr] ●5 mi N Troy [May] ●Spring Valley Reservoir [Apr, Jul]; Lewis Co. ●Winchester [May]; Lincoln Co. ●Shoshone [Jul]; Madison Co. ●Driggs [Jun]; Nez Perce Co. ●Gifford [May]; Oneida Co. ●Sublette Canyon [Apr]; Owyhee Co. ●17 mi W Silver City; Shoshone Co. ●Wallace [Jun, Aug] ●3.4 mi N Clarkia; Teton Co. ●Tetonia [Jul]; Twin Falls Co. ●Timber Canyon [Jul] ●Steamboat Rock [Jun]; Valley Co. ●Bear Valley [Jul] ●Cascade [May] ●Donnelly [May]; Washington Co. ●7 mi NW Midvale [Apr]. MONTANA: Beaverhead Co. ●Centennial Valley [May] ●Dillon [Jul] ●Birch Creek [Aug]; Broadwater Co. ●Limestone Hills (near Townsend), 46°17.54ʹ N 111°36.45ʹ W; Flathead Co. ●Big Creek, Flathead National Forest ●Glacier National Park, Colby Meadow [Jun-Sep]; Gallatin Co. ●7 mi S Logan, Madison Buffalo Jump State Park [May] ●Willow Creek [Jun]; Hill Co. ●Beaver Creek, 6300 ft; Lake Co. ●Ronan [Jul-Aug]; Lewis and Clark Co. ●Helena [Jul]; Madison Co ●9 mi S Alder, Ruby Reservoir [Jul]; Meagher Co. ●Ringling [Apr]; Missoula Co. ●Salmon Lake, 7.5 mi N jct. hwys 200 & 83 [Jul]; Park Co. ●Crazy Mountains, 5000 ft [Jun]; Ravalli Co. ●Hamilton [Jun] ●Camp Creek [Jul] ●Gird Creek [Jul] ●Flathead Lake [May] ●Darby [Aug]; Sanders Co. ●Thompson Falls [Jun]; Sheridan Co. ●Sheridan [Aug]. NEVADA: Clark Co. ● 10 mi N Paradise [Jun] ● Elko Co. ●Elko [Jul] ●Wildhorse Crossing State Park [Sep] ●Jarbidge [Jul]; Lander Co. ●18 mi S Austin [Jun]. NEW MEXICO: Catron Co. ●Luna [Apr]; Sandoval Co. ●Valles Caldera National Preserve, Valle Grande, 35°53ʹ N 106°30ʹ W [Jul]. OREGON: Baker Co. ●Whitman National Forest, Eagle Mountains, Skookum Creek ●Pine Creek, Blue Mts, 4000 ft [Aug] ●Baker [Apr, Jul]; Benton Co. ●Monroe [Jun] ●Corvallis [May-Aug] ●McDonald Forest, Oak Creek [Apr]; Deschutes Co. ●16 mi SE LaPine [May] ●7 mi NW LaPine, Pringle Butte, 4000 ft [Apr] ●Paulina Creek [Aug]; Douglas Co. ●Riddle [May]; Grant Co. ●Graylock Butte [Jul] ●Dixie Pass, 5200 ft ●Silvies [Jun]; Harney Co. ●Steens Mountain, 6500 ft [Jun-Jul] ●Hwy 20, mile marker 156, near Buchanan [Mar-Jun] ●Steens Mt., above Fish Lake, 8000-9000 ft [Jun]; Hood River Co. ●4 mi W Oak Grove [May-Jun]; Jackson Co. ●Medford [Apr-May] ●Colestin [Jul] ●Gold Hill [Apr]; Jefferson Co. ●0.3 mi E Horseshoe Lake [Jul-Aug] ●Santiam Pass, 5000 ft [Jun]; Josephine Co. ●Grants Pass [Jul]; Klamath Co. ●Worden [Jul] ●Collier State Park [Jul] ●Klamath Lake [Jun] ♦Fremont National Forest [Jun] ●Crater Lake [Jul] ● Bly, Meryl Lake [Jun] ●7 mi NE Bly, Deming Creek [Jul] ●8 mi N Bly, Preacher Creek [May] ●Spencer Creek [Jun] ♦Algoma [Apr]; Lake Co. ●15 mi E Lakeview [Jun] ♦Warner Canyon, near Lakeview [Jun-Jul] ●12 mi SW Silver Lake [Aug]; Lane Co. ●10 mi NNW Junction City [Sep] ●11 mi NE Blue River, H.J. Andrews Experimental Forest [Jun]; Marion Co. ●Pratum; Multnomah Co. ●Portland [Jun]; Umatilla Co. ●Tollgate [Jun] ●Meacham, 4300 ft [May-Jun] ●Cold Springs [Jun]; Union Co. ●28 mi SE Union 4280 ft [Jul] ●La Grande [Jun] ●0.2 mi N Rd. 77 on Rd. 600, 5550 ft [Jun] ●Kamela [Jun]; Wallowa Co. ● Hwy 3 at Day Ridge Road, 25 mi N Enterprise, 4700 ft [Sep] ●Zumwaldt Prairie Reserve [Jun] ●Blue Creek, 4170 ft; Wasco Co. ●16.5 mi E Hwy 35 on Hwy 44, 2800 ft [Jun-Jul]; ●6.4 mi E Rd. 1720 on Upper Fivemile Rd.; 14.8 mi E Hwy 35 on Hwy 44, Hood River National Forest [Apr] ●10.1 mi W Dufur Road on Hwy 44, 45°24.882ʹ N 54 · October 14, 2022 Edmonds 121°19.524ʹ W, 2732 ft [May] ●Mt. Hood National Forest, Clear Creek Campground [Jun]; Washington Co. ●Forest Grove [May]; Wheeler Co. ●Walton Lake [Jun]; Yamhill Co. ●McMinnville [Apr-May]. UTAH: Box Elder Co. ●Rosevere Creek, Raft River Mountains [Jun]; Cache Co. ●Logan Canyon, 7200 ft [May] ●Bear Lake, summit Logan Canyon [Jun] ●Tony Grove [Jul] ●Sardine Canyon ●Wellsville Canyon [Jun]; Carbon Co. ●8 mi NE Castlegate [Jun] ●Scofield Reservoir [Jun]; Duchesne Co. ●Hanna [Jul]; Juab Co. ●Nephi Canyon [Jun] ●Chicken Creek Campground, 5 mi E Levan [Sep]; Morgan Co. ●Cottonwood Canyon, Cache National Forest [Jul] ●Lost Creek, above Lost Creek Reservoir [Jul]; Rich Co. ●Logan Canyon [May-Jun] ●Birch Creek Reservoir [Jul] ●Otter Creek Spring, 12 mi W Randolph [Jun] ●Laketown Canyon; Salt Lake Co. ●Little Cottonwood Canyon, Wasatch Mountains [Jul]; Sanpete Co. ●Boulger Lake, Wasatch Plateau [Jun] ●Indianola [May]; San Juan Co. ●Blanding [Jul] ●Ute Mountains (Colorado state line) [Jun]; Tooele Co. ●Oquirrh Mountains, Settlement Creek Canyon [Jul]; Utah Co. ●Timpanogos, Wasatch Mountains [Jun] ●Aspen Grove, 7000 ft [May, Aug] ●Provo [Apr, Sep]●Provo Canyon [Sep] ●American Fork Canyon, 9500 ft [Aug] ●Payson Canyon [Jul] ●Payson Lakes [Jun] ●Sundance [May] ●Timpanogos Lodge [May] ●Squaw Peak [Aug] ●Alpine [May]; Wasatch Co. ●Strawberry Reservoir [Jun-Jul] ●Provo River [Jun] ●Silver Lake [Jul]; Wasatch Mt. State Park [May] ●Cascade Springs [AprMay] ●Daniels Canyon, 8000 ft [Jul]; Washington Co. ●St. George ●Leeds Canyon [Apr] ●Lytle Preserve, 37.14490° N 114.02283° W, 836 m [Sep] ●Lytle Ranch, Welcome Spring [Jun] ●Snow Canyon Dunes [Mar-Apr] ●Red Cliffs Recreation Area, Red Cliffs Campground [Apr] ●Toquerville. WASHINGTON: Asotin Co. ●8 mi SE Clearwater Ranger Station (near Mt. Misery); Benton Co. ●Prosser [May]; Chelan Co. ●Riverhead Campground, Wenatchee National Forest [Jun]; Ferry Co. ●10 Mile Creek Campground, 48.515 -118.739, 1800 ft [Jun]; Klickitat Co. ● Spearfish [Jul] ●Glenwood, 0.2 mi N jct. Ladiges Rd on Bird Creek Road., 46°01.955ʹ N 121°18.255ʹ W, 1993 ft [May]; Kittitas Co. ●13 mi NE Kittitas [May] ●Easton ●Lake Cle Elum [Jun]; Pend Oreille Co. ●5.8 mi SW Tiger, Nile Lake, 3400 ft [Aug] ●Leo Lake, 3150 ft [Jun]; Snohomish Co. ●Lewis Peak, Blue Mts. [Jun]; Spokane Co. ●Mt. Spokane, 5000-5800 ft; Stevens Co. Northport [Jun] ●Pleasant Valley [May, Oct]; Walla Walla Co. ●Lowden [May] ●Walla Walla [Apr-May]; Whitman Co. ●Pullman [Mar-May, Jul] ●Palouse, Kamiak Butte [AprMay]; Yakima Co. ●Bird Creek Ford, Yakima Indian Reservation, 2500 ft [Jun] ●5 mi W, 2 mi N Wenas Lake ●Toppenish [May]. WYOMING: Albany Co. ●5 mi W Garrett [Jul] ●Medicine Bow National Forest, Curtis Gulch Campground [Jul]; Big Horn Co. ●Trout Creek [Jun]; Fremont Co. ●Sweetwater Station [Jun]; Natrona Co. ●Bates Creek, 22 mi S Casper, 7000 ft [Aug]; Park Co. ●Lake Creek Camp, 13 mi SE Cooke City, MT, 7200 ft [Jul]; Sublette Co. ●Fremont Lake [Jul] ●Half Moon Lake [Jun]; Sweetwater Co. ●Burntfork [Jun] ●Green River [Jul]; Teton Co. ●43.61735° N 110.42963° W, 7300 ft [Aug] ●Yellowstone National Park, Lamar Valley [May] ● Yellowstone National Park, Trumpeter Swan Lake, 6080 ft [Sep] ●Jackson Lake, Teton National Park [May] ●Jackson Hole, 6300 ft [Jul, Sep] ●Jenny Lake [Jun]; Washakie Co. ●12.5 mi ESE Ten Sleep, Ten Sleep Preserve (The Nature Conservancy) [Jun]. Comments. There is potential confusion regarding the date of publication (1857 or 1860) of this LeConte species. Moore (1986) pointed out that the various parts of the Steven’s survey were published hastily and piecemeal and only later grouped and formally published in book form in 1860 — making it, in Moore’s words, a “collater’s nightmare.” LeConte, and perhaps other contributors, evidently opted to commission a private “preprint” of his results, which he distributed to colleagues years in advance of the issue of the complete report. I have not seen a copy of LeConte’s “preprint” but, in the light of Moore’s commentary and the fact that the 1857/1860 duality is of no nomenclatorial consequence, I accept the long-standing reported date of publication as 1857. Thanks to Dr Jiří Hájek of the Czech National Museum, Prague, I have examined photographs of the two syntypes there of Canthon bisignatus Balthasar. In his study of Canthon, Balthasar (1939) considered eleven species here assigned to Boreocanthon, including B. simplex and a new species, Canthon bisignatus. The latter he separated from the remaining species, including B. simplex, in couplet 77(232) of his identification key (p. 190), [“Clypeus bidentate, lateral teeth either completely absent or indicated at most only by a weakly angulate thickening of the clypeal margin.“], terminating at couplet 146(147). Both syntypes are clearly highly worn individuals of B. simplex whose originally quadridentate clypei have been rendered weakly bidentate by severe abrasion; moreover, both have bright red/orange elytral umbones. One bears the label, Canthon simplex humeralis Horn, Det. B. D. Gill ’96. Since, to my knowledge, the synonymy of Canthon bisignatus Balthasar with Boreocanthon simplex has not heretofore been established formally, I do so here. Taxonomic review of the dung beetle genus Boreocanthon Insecta Mundi 0952 · 55 In addition to the syntype for Canthon simplex LeConte cited above (labeled by blue disk as Oregon/ Washington), there are two additional likely syntypes also present in the LeConte Collection at the Museum of Comparative Zoology (Crystal Maier, pers. comm). These are labeled (brown disk) as “Russian America”, a possible reference to Fort Ross in present day Sonoma County, or other colonial outpost in California operated by the Russians until 1841 (Ballard 1997). Potential syntypes of Horn’s (1870) varieties humeralis and militaris also reside in the MCZ collection. But in each case, the specimen depicted in the MCZ database (see photographs referenced above) as putative type material was, for an unknown reason, drawn from the LeConte Collection (Crystal Maier, pers. comm.) and, therefore, not strictly attributable to Horn. For this reason, I have added “?” to the syntype designations above. Canthon simplex antiquus was one of two fossil taxa described by Pierce (1946) from asphalt deposits at Rancho La Brea in Los Angeles, California. It, along with C. praticola vetustus Pierce, are both considered conspecific with modern species (Miller et al. 1981; see Comments for B. praticola). Horn (1870) recognized four varieties of B. simplex, viz., simplex, corvinus, militaris and humeralis. The second, corvinus, was in reference to Harold’s (1860) species, which he explicitly rejected as deserving species status (“I cannot agree with Harold in separating corvinus as distinct.”). Similarly, it is reasonable to assume that his “var. simplex” was a direct reference to LeConte’s species. Only the remaining two varieties, var. humeralis and var. militaris, are here considered Horn taxa and, in accordance with ICNZ Article 45.6.4, are to be treated as subspecific names. Horn’s (1870: 46) estimation of his varieties was blunt: “My own series is large, and my observation in California leads me to consider all these forms as mere local varieties and not distinct species.” Robinson (1948: 88) later came to a similar conclusion, although he went farther by rejecting Horn’s notion of local varieties altogether: “I think simplex is just a very variable species without any correlation between the internal and external variations and geographic localities, and therefore see no reason for continuing to split it into several varieties.” I agree with Robinson’s viewpoint. I am inclined to believe that my taxonomic view of this species could be too simplistic — Boreocanthon simplex is anything but “simplex.” As for morphological features, its only real constant is the shape of the head margin, with reduced clypeal teeth and weak paraocular notch (Fig. 92). Most specimens also show weakly expressed lateral elytral striae (7th and 8th), which, especially viewed obliquely from the side (Fig. 95), appear almost effaced basally. A common variable is the conspicuousness (“crispness”) of pronotal puncturing, which varies continuously from nearly effaced to clearly expressed; the latter state figures in Harold’s definition of his “var. corvinus.” The elytral umbones, while dull black (Fig. 95) over most of its range, can be colored (Fig. 96–97, see below). I have not been able to sort these variables and others in any useful taxonomic way, and thus I, like previous observers (see below), have concluded that it is but a very common, widely distributed, highly variable species. A future student of the group might find it to be a complex array of populations with greater or lesser connections inter se — in other words, with other data (e.g., molecular) it might be possible to delimit multiple species, or semispecies or subspecies. Halffter (1958) fittingly recognized B. simplex as the most variable species in the genus and suggested that it could represent a “… [a Rassenkreiss or polytypic species with geographic subspecies … or {several} well-defined species]”. Perhaps the best known manifestation of variation in B. simplex is the color and luster of the elytral umbones. The most common condition throughout its range is the combination of dull, black umbones (Fig. 90, 95). Umbone luster varies irregularly but can be weakly shiny in contrast to the dull surrounding elytral surface in certain populations (Fig. 99). In some places, preponderantly in the mountainous zones surrounding the Central Valley of California (see Fig. 101 and collection localities listed above marked with red diamond ♦) a portion of the population bears umbones marked by an orange (reddish/red) spot. In most of these places, the colored umbones vary in size and intensity and occur along with all-black specimens. The types of the “varieties” militaris, humeralis and corvinus, as well as that of Canthon bisignatus, all bear orange markings, but only in the case of militaris are they cited in Horn’s formal descriptions (1870:46). However, Harold (1868) does say, in his comments on corvinus, “[The number of individuals with red umbones is almost more than the solid black form.]” Moreover, he cites “Sorona, California” as type locality for his species, which I regard as a typographical error meaning Sonora, California (Tuolumne Co.). Individuals with orange umbones almost always occur with black individuals and are absent outside of California and southern Oregon. Among the 3000+ specimens examined for this study, I have seen only one outlier specimen with orange umbones; it is labeled LaGrande (Union Co.) 62 · October 14, 2022 Edmonds Blackwelder RE. 1944. Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 2. 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