Aneflomorpha Casey, 1912

(Fig. 1–3, 5–16)

Aneflomorpha Casey 1912: 293 . Type species: Elaphidion subpubescens LeConte 1862 . Original designation.

Discussion. Aneflomorpha species are generally nondescript and varying shades of brown in color, ranging from mostly pale testaceous (as in A. gilana Casey and A. linearis (LeConte), Fig. 1h, i), rufous (as in A. aculeata (LeConte) and A. linsleyae Chemsak, Fig. 1a, 2a), brunneous (as in A. fisheri Linsley and A. luteicornis Linsley, Fig. 1g, 2b), or piceous (as in some A. cazieri Chemsak and A. rectilinea Casey, Fig. 1c, 2h). Most specimens range between 11–15 mm long, but a few are 7–10 mm (e.g., A. crypta, n. sp., Fig. 1e, and A. minuta Chemsak, Fig. 2c), and others can be 18–20 mm (e.g., some A. fisheri, Fig. 1g, and A. unispinosa, Fig. 1b, 3e). Most species are quite narrow bodied with the pronotum nearly always longer than wide, ranging from 0.95 times longer than wide (as in A. gilana, Fig. 5h) to 1.3 times longer than wide (as in A. tenuis (LeConte), Fig. 6u), with most species having the pronotum 1.1–1.2 times longer than wide. The elytra together range from 3.1 times longer than wide (as in A. texana Linsley, Fig. 3c, d) to 3.93 times longer than wide (as in A. crypta, n. sp., Fig. 1e) with most species having the elytra 3.3–3.5 times longer than wide. This elongate and narrow body form is well adapted to, and a consequence of, the larval habits of developing in small diameter twigs and branches which constrain the adult size and proportions (Craighead 1923; Heffern et al. 2018).

Most species have a well-developed and acute spine on antennomere three that is longer than the second antennomere and successively smaller spines at the apex of antennomeres four and five (Fig. 9), however some species have the spine on antennomere three either blunt (as in A. tenuis, Fig. 9r) or shorter than the second antennomere (as in A. unispinosa, Fig. 9s) and may have subsequent spines dentiform or absent as in A. texana (Fig. 9t). Some species, such as A. rectilinea (Fig. 9o), which have well-developed spines that project from the antennal plane, use the spine of the third antennomere, primarily, as a defense against predation by rapidly moving the antennae back toward the elytra and stabbing the offender (pers. obs.). Most species have a variably developed carina on the basal antennomeres that can be quite bold (as in A. luteicornis, Fig. 9j, and A. rectilinea, Fig. 9o), or absent (as in A. subpubescens, Fig. 9q and A . cazieri Fig. 9b).

All species have the pronotum punctate (Fig. 5, 6), but vary in the density, size, and distribution of those punctures. In most species the punctures are partially hidden by pubescence, but in some species, such as A. subpubescens (Fig. 6t) the punctures are nearly completely exposed, and in others, like A. yumae Giesbert and Hovore (Fig. 6x), they are nearly completely hidden. Aneflomorpha species vary with regard to the presence of a medial, impunctate pronotal callus (Fig. 5, 6). Most species do not have a callus. Some species are variable and have only a small callus in some specimens (e.g., some A. cazieri Fig. 5c, A . delongi (Champlain and Knull) (Fig. 5f), A. crypta (Fig. 5e), A. minuta (Fig. 5l), A. tenuis, Fig. 6u, and A. texana, Fig. 6w). The sides of the pronotum in most Aneflomorpha are broadly, gradually rounded, or nearly straight, without tubercles. However, A. gilana (Fig. 5h) and A. linearis (Fig. 5i, 6o) have weakly produced tubercles at the middle and some specimens of A. minuta (Fig. 5l) have the posterior fifth of the pronotum more abruptly constricted than the anterior margin. Most species have the procoxal cavities closed or nearly closed posteriorly by an expanded procoxal process (Fig. 10), but a few have the intercoxal process barely expanded and have widely open cavities (as in A. linearis, Fig. 10i, o, A . subpubescens, Fig. 10t, and A. yumae, Fig. 10x).

Most species have the elytral apices symmetrically dentiform or bispinose (Fig. 8), although a few species have the apices unarmed and truncate (as in A. minuta, Fig. 8k and A . crypta, n. sp., Fig. 8d). A few species have the apices asymmetrical, with the outer apex rounded to a dentiform or spinose suture (as in A. unispinosa, Fig. 8t). Most species have moderate pubescence on the elytra comprised primarily of recumbent, recurved setae (e.g., Fig. 7e, q, v), however, a few species have a nearly equal amount of erect or semi-erect setae in addition (as in A. aculeata, Fig. 7a and A . tenuis, Fig. 7u), and a few have only erect and suberect setae and lack recumbent setae entirely (as in A. linsleyae, Fig. 7j and A . subpubescens, Fig. 7t).

Aneflomorpha is most similar to species in the genera shown in Figure 4. These include Aneflus LeConte (Fig. 4a), Anelaphus Linsley (Fig. 4b), Anopliomorpha Linsley (Fig. 4c), Elaphidion Audinet-Serville (Fig. 4d), Micraneflus Linsley (Fig. 4e), Micranoplium Linsley (Fig. 4f), Neaneflus Linsley (Fig. 4g), Orwellion Skiles (Fig. 4h), Parelaphidion Skiles (Fig. 4i), Pseudoperiboeum Linsley (Fig. 4j), Psyrassa Pascoe (Fig. 4k), and Stenelaphus Linsley (Fig. 4l). Characters distinguishing each of these genera are discussed below and extracted from the following papers where these genera were fully diagnosed: Lingafelter (1998, 2007, 2020), Lingafelter and Ivie (2004), and Linsley (1963).

Aneflus (Fig. 4a) have an elongate morphology similar to Aneflomorpha, but the ratio of elytra length to width is lower, with most species having both elytra less than 3.3 times as long as wide. Likewise, the pronota of many species of Aneflus are as wide as long (or wider), and this is very rare in Aneflomorpha . With few exceptions, such as small individuals of Aneflus levettei (Casey) (Fig. 4a), Aneflus species are more robust and are at least 20 mm in length. Aneflomorpha unispinosa Casey (including its new synonym A. arizonica Linsley) (Fig. 1b, 3e) and A. tenuis (Fig. 3b) are regularly in that size range and rarely, a few very large individuals of other species such as A. fisheri Linsley (Fig. 1g) and A. rectilinea (Fig. 2h) also approach this size. Most species of Aneflus have pronounced mesal antennal spines present on antennomeres 3–7 (rarely more than dentiform beyond 5 in Aneflomorpha) and sometimes also apicolaterally (apicolateral spines are absent in Aneflomorpha). Nearly all specimens of Aneflus have strongly bispinose elytral apices, while in Aneflomorpha, the apicolateral spine is usually reduced, dentiform, or absent, however, exceptions are found such as in A. aculeata .

With a few exceptions such as Anelaphus villosus (Fabricius) (see Lingafelter (2020) for full description of this species), Anelaphus species (Fig. 2b) are not as elongate and narrow-bodied as Aneflomorpha and have elytra length to width ratios much less than 3.3. Further, most species of Anelaphus have the pronotum as wide as, or wider than long. Most Anelaphus species have a rather distinct pubescent patch on the antennal tubercles, while in Aneflomorpha, it is lacking or inconspicuous. The antennae are not carinate in Anelaphus but are in most species of Aneflomorpha .

Anopliomorpha species (Fig. 4c) are quite small (generally less than 8 mm) and would be at the lowest size range of Aneflomorpha, comparable to small individuals of A. cazieri Chemsak (Fig. 1c), A. minuta Chemsak (Fig. 2c), and A. crypta Lingafelter, n. sp. (Fig. 1e). Like most Aneflomorpha, Anopliomorpha species have the antennae carinate on the basal segments. The pronotum in Anopliomorpha is densely, confluently alveolate-punctate while in Aneflomorpha, the punctures are usually not all confluent and are never alveolate. The presence of very long “flying” setae scattered over the body and appendages (present in only a few species of Aneflomorpha, but not as strongly developed), combined with a very densely white pubescence scutellum are also distinctive for Anopliomorpha .

Almost all species of Elaphidion (Fig. 4d) have a very pronounced impunctate median pronotal callus. The callus, when present in Aneflomorpha, is typically very narrow and not as conspicuous. Most species of Elaphidion have strongly developed elytral spines (often strongly bispinose), and more developed than in most Aneflomorpha species. In Aneflomorpha, only A. aculeata (Fig. 8a) has the elytral apices typically strongly bispinose. Nearly all species have a pronounced, abruptly declivous prosternal process that angles behind the procoxal cavities. In Aneflomorpha (and other genera of Elaphidiini), the process is gradually declivous. The antennae are not carinate in Elaphidion, unlike in most species of Aneflomorpha . Elaphidion species are much more robust than Aneflomorpha and have elytra length to width ratios less than 3.1 in most individuals. Further, most species of Elaphidion have the pronotum as wide as, or wider than long. The antennal spines are more pronounced in many species of Elaphidion and often are present apicolaterally on many antennomeres (apicolateral antennal spines are absent in Aneflomorpha .)

Micraneflus is a monotypic genus containing only M. imbellis (Casey) (Fig. 4e) and resembles Aneflomorpha but is easily distinguished since it lacks antennal spines. All species of Aneflomorpha have at least a small spine on antennomere three. The elytral apices are rounded apicolaterally in Micraneflus and this is present in only a few Aneflomorpha such as A. unispinosa Casey (Fig. 8t,w).

Another monotypic genus, Micranoplium (Fig. 4f), which is known only from the eastern United States, is, like Micraneflus, distinguished from Aneflomorpha by its lack of antennal spines. Further, most individuals are smaller than 10 mm in length, so only small individuals of A. cazieri Chemsak (Fig. 1c), A. minuta Chemsak (Fig. 2c), and A. crypta Lingafelter, n. sp. (Fig. 1e) are comparable.

Neaneflus (Fig. 4g) is very similar to Aneflomorpha and one species of the latter ( A. opacicornis Linsley) is transferred to the genus herein (Fig. 17). This genus differs from all Aneflomorpha in having strongly apicolaterally expanded antennomeres and pronounced sexual dimorphism in antennal length (Fig. 18). Neaneflus differs from most Aneflomorpha by its lack of antennal carinae. Further, the elytral and pronotal proportions are broader than all Aneflomorpha species with only A. texana (Fig. 3c, d) and A. gilana (Fig. 1h) having nearly as broad elytra and pronota, respectively. As in Micraneflus, the elytral apices are rounded apicolaterally, a rarely occurring character state in Aneflomorpha .

Orwellion (Fig. 4h), like Elaphidion, has much broader pronota (wider than long) and elytra (less than 3.1 times longer than wide) than Aneflomorpha species. Antennomere three is at least two-thirds the length of the pronotum in Orwellion (shorter in Aneflomorpha). The elytra and pronotum in Orwellion species have scattered dense pubescent patches while the pubescence is more uniformly distributed in Aneflomorpha .

Parelaphidion (Fig. 4i), like Elaphidion and Orwellion, has broader proportions and a less elongate facies than Aneflomorpha . The pronotum is wider than long and has multiple well-developed glabrous calli in Parelaphidion (longer than wide in nearly all specimens of Aneflomorpha and with, at most, a small median callus). The antennae are not carinate in Parelaphidion, unlike most species of Aneflomorpha .

One species of the small genus Pseudoperiboeum ( P. lengi (Schaeffer), Fig. 4j), was included in Aneflomorpha by Linsley (1963) in part due to the carinate antenna. It is distinguished by having a moderately developed lateral pronotal projection or tubercle on each side (the pronota of most Aneflomorpha are evenly rounded (or nearly straight) at the sides, Fig. 5, 6), thus making the pronotum about as wide or wider than long (longer than wide in nearly all species of Aneflomorpha). The integument is covered with long, flying setae unlike nearly all species of Aneflomorpha (except A. aculeata (LeConte), Fig. 7a).

Psyrassa (Fig. 4k) (and the recently synonymized Megapsyrassa, García and Santos-Silva, in press) is most similar to Aneflomorpha in terms of the elongate and narrow pronotum and elytra and presence of antennal carinae in some species. It is distinguished from Aneflomorpha by having the pronotum mostly smooth, shiny, glabrous, and nearly impunctate (heavily punctate with punctures usually partially obscured by pubescence in Aneflomorpha).

Stenelaphus (Fig. 4l) is a monotypic genus that could be confused initially with Aneflomorpha, but upon examination, the combination of antennae lacking carinae, broad pronotum with smooth, impunctate calli; nearly glabrous elytra except for widely scattered, long, erect setae; densely golden pubescent scutellum; and rounded elytral apices to a spinose or dentiform suture are unique to Stenelaphus and not present in any Aneflomorpha species.