Review of the Nearctic genus Lacconotus LeConte (Coleoptera, Mycteridae, Eurypinae)

Abstract Lacconotus LeConte, the sole Nearctic representative of the eurypine Mycteridae, is revised, based on morphological features of adults. The following syn. n. is proposed: Lacconotus pallidus Van Dyke, 1928 = Lacconotus pinicola Horn, 1879. The former is a light-colored form with a southern California distribution. A subgen. n., Alcconotus, is described for Lacconotus pinicola, producing the following comb. n.: Lacconotus (Alcconotus) pinicola (Horn). A lectotype is designated for Lacconotus pinicola. A key separating the two subgenera and species is provided, as are photographs and illustrations of salient structures of adults, and maps showing collection localities. Lacconotus punctatus is newly recorded in Alabama, Arkansas, Massachusetts, Oklahoma, Texas, and Wisconsin; Lacconotus pinicola is newly recorded in Arizona and Utah in the USA, and Baja California Norte in Mexico. Phenology information shows a north-to-south gradation in occurrence time.


Introduction
Among the three subfamilies of Mycteridae, the Eurypinae (= Lacconotinae) are the most diverse with 26 genera and 160 species recognized worldwide, the greatest diver-

Methods and conventions
Standard taxonomic methods were used in this study. Habitus photographs were taken with a Nikon Coolpix 5000 ® digital camera fitted to a Leica MZ95 stereoscope. Approximately 30 separate photographs were taken for each specimen/structure; these were imported into Combine ZP (Hadley 2010), which stacked and aligned the individual images to create a final photograph completely in focus.
Several measurements were used: HL = length of head from anterior margin of pronotum to labrum; PL = length of pronotum along middle; EL = length of elytron from anterior to posterior extent; GHW = maximum width of head, across eyes; GPW = maximum pronotal width; GEW = maximum width of both elytra; TL = HL + PL + EL.
Label data on type specimens are recorded verbatim, with all label data enclosed in quotes and individual labels separated by a slash (/). Information added by the authors for clarity is enclosed in square brackets ([ ]
Head relatively short, narrowed slightly posterior of eyes; eyes moderately large, distinctly convex, inner margins convergent anteriorly; facets moderately coarse, with intrafacetal setae (especially noticeable in Alcconotus); frontoclypeal suture indistinct, indicated by slight furrow only; antennal insertions slightly concealed dorsally by raised lateral margins of frons; labrum transverse, anterior margin straight to shallowly sinuate.
Elytra elongate, subovate, disc flat (Lacconotus) to slightly convex (Alcconotus), upper surface uniformly and moderately coarsely punctate and setose (slightly more coarse in Lacconotus), setae closely appressed to elytral surface; apical elytral patch present, but not conspicuous dorsally, not contrasting in color with respect to remainder of elytron; epipleuron narrow, traceable only to abdominal ventrite 3 or 4; mesosternum with posterior intercoxal process parallel-sided, extended posteriorly to near pos-  terior extent of mesocoxae; mesocoxae narrowly but completely separated, trochantins exposed; coxal cavities partly closed laterally by mesepimera; metasternum large, convex, anterior margin with indistinct (Lacconotus) to distinct (Alcconotus) process, in contact with posterior mesosternal process; median impressed line distinct to at least half distance to anterior margin of metasternum; metendosternite with long, relatively wide stalk; anterior tendons inserted on anterior margin of metendosternite body; laminae large, produced and somewhat angulate laterally.
Legs well developed, similar in relative shape and size on all thoracic segments; middle and hind femora slightly more expanded than front femora; femora relatively slender, but distinctly widened toward midlength; tibiae straight, about same length  as femora, tibial spurs very short, equal in length; tarsomeres slender, 5-5-4; all tarsomeres simple ventrally, except for penultimate tarsomere with large ventral lobe; basal tarsomere on hind tarsus subequal in length to other tarsomeres combined; tarsal claws with large blunt tooth.
The new records above make clear that the distribution of L. punctatus in North America is much wider than previously known (Fig. 15). Less than a decade ago, Pollock (2002) reported the species from only four jurisdictions in North America (Québec, Pennsylvania, Ohio, and Michigan). There are now records from 14 states and provinces on the continent. The records from Wisconsin constitute a northwestern range extension of 650 km; and those from Oklahoma a southwestern range extension of 1,000 km. The present distribution indicates that L. punctatus is found over much of eastern North America, from a latitude of 33.6° to 44.3°N, and between longitudes of 64.5° and 95.3°W, much of the continent west of the prairies.
There is also much more information on the range of habitats that L. punctatus occupies. In Nova Scotia a specimen was found in a mixed forest of white pine (Pinus strobus L.), balsam fir (Abies balsamea (L.) Mill), eastern hemlock (Tsuga canadensis (L.) Carr.), and maple (Acer spp.) (Majka and Selig 2006). In New Hampshire, W.J. Morse and D.S. Chandler collected 26 specimens at a water tower in a mixed hardwood forest with eastern hemlocks. In Oklahoma a specimen was collected on a dead oak and in Wisconsin a specimen was found in an oak savanna. In Arkansas a specimen was found in a mixed forest/old field.
The phenology information that is available (Fig. 16) indicates that adults can be found between 31 March and 16 June. Specimens from southern areas (i.e., Georgia) were found in mid April (Ulyshen et al. 2010), whereas those from northern latitudes (i.e., New Hampshire) occur mainly during the last two weeks of May and first week 1 Although listed from Ontario in Campbell (1991), we have not been able to find any published record, or any vouchers specimen in any North American collection that would substantiate this report. Consequently, pending verification of its occurrence in this jurisdiction, we remove Ontario from the known distribution of this species. 2 In addition to the specimen from Montreal in the CUIC (Appendix 1) a second specimen from Québec was formerly in the Ouellet-Robert collection of the Université de Montréal, however, the specimen was stolen before being databased so its collection date and locality in the province are unknown (pers. com., Louise Cloutier).
of June (D.S. Chandler, pers. comm.), indicating a north-to-south gradation in occurrence period.
Derivation of name. a partial anagram of Lacconotus, in which its species was formerly placed.
Taxonomic notes. Pollock (2002) stated that it might be necessary to establish a new genus for the two western species of Lacconotus, although no details were given to justify this possibility. There are many differences between the eastern and western  species of Lacconotus (see description above, for Lacconotus), but the most significant reason for proposal of this new subgeneric name is the structure of the male sex patch, which differs greatly between L. punctatus and L. pinicola. It could be argued that this might justify separation into two genera; however, within the related genus Mycterus Clairville (Mycterinae) there are also significant differences in this structure. Also, there are significant differences in the structure of the bursa copulatrix and spermatheca between the two groups (see ; it is impossible to compare these intrageneric differences with other eurypine or mycterid taxa, since the internal female genitalia have yet to be studied in detail in most groups. Description. See description above, for Lacconotus; characteristics unique to Alcconotus are indicated with the alternatives for Lacconotus (s. str.).  1902: 297;Woodworth 1913: 194;Seidlitz 1917: 99;Leng 1920: 240;Blair 1928: 33;Van Dyke 1928: 257;Spilman 1951: 50, fig. 15;Hatch 1965: 88;Arnett 1983: 3;Campbell 1991: 267;Poole and Gentili 1996: 315;Pollock 2002: 532. Lacconotus pallidus Van Dyke 1928Fall 1901: 32 (as Lacconotus pinicola), 177; Leng and Mutchler 1933: 36;Spilman 1951: 50;Crowson and de Viedma 1964;Pollock 2002: 531 (fig . 2.112 Diagnosis. Lacconotus (Alcconotus) pinicola may be distinguished from L. punctatus by the following features: body color ranging from testaceous to dark brown, uniform dorsally (Figs 2-3); antennae relatively long, subserrate; male sex patch on ventrite 2 small, oval, densely pubescent (Fig. 6); distribution in western North America (Fig. 15).

Lacconotus (Alcconotus) pinicola
Re-description. (see Horn 1879 and Van Dyke 1928) -With general features of subgenus Alcconotus (as described above) with the following: TL 4.8-7.5 mm; GEW 1.5-2.3 mm; TL/GEW 2.8-3.4. Dorsal body surface uniformly testaceous to dark brown or piceous (Figs 2-3), without any color contrast; antennomeres 5-10 relatively elongate, subserrate; antennal sensilla not completely surrounding opening of antennomere, restricted to triangular side of antennomeres; wing membrane distinctly pigmented, veins very conspicuous (Fig. 8); male sex patch longitudinally oval, occupying about 2/3 length of ventrite 2, densely covered with short setae (Fig. 6), not bulbous or contrasting in color; tegmen of male genitalia (Fig. 11) short, parameres stout; bursa copulatrix ( Fig. 14) very large, spherical. Notes. Van Dyke (1928) established L. pallidus (as distinct from L. pinicola) based on its lighter color, relatively narrower pronotum, shorter relative length of the elytra, and deeper punctation. However, upon examination of the type series and other specimens, we have determined that the only feature of Van Dyke's that withstands scrutiny is the habitus color. As well, more detailed examination has revealed that the male and female genitalia and male sex patch are virtually identical between L. pinicola and L. pallidus. One feature, mentioned by Van Dyke (1928) that does seem noteworthy is the somewhat restricted distribution of L. pallidus in southern California. We herein consider L. pallidus a pale "form" of L. pinicola. Lacconotus (Alcconotus) pinicola is newly recorded from Arizona, Utah, and Baja California Norte in Mexico (see Appendix A). Published records of L. pinicola are from British Columbia (Hatch 1965), California (Fall 1901;Van Dyke 1928), Colorado (Horn 1879;Cockerell 1893;Van Dyke 1928), western Nevada (Horn 1879), and New Mexico (Snow 1882(Snow , 1906Knaus 1907). The range of the species (Fig. 15) shows it to be widely distributed in the southwestern United States (Arizona, California, Colorado, Nevada, New Mexico, and Utah) extending south to Baja California Norte in Mexico, and in southeastern British Columbia. Specimens should be sought in intervening areas in Idaho, Oregon, and Washington to ascertain if these populations are actually disjunct.
A number of specimens examined were found on ponderosa pine (Pinus ponderosa Douglas ex. C. Lawson), including one specimen which was recorded as emerging from a dead P. ponderosa branch. It has also been found on Engelmann spruce (Picea engelmanni Parry ex Engelm.), scrub oak (Quercus turbinella Greene), and cherry (Prunus sp.). Specimens have been collected at UV lights, with malaise and flight-intercept traps, and by beating vegetation. Fall (1901: 177) wrote "...rare during May and June; found always on oaks, notwithstanding its name." The larva of L. pinicola has been illustrated by Lawrence (1991), but not described in detail. The phenology information that is available (Fig. 16) indicates that adults can be found between 13 May and 29 August with two specimens having been found in the autumn (6 October and 6 November). The peak in adult numbers appears to be in the first half of July.