The five tribes represented in the New World are unlikely to represent a monophyletic group and as such cannot be satisfactorily diagnosed by morphological characters or separated from the Old World tribes as a whole, even when excluding the aberrant genera.

This monotypic tribe has been enigmatic in its placement in Curculionidae since the description of its sole genus by Chevrolat in 1879. Trichodocerus Chevrolat, 1879 was originally considered by Chevrolat to be near Conotrachelus Dejean, 1835 and it has since been treated as or had its species described in the Cryptorhynchinae (Champion 1906: 713, Hustache 1936, Papp 1979, O’Brien and Wibmer 1982, Wibmer and O’Brien 1986, Zherikhin and Gratshev 1995), Baridinae (Bondar 1946) and Conoderinae (Wibmer and O’Brien 1989: 15, Wolda et al. 1998, Alonso-Zarazaga and Lyal 1999, Lyal et al. 2006, Bouchard et al. 2011, Prena et al. 2014), where it currently resides.

Trichodocerus spinolae Chevrolat, 1879 [by subsequent designation: Champion 1906: 713].

Masculine.

Figures 67–70. 

Lateral and dorsal habitus images of Trichodocerini and Piazurini. Scale bars = 2 mm unless otherwise specified. 67a–b Trichodocerus spinolae [ARTSYS0000534] 68a–b Cratosomus lafontii [SSAC0001133]; scale bars = 10 mm 69a–b Lobops bonvouloiri [ARTSYS0000527] 70a–b Piazurus caprimulgus [SSAC0001113].

Trichodocerus is easily separated from all other genera treated here by the loose antennal club, the presence of elongate setae on the club and funicular articles, contiguous procoxae, a prosternal rostral channel with the lateral margins strongly converging and meeting below the procoxae, a mesoventrite with a transverse ventrally produced ridge anterior to mesocoxae and the mesoventrite posterior to ridge with dense yellow scales (Fig. 1), and the presence of stridulatory plectra on the male seventh abdominal tergite. Funicular article 1 is short and globose, the hind femora are not carinate and ventrally with one or no teeth, the femoral apices are always unarmed at the lateral and mesal faces, the eyes of most species have a distinct lower constriction lateral to rostral insertion (Fig. 37), the abdominal ventrites are slightly ascending, and the pygidium is concealed (slightly exposed in male T. spinolae and male of at least 1 undescribed species).

The contiguous procoxae efficiently separates Trichodocerus from all genera except some species of the very different Zygops. The yellow scales of the mesocoxae and mesoventrite have also been observed in Arachnomorpha Champion, 1906 (on the pro- and mesocoxae), Microzygops Champion, 1906 (on the procoxae), and some species of Lechriops and Macrocopturus (on the pro- and mesocoxae and ventrally on the mesofemora), but in those genera they are not also present on the mesoventrite between the mesocoxae.

The numerous features that easily separate Trichodocerus from the rest of the conoderines are potentially indicative of improper placement in the Conoderinae, however, the same unique characters prevent confident reassignment to another group of Curculionidae. Champion (1906: 713) noted the similarity of the antennal funicle to Hedycera Pascoe, 1870, a South American genus here reassigned to the Piazurini.

The genus is apparently nocturnal, with most observed specimens being collected at UV light traps – 488 specimens were collected by Wolda et al. (1998), representing three of the 17 species of Conoderinae with more than 10 specimens collected. One undescribed species has been reared from balsa, Ochroma pyramidale (Cav. ex Lam.) Urb. (Malvaceae: Bombacoideae Burnett) (Hespenheide, personal communication). An observed specimen of T. brevilineatus Champion, 1906 [STRI_ENT_008474] was recorded from dead branches of balsa and an undescribed species [SSAC0001000] was collected on balsa leaves; another observed undescribed species has been collected on dead branches of Pachira sessilis Benth. (Bombacoideae) [STRI_ENT_0084793].

Two species are known from the focal region and one additional described species occurs in South America. I have accumulated and distinguished over 20 species of Trichodocerus new to science as part of a revision currently in preparation.

Guatemala, Costa Rica, Panama; South America. Undescribed species are also known from Mexico.

This tribe was originally characterized by Lacordaire (1865: 144) for the genera Cratosomus, Pinarus, and Piazurus in recognition of the strongly canaliculate prosternum, the “gutter-like” modification to the mesoventrite and the clavate, non-carinate hind femora that do not or only slightly exceed the abdominal apex. Heller (1906: 31) produced a key to Piazurini that includes 8 of the 12 currently recognized genera – not included are Lobops, Latychellus Hustache, 1938, Hedycera, and the Old World Guiomatus – based largely on the relative size of abdominal ventrites and the amount that they ascend, relative lengths of funicular articles, and the construction of the mesoventrite.

The monotypic South American genus Hedycera is moved to the Piazurini despite the occurrence of the genus outside the geographic focus of this paper. The exposed pygidium that is not completely visible in dorsal view, the large triangular tooth on the hind femur, the transverse posteromedial depression on the metaventrite (discussed further below), and the unarmed femoral apices place the genus not only in the Piazurini but in a hypothesized clade containing Piazurus, Pseudopiazurus Heller, 1906, Pseudopinarus Heller, 1906 and the South American Piazolechriops Heller, 1906. Hedycera megamera Pascoe, 1870 would key out to couplet 7 of Heller’s 1906 key (containing Pseudopinarus and Piazolechriops), for having abdominal segments only slightly ascending, abdominal segment 2 not being longer than 3 and 4 combined, and the presence of “superciliarleisten”, referring to the arcuate carina at the vertex of the head found in most members of these genera (though not in a few species of Pseudopinarus), a greatly elongate antennal funicular article 2, and a slender rostrum. Hedycera can be differentiated from these by the shape of the pronotum in dorsal view, which is widest in the anterior half just before the subapical constriction, and in having elongate setae on the antennal funicular articles. When originally describing the monotypic genus, Pascoe (1870) stated that it was most closely related to Piazurus, which was later agreed with by Champion (1906: 713). Hedycera is the first genus separated in Heller’s key (1895) for having similar-sized abdominal ventrites 2, 3, and 4, but is not treated further in that publication. In the catalogs of Hustache (1934: 45) and Blackwelder (1947: 884) Hedycera is listed under the otherwise entirely Old World-distributed conoderine tribe Mecopini Lacordaire, 1865 and was moved to Lechriopini in Wibmer and O’Brien (1986: 19), without a justification provided in either placement.

The modification to the mesoventrite in the genera treated here in the Piazurini varies from being a cup-shaped receptacle (as in Lobops; Fig. 2) to structured similarly to a cup-shaped receptacle but with the posterior margin flattened and depressed at least slightly below the level of the lateral margins of the channel (Figs 3, 4) allowing the rostrum to pass through to the metaventrite if long enough. The eyes are often smaller and more separated and are not or not as sharply acuminate ventrally or laterally inflexed (Figs 38, 39) as in many Lechriopini and Zygopini, but can be quite large and contiguous or subcontiguous (Figs 40–41), taking up most of the surface of the head as well as be slightly ventrally acuminate to slightly laterally inflexed. The pygidium is exposed but not entirely visible in dorsal view (somewhat concealed from above by the elytral apex; e.g. Fig. 68), usually only visible completely in posterior or ventral view. Abdominal ventrites are flat to slightly, evenly ascending. The vestiture consists of thick setae to small scales, usually not covering most of the body surface except in Lobops, which has large, flat and round scales. The femora are at least slightly clavate, the hind femur is without a lateral carina and lacks teeth at the mesal and lateral apical faces (Fig. 60; in most lechriopines and zygopines, a tooth is usually present at the mesal and/or lateral face of the femoral apex on the middle and/or hind legs as in Figs 61–63), and several genera have a large, laterally compressed, triangular ventral tooth. This large triangular tooth is also found in other conoderine tribes (e.g. Menemachini Lacordaire, 1865; Campyloscelini Schoenherr, 1845) as well as other groups of weevils (e.g. Hylobiini Kirby, 1837). Despite this homoplasious distribution in Curculionidae it likely represents a single origin within the Piazurini, with the genera having it also sharing additional characters; it is also not found in other New World Conoderinae, making it useful for diagnosing the group of Piazurines that bear it.

The metaventrite posteromedially has a transverse depression, not with a narrow longitudinal sulcus extending variably anteriorly as in most Lechriopini and Zygopini (but many species of Cratosomus have a broad longitudinal depression). The antennal club is typically more spherical to ovoid, with the suture between at least articles 2 and 3 sinuate (but also found in a few lechriopines and zygopines). A mesal process of the procoxae is absent in most piazurines and found in many lechriopines and zygopines (though present, among the Central American species observed, in Pseudopinarus, Lobops bonvouloiri (Hustache, 1932), and in the species Piazurus alternans Kirsch, 1875). Sclerolepidia are absent in Piazurini (Lyal et al. 2006: 237). Additionally, piazurines are quite different behaviorally from the remainder of the New World Conoderinae, typically being less active in the daytime and no species are known to be part of the several widespread mimicry complexes found in the tribes Lechriopini and Zygopini (Hespenheide 1995).

Fifty-two species are currently known from north of South America in five genera. Six additional genera are known only from South America, and one genus, Guiomatus, occurs in Papua New Guinea.

Rhynchaenus herculeanus Dalman, 1823 [by original designation] (=Rhynchaenus roddami Kirby, 1819).

Masculine.

Cratosomus can be differentiated from the other Piazurines treated here by the setal tufts of the anterior margin of the tibial apex being composed of thick fascicles of golden setae (Fig. 21), the dense setal brush at the posterodistal face of the meso- and metatibia (Fig. 21), the thick rostrum that is apically dorsoventrally compressed, and generally larger body size. The eyes can be small and widely separated (Fig. 38) or large and approximate; the femora are ventrally with 0-2 teeth, usually with a distinct ventral carina distally; and the elytra and pronotum are often tuberculate or spinose.

This genus includes some of the largest Neotropical weevils (Champion 1906: 1). Males of some species have lateral tusk-like processes of the rostrum and are presumably under sexual selection and used during male-male competitions.

Emden 1933 (Central and South America), Champion 1906: 2 (Central America).

Schoenherr’s (1838) classification included Cratosomus in a separate Cohors of Cryptorhynchides, thereby distinguished from the rest of the then-described Conoderinae. Although somewhat dissimilar in appearance to the Piazurini treated here, it resembles the South American piazurine genera Latychus Pascoe, 1872 (and likely also the South American Costolatychus Heller, 1906, and Latychellus Hustache, 1938, but no specimens were observed of those genera) in the thickened dorsoventrally compressed rostral apex, the smaller and relatively widely separated eyes (as in some Cratosomus), and the small ventral femoral tooth.

R.S. Anderson (1993: 218) lists Annonaceae Juss., Rutaceae Juss., Lauraceae Juss., Myrtaceae Juss., and Sapotaceae Juss Hosts for several species of Brazilian Cratosomus are recorded by Costa-Lima (1956: 213).

Twenty-five species are known from the focal region (with 14 subspecies or forms) and an additional 126 species (and many subspecies or forms) are known exclusively from South America (Wibmer and O’Brien 1986: 254; Rheinheimer 2011: 66 described one more).

Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama; South America. Distributions of Guadeloupe and the Lesser Antilles are listed with doubt by Emden (1933: 532) and subsequent catalogs.

Lobops setosus Fåhraeus, 1845 [by original designation].

Masculine.

Lobops is unique among the Piazurines in having a prominent cup-shaped receptacle for receiving the rostrum on the mesoventrite (Fig. 2), a dense covering in flat, round scales, and strongly concave interocular space (Fig. 39). The metafemoral tooth is not especially large, the femora are not strongly clavate, and the second funicular article is relatively short (not longer than the first); these characters are also shared with Cratosomus but not other piazurine genera treated here.

Of the five piazurine genera covered here, Lobops has the least certain placement in the tribe. Schoenherr (1845) originally indicated for the South American type species a relationship with Conotrachelus, and the genus was previously placed in the Ithyporini Lacordaire, 1865 (O’Brien and Wibmer 1982: 125, as Cryptorhynchinae; the tribe is currently placed in the Molytinae), overlooking a transfer to the Piazurini by Kuschel (1955: 271). The only known Central American species, L. bonvouloiri, was originally described in the genus Pseudopinarus. The structure of the mesoventrite is suggestive of placement in the Cryptorhynchinae, but that type of receptacle has been shown to not be exclusive to the subfamily (Riedel et al. 2016: 5). Lobops is certainly better placed in the Piazurini than in the other tribes reviewed here – despite differences in a number of characters, the exposed pygidium that is not visible in dorsal view, the unarmed femoral apices, non-carinate femora, and lack of sclerolepidia, in combination, are unique to the Piazurini.

Unknown.

One species is known from the focal region and two additional species are known from South America (Wibmer and O’Brien 1986: 263).

Panama; South America.

Poecilma stipitosum Germar, 1824 [by original designation].

Masculine.

An elongate second funicular article and broad triangular femoral tooth place Piazurus near Pseudopinarus and Pseudopiazurus, and it can be differentiated from them by the longer second abdominal ventrite (which is as long as the third and fourth ventrites when seen from the side) and the protibial apex that bears a premucro. Being a much more diverse genus than Pseudopinarus and Pseudopiazurus, it is easiest to arrive at an identification by a negative identification of those two smaller genera: namely, species of Piazurus never have a carinate vertex of the head (as in many Pseudopinarus), a strongly impressed first abdominal ventrite (as in Pseudopiazurus), or a subapical premucro (as in some Pseudopinarus and Pseudopiazurus) and usually do not have a mesal procoxal process (which is found in most Pseudopinarus and a South American Pseudopiazurus; it is present at least in Piazurus alternans).

Fiedler (1936) divided Piazurus into seven groups based mainly on the shape of the elytra and the presence, location and shape of elytral tubercules.

Fiedler 1936 (Central and South America), Heller 1906: 33 (Piazuruss. str. of Central and South America), Champion 1906: 9 (Central America).

Of the genera with a broad ventral metafemoral tooth, Piazurus is most similar to Pseudopiazurus with a conical prothorax and lack of a mesal process of the procoxae (though it is present in at least one species of each genus).

Some species have been reared from branches of various genera of Lecythidaceae (Fassbender 2013, Fassbender et al. 2014). Maes and O’Brien (1990) report Piazurus trifoveatus Champion, 1906 from Coffea L. (Rubiaceae Juss.) and Costa-Lima (1956: 218) reports a Brazilian species from fruits of Myrtaceae.

Nineteen species are known from the focal region and an additional 58 species are exclusive to South America (Wibmer and O’Brien 1986: 260).

Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama; South America.

Piazurus obesus Boheman, 1838 [by subsequent designation: Rheinheimer 2011: 76].

Masculine.

Figures 71–74. 

Lateral and dorsal habitus images of Piazurini, Othippiini and Lechriopini. Scale bars = 2 mm unless otherwise specified 71a–b Pseudopiazurus centraliamericanus [ASUHIC0086627]; scale bars = 5 mm 72a–b Pseudopinarus condyliatus [ASUHIC0086626]; scale bar for 72a = 1 mm 73a–b Acoptus suturalis [ASUHIC0016914] 74a–b Copturomimus cinereus [ASUHIC0086628].

Pseudopiazurus centraliamericanus (Heller, 1906), the only recorded Central American species of Pseudopiazurus, can be separated from other Central American piazurines by the deep U-shaped depression on the first abdominal ventrite (Fig. 64; also found in a few Pseudopinarus, but when present in that genus there is always also present the arcuate carina on the vertex of the head which is never found in Pseudopiazurus), the large subapical premucro of the metatibia (Fig. 22; but also at least in Pseudopinarus guyanensis Hustache, 1938), the absent premucro at the protibial apex (also in species of Pseudopinarus), the very large, ovoid, contiguous eyes (Fig. 40), and the large, deep, ovoid punctures of the elytral striae (but also found in some Cratosomus species).

First described by Heller (1906) along with Pseudopinarus as subgenera of Piazurus. As documented by Champion (1906: 18), after the subgenus was first introduced in the key the name was erroneously switched with that of Pseudopinarus and the key to species of Pseudopiazurus is given under the name Pseudopinarus (Heller 1906: 34). The catalog of South American species by Wibmer and O’Brien (1986: 262) makes a correction to the two species given for Central America by O’Brien and Wibmer (1982: 160), which overlooked an error by Hustache (1934), leaving P. centraliamericanus the only species known from Central America.

Marshall (1922: 69), Fiedler 1936: 28 and Heller 1906: 34 (under Pseudopinarus, in error) also contain keys to species.

This genus is most similar in overall appearance to Piazurus (see above), but the genus as a whole is incompletely distinguishable from Pseudopinarus by obvious characters with the exception of the much larger eyes, usually larger and more ascending mesepipleura, and absent ventral profemoral tooth of Pseudopiazurus – the deep arcuate sulci of the first ventrite in Pseudopiazurus is apparently also found in Pseudopinarus, e.g. in Pseudopinarus quadratus (Champion, 1906); the mesal process of the procoxae is absent in Pseudopiazurus centraliamericanus but present in the South American Pseudopiazurus spiniventris Marshall, 1922 as well as in many Pseudopinarus, and the subapical premucro of the metatibia is also found in Pseudopinarus guyanensis. Despite the overlap in these characters, Pseudopinarus is interpreted here as more closely related to the South American Piazolechriops, Pinarus, and Hedycera.

R.S. Anderson (1993: 218) lists Guttiferae Juss. (=Clusiaceae Lindl.). Marshall (1922: 67) records the South American P. obesus (=P. papayanus (Marshall, 1922)) as a borer of the “leaf-stems” of papaya (Caricaceae Dumort: Carica papaya L.).

One species is known from the focal region and three additional species are known exclusively from South America (Wibmer and O’Brien 1986: 262, including Costa-Lima’s (1956: 217) synonymy of P. papayanus with P. obesus, which was either overlooked or disputed, as they are treated as separate by Wibmer and O’Brien).

Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama; South America.

Piazurus rana Heller, 1906 [by subsequent designation: Rheinheimer 2011: 76].

Masculine.

As a subgenus of Piazurus, Pseudopinarus was separated from Piazuruss. str., along with the other subgenus, Pseudopiazurus, by Heller (1906) and Fiedler (1936) for having a second abdominal ventrite that is shorter in length than the third and fourth ventrites combined. Pseudopiazurus is easily distinguished from Pseudopiazurus in body shape, which is much more robust in Pseudopiazurus, but as a whole Pseudopinarus is the most difficult piazurine genus to characterize. Most species can be further distinguished from Pseudopiazurus by the presence of a ventral tooth on the profemora. Some Pseudopinarus have an arcuate carina on the vertex of the head (Fig. 41) similar to the South American Piazolechriops and Hedycera. The eyes are generally smaller and more separate than in Pseudopiazurus, but can be large and subcontiguous (as in Fig. 41). The structure of the mesoventrite varies as well, with some species bearing ventrally produced posterolateral tubercles (as in Pseudopiazurus) and others with nearly a cup-shaped receptacle. Additionally, the relatively small mesepipleura and the procoxae with a mesal process differentiates some Pseudopinarus from most species of Piazurus and the single Central American species of Pseudopiazurus.

Fiedler 1936: 29, Heller 1906: 34.

The species of Pseudopinarus that have the arcuate carina on the vertex of the head are very similar to the South American genera Piazolechriops and Hedycera than to other Central American genera. The only character given by Heller (1906) to separate Pseudopinarus from Piazolechriops is the shorter hind femur of Pseudopinarus, which do not, or only very slightly, extend beyond the apex of the abdomen. Pseudopinarus differs from Hedycera by the short antennal setae and the pronotum in dorsal view, which is not widest just before the apex. Other species of Pseudopinarus, e.g. P. cerastes (Fabricius, 1801), are more similar to smaller species of Piazurus (e.g. P. alternans), raising the question of the monophyly of the genus and the validity of the shorter second abdominal ventrite as a character separating monophyletic groups from Piazurus.

Some species have been reared from branches of various genera of Lecythidaceae (Fassbender et al. 2014). Pseudopinarus guyanensis has been reared from seeds of Gnetum L. (Gnetaceae Blume) [ASUHIC0086636, STRI_ENT_0082031].

Seven species are known from the focal region (Wibmer and O’Brien 1986: 263 add P. guyanensis to the Central American fauna) and an additional 13 species are known only from South America (Wibmer and O’Brien 1986: 262).

Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama; South America.

Eleven genera were first grouped into tribe Othippiini by Hustache (1938: 63), which was not treated as a valid name in the catalog of Alonso-Zarazaga and Lyal (1999: 113) for lacking a description. Morimoto (1962: 47) provided some clarification on the distinction of the tribe, and this was further refined in Kojima and Lyal (2002), where seven genera were transferred out of the Othippiini in order to redefine it. Othippiines can be distinguished (sensuKojima and Lyal 2002: 172) by the following combination of characters: the mesepisterna are non-ascending, the scutellum is exposed, the prosternum of most is canaliculate, and the antennal funiculus has 7 articles. However, the monophyly of the tribe has yet to be shown (Kojima and Lyal 2002).

The mesoventrite of othippiines can have a rostral channel or be unmodified (Kojima and Lyal 2002). The number of antennal funicular articles for othippiines given by Kojima and Lyal (2002) serves to separate this tribe from the Mecopini which have been considered to have 6 articles since originally described. However, one of the genera currently treated in the Mecopini (Emexaure Pascoe, 1871) has a funiculus with 7 articles (Pascoe 1871: 216), some of the genera are similar in appearance and also in eye shape (Kojima and Lyal 2002: 171) and distinction between the tribes requires further study.

Othippiini now contains eight genera, six of which are currently monotypic. The monotypic genus Acoptus is the sole New World representative.

Acoptus suturalis LeConte, 1876 [by monotypy].

Masculine.

Acoptus can be easily recognized from the rest of the conoderines treated here by the following characteristics: the inner margin of eyes towards the top has a large lanceolate space (Fig. 42) and the eyes are nearly touching above and below the lanceolate space, the mandibles are somewhat falcate and are in contact only at the apex (visible in Fig. 5), and the tibial apex distally has the dorsal margin dilated and premucro absent from all tibiae (Fig. 24). The relatively forward facing eyes that do not extend much on lateral portions of head (genae large), the first funicular article that is longer than the second, the unmodified mesoventrite, the ventrally toothed and non-carinate metafemora, the unarmed femoral apices, the small and non-ascending mesopleura, the absent sclerolepidia, and the abdominal ventrites that are not rapidly ascending additionally help diagnose the genus and in combination separate it from all Lechriopini and other New World Conoderinae.

The species Homogaster quebecensis was first placed in synonymy with Piazurus subfasciatus LeConte, 1876 (=Lechriops subfasciatus (LeConte)) by LeConte (1880: xii), where it remained in catalogs until Blackwelder and Blackwelder (1948: 48) listed it in synonymy with Acoptus suturalis. Provancher’s description agrees with that of Acoptus.

In keys to North American genera, Acoptus is always separated from the rest of the North American genera along with Psomus by the flat abdominal ventrites (e.g. Hespenheide 2002: 755). Casey (1892: 458, 1897: 666) suggested a relationship to Psomus on these grounds, but this similarity only suggests they are both aberrant in their placement in the Lechriopini – they are otherwise very different in appearance, as also noted by Casey (1892: 458). LeConte (1876: 264) originally distinguished his genus from Copturus (which, with the geographic scope and time of publication of the key included only species currently placed in Cylindrocopturus and Eulechriops) and Zygops by the nearly flat abdominal ventrites and elongate first funicular article. Provancher (1877: 530), apparently independently, separated his genus Homogaster from Zygops and Copturus (as well as the South American Timorus) for the same reasons. A phylogeny by Davis (2014) recovered Acoptus as closer to the Old World genus Mecopus (Conoderinae: Mecopini) than any of the included New World conoderines, a result consistent with the new placement of the genus (the study did not include Othippiini).

Mentions in the literature of host plants include Fagus L. (Fagaceae Dumort) (Chittenden 1890: 171), Ulmus americana L. (Ulmaceae Mirb.) (Hoffman 1942: 12) and Juglans cinerea L. (Juglandaceae DC. ex Perleb) (Halik and Bergdahl 2002). Sleeper (1963: 215) additionally reports Quercus L. (Fagaceae), Cercis canadensis L. (Fabaceae Lindl.), Carya Nutt. (Juglandaceae), and Platanus occidentalis L. (Platanaceae T. Lestib.). Adults of A. suturalis have been implicated as vectors of the chestnut blight fungus, Cryphonectria parasitica (Murrill) Barr (Pakaluk and Anagnostakis 1977) and the butternut canker fungus, Sirococcus clavigignenti-juglandacearum Nair, Kostichka & Kuntz (Halik and Bergdahl 2002).

One.

Eastern Canada, Eastern U.S.A., extreme northeastern Mexico (Sleeper 1963: 215). Specimens have not been observed from Mexico or even Texas to confirm Sleeper’s range extension; recently the genus was reported for the first time from Arkansas (Skvarla et al. 2015).

This tribe was originally characterized by Lacordaire (1865: 149) for the genus Lechriops by the rostral channel, which is closed (horseshoe-shaped) posteriorly to receive the rostrum and the linear, carinate femora that may or may not exceed the apex of the abdomen.

While a subclassification for the Lechriopini is not formally proposed here without also examining the South American genera, the following groups of genera are hypothesized to be related: the “Eulechriops genus complex”, including Eulechriops, Macrolechriops Champion, 1906, Copturomorpha Champion, 1906, Cylindrocopturinus Sleeper, 1963, Coturpus R.S. Anderson, 1994, and Turcopus R.S. Anderson, 1994 and the “Macrocopturus genus complex”, including Macrocopturus, Copturomimus Heller, 1895, Lechriops, Pseudolechriops, Hoplocopturus Heller, 1895, and Mnemynurus Heller, 1895. The genera Microzygops, Paramnemyne, Poecilogaster, Euzurus, Copturus, Microzurus and Psomus do not fit into either complex as currently conceived. Until the inclusion of the South American lechriopine genera a subtribal classification for the Lechriopini will not be further speculated here.

Among the genera currently placed in the tribe (sensuLyal et al. 2006), the only characters that distinguish them (after the exclusion of Acoptus, Philinna, and Philides) are a concealed pygidium with rapidly ascending abdominal sclerites, the presence of modification to the mesoventrite and/or the presence of sclerolepidia (just sclerolepidia in Copturomimus, most Macrocopturus and Psomus). The mesepipleura are usually large and somewhat ascending (except in Paramnemyne and Psomus). Other characters given by Lyal et al. (2006: 229) that separate lechriopines from zygopines are: “larger eyes, extending half-way or more down the side of the head; a longer rostrum, reaching at least the middle coxae; the middle and hind femora with the posterior distal margin extended into an acuminate projection extending beyond the anterior distal margin”, but these appear to be homoplastic – many lechriopines, especially some Eulechriops and related genera, have smaller eyes like many zygopines, and many zygopines have a similar femoral apex. The presence of a carina and ventral tooth on the hind femora, and the relative lengths of the first two funicular articles are potentially indicative of infratribal relationships; in the Eulechriops genus complex the hind femora are not carinate and unarmed ventrally and the second funicular article is at most subequal to the first, while in the Macrocopturus genus complex the hind femora are ventrally toothed and carinate and the second funicular article is longer than the first.

Modification to the meso- and metaventrite to receive the rostrum varies quite a bit in this group, with the typical forms (i.e. deviating the least from Lacordaire’s original tribal construction of a closed, horseshoe-shaped channel), being found in most members of the following genera: Lechriops, Poecilogaster, Eulechriops, Macrolechriops, Copturomorpha, Coturpus, Turcopus, Copturus, Microzurus, Euzurus, Microzygops and Pseudolechriops. These genera likely do not represent a monophyletic group, and the mode of closure (whether a simple depression or a strongly carinate apex of the channel) and the location of closure (on the mesoventrite or metaventrite) can vary significantly within genera. Pseudolechriops has arcuate lateral margins of the channel forming an ovoid carina that encircles a deep excavation on the mesoventrite and the anterior margin of the metaventrite (Fig. 13). A few species of the genus Macrocopturus (e.g. M. albidus Champion, 1906) and the genus Microzygops have a similarly constructed mesoventrite (Fig. 11) but the majority of Macrocopturus species and the very similar Copturomimus species have the unmodified “zygopine type” of mesoventrite. The mesoventrites of the genera Hoplocopturus and Mnemynurus are interpreted as of the lechriopine type, with the sides of the channel strongly arcuate and meeting medially, forming an inverted U-shaped carina that no longer appears to serve the function of receiving the rostrum (Fig. 9). Paramnemyne, Euzurus, Copturus, and Microzurus have a mesoventrite that would be classified here as the piazurine type (Heller 1895: 5 also notes the resemblance), with the rostral channel on the mesoventrite open (in Paramnemyne and Euzurus, Figs 8, 12) or closed (in Copturus and Microzurus, Fig. 6) and without anteriorly extending carinae; at least the latter three genera likely belong in the Lechriopini considering other characters. The mesoventrite of Psomus is unmodified.

Two hundred and forty-two species are currently known from north of South America in nineteen genera, comprising nearly half of the genus- and the majority of the species-level diversity of North and Central American Conoderinae. An additional eight genera are known only from South America.

Copturomimus cinereus Heller, 1895 [by present designation].

Masculine.

Copturomimus is similar to the large genus Macrocopturus with the elongate second funicular article, unmodified mesoventrite, and carinate and ventrally toothed hind femora, and can only be distinguished externally from that genus by the obliquely striolate area dorsally on the profemora (Fig. 66). The other genus with a striolate patch on the profemora, Copturomorpha, tends to have a striolate patch that is less obvious, being more finely striolate and more often concealed by scales; that genus otherwise is more similar to Eulechriops.

The function of the striolate profemora is unknown – the first conoderine species described with it, Copturomorpha musica (Kirsch 1875b), was named, as the specific epithet suggests, for its hypothesized stridulatory function (Kirsch 1875b: 248). The function of the patch was instead suggested to be for antennal grooming purposes (Champion 1906b: 60) due to the lack of an obvious corresponding file structure required for stridulation and the position of the leg relative to the antennal club – observation of Copturomimus caeruleotinctus Champion, 1906 [SSAC0001059] revealed the use of the setal comb at the protibial apex (and not the striolate femoral patch) for antennal cleaning purposes.

Champion 1906b: 60 (for Central America), Muñiz 1965: 5 (for three species on avocado, key modified from Muñiz and Barrera 1958: 2).

Heller (1895: 63) originally implied a relationship with his South American genus Copturosomus Heller, 1895, which is also difficult to distinguish from Macrocopturus. The relationship of both genera with Macrocopturus requires much more study to identify natural groupings of species. Whether the striolate femoral patch identifies a natural group is unknown but unlikely (Hespenheide 2009: 337). See also entry on Macrocopturus for discussion of the relationships of that hypothesized complex of genera.

. Copturomimus perseae (Guenther, 1935) and two other South American species are wood-boring on avocado (Lauraceae: Persea Mill.) (Hustache in Mariño M. 1947, Kissinger 1957, Muñiz 1965). Associations of other Central American species are unknown.

Twelve species are known from the focal region (one species described by Hespenheide 2009) and five additional species are known from South America (Wibmer and O’Brien 1986: 271; Muñiz 1965 transferred one species from Copturus).

Mexico, Guatemala, Honduras, Costa Rica, Panama; South America.

Copturomorpha interrupta Champion, 1906 [by original designation].

Feminine.

Figures 75–78. 

Lateral and dorsal habitus images of Lechriopini. 75a–b Copturomorpha sp. [ASUHIC0086641] 76a–b Copturus aurivillianus [ASUHIC0024140] 77a–b Coturpus arcuatus [ARTSYS0000531] 78a–b Cylindrocopturinus pictus [SSAC0001288]. Scale bars = 2 mm.

Like Copturomimus, Copturomorpha can only be distinguished from a much larger genus (Eulechriops) by the presence of a striolate patch dorsally on the profemur, and shares the short second funicular article that is not longer than the first, the unarmed and non-carinate metafemora and the excavate mesoventrite.

The presence of the striolate patch on the dorsal surface of the profemora is shared only with Copturomimus, where the patch is typically larger and more coarsely striolate. Copturomorpha will key out to Eulechriops if that character is overlooked – it is often indistinct and obscured by scales. Several South American species described in the genus by Hustache (1938) have a second funicular article that is longer than the first and a carinate and toothed hind femora in addition to the excavated mesoventrite, necessitating further study on the identity of Copturomorpha and the species currently placed there.

Champion 1906b: 65 (for Central America).

The combination of characters from the antenna, mesoventrite, and femora place Copturomorpha in a hypothesized genus complex including Eulechriops; whether the striolate femoral patch identifies a natural group separate from or within Eulechriops needs investigation.

Hosts of all described species are unknown; Fassbender (2013) and Fassbender et al. (2014) reared specimens from branches of Lecythidaceae that potentially represent a species of this genus.

Eight species are known from the focal region (one species described by Hespenheide 2011) and an additional 16 species are known from South America (Wibmer and O’Brien 1986: 271).

U.S.A.: Texas, Mexico, Guatemala, Panama; South America.

Poecilma papaveratum Germar, 1824 [by original designation].

Masculine.

Copturus is very similar to Microzurus with a concealed scutellum and a closed receptacle of the mesoventrite that is laterally flanged near the apex (Fig. 6), and can be distinguished from Microzurus by the ventrally toothed pro- and mesofemora, larger tarsal claws, and flattened (not costate) elytral intervals. The first two funicular articles vary in length among the species, with the second article being longer than or subequal to the first (Champion 1906b: 87).

The usage of this generic epithet has a particularly complicated history, as explained by Muñiz-Vélez and Ordóñez-Reséndiz (2010). The first usage of the name Copturus was as a subgenus of Zygops (Schoenherr 1825: col. 586), where the type species was designated as Poecilma papaveratum Germar, 1824. The subgenus was elevated to genus by Dejean (1835), and dozens of additional species were described to the genus (e.g. Schoenherr 1838, 1845, Kirsch 1875a, b). Heller (1895) created several genera out of specimens included in Copturus including the genus Zurus Heller, 1895 for the species of Copturus that have a concealed scutellum, second funicular article that is barely longer than the first, and a unique, horseshoe-shaped modification to the mesoventrite. Among the species moved from Copturus into Zurus was Poecilma papaveratum, which Heller recognized as the previously designated type species of Copturus. Neozurus O’Brien & Wibmer, 1982 was created as a replacement name for Zurus (O’Brien and Wibmer 1982: 4) which was preoccupied by Zurus Amyot, 1846 and was later synonymized with Copturus as an unjustified replacement name (Wibmer and O’Brien 1986, Alonso-Zarazaga and Lyal 1999: 111). As a result, all species treated as Copturus by Heller became newly recombined as Macrocopturus, one of Heller’s original subgenera of Copturus elevated to genus (Wibmer and O’Brien 1986: 17), and all species treated as Zurus by Heller and subsequent authors until Wibmer and O’Brien (1986) became newly recombined as Copturus, returning the type species Poecilma papaveratum to its original genus.

Champion 1906: 87 (to Zurus of Central America), Heller 1895: 5 (to Zurus of Central and South America).

Hespenheide (1984: 315) suggests a relationship with Microzurus, Euzurus, and Cylindrocopturinus. Of those three, it is most similar to Microzurus, the only differences given by Champion (1906: 87) being the ventrally toothed pro- and mesofemora, the larger body size and comparatively proportionate tarsal claws. Euzurus also has a scutellum concealed by a posterior lobe of the pronotum, but the manner of it’s concealment differs from Copturus and Microzurus: where the posterior pronotal lobe of Copturus and Microzurus subducts the elytra, completely concealing the scutellum, in Euzurus the posterior lobe is only extended posteriorly and not below the elytral base, leaving the scutellum visible in posterior view. Despite this difference, Lyal et al. report Type II sclerolepidia to be present in Copturus, Microzurus, and Euzurus, which, among the sclerolepidia-bearing lechriopines, is only also known in the very different Psomus. See entry on Cylindrocopturinus for the present interpretation of the relationship of that genus. The relationship of Copturus and Microzurus within the Lechriopini and whether or not Euzurus is found to be the sister-genus is not easily hypothesized by the external characters examined thus far.

The widespread Central and South American species Copturus aurivillianus (Heller, 1895) is reported by Costa-Lima (1956: 219) to bore stems of Canavalia Adans., Dolichos L., and Phaseolus L. (Fabaceae) as larvae in Brazil.

Six species are known from the focal region and an additional 27 species are known only in South America.

Mexico, Honduras, Nicaragua, Costa Rica, Panama, Guadeloupe; South America.

Coturpus arcuatus R.S. Anderson, 1994.

Masculine.

Within the Eulechriops complex of genera, Coturpus can be identified by lacking a striolate profemoral patch (as in Copturomimus), lacking a very prominent receptacle on the mesoventrite (as in Turcopus), lacking the premucro-like inner flange at the tibial apex (as in Cylindrocopturinus), and can be differentiated from the observed species of Eulechriops by bearing elongate setae on the ventral surface of strongly arcuate hind legs at least in the males and by lacking a procoxal mesal tooth (though with the vast numbers of undescribed Eulechriops it is difficult to rule out the absence of this character from that genus).

Females are unknown, and R.S. Anderson (1994: 482) suspects the modified hind legs to be found only in males as similar modification to the hind legs is known only in male Cylindrocopturinus.

R.S. Anderson (1994: 462) proposed a relationship to Cylindrocopturinus based on the presence of elongate setae on the ventral surface of the hind legs in males. This genus is difficult to separate from large and variable Eulechriops. Coturpus can be further separated from Turcopus and Cylindrocopturinus by genitalic characters given by R.S. Anderson (1994).

The genus has been collected on mistletoe, Phoradendron Nutt. (Santalaceae) on Quercus (R.S. Anderson 1994: 484).

One (R.S. Anderson 1994).

Mexico.

Eulechriops pictus Schaeffer, 1908 [by monotypy].

Masculine.

Cylindrocopturinus can be differentiated from genera in the Eulechriops complex of genera by the tibial apex, which has a modified inner flange that resembles the premucro (Fig. 25), a rostral channel that is laterally carinate on the mesoventrite and not closed posteriorly by carina (Fig. 7), and no striolate profemoral patch (a similar mesoventrite has been seen in Copturomorpha, but members of this genus always have a striolate profemoral patch).

Sleeper’s (1963) key to U.S. genera inexplicably contains two couplets that lead to Cylindrocopturinus: couplet 6a leads to genus “7. Cylindricopturinus, new genus” and couplet 8a leads to genus “7. Cylindrocopturinus, new genus”. Couplet 6a is where C. pictus Schaeffer actually would key out to (in order to reach couplet 8a, couplet 4a would have to be selected, which states that the mesoventrite is not excavated, leading to couplet 8, where Cylindrocopturinus is differentiated from Cylindrocopturus for having such excavation). The use of the epithet in couplet 6a is the first appearance of the name and is spelled differently than the usage in the remainder of the text. Subsequent authors (Kissinger 1964, Hespenheide 1984, Anderson 1994) did not choose among the original spellings as a First Reviser (International Code of Zoological Nomenclature article 24.2.3) but used the spelling “Cylindrocopturinus” in their work. Cylindrocopturinus is thus selected here as the correct original spelling, making “Cylindricopturinus” an unavailable name as an alternative original spelling.

R.S. Anderson 1994: 463.

See “Phylogenetic relationship” section for Coturpus. Hespenheide (1984: 315) suggested a relationship of Cylindrocopturinus with Zurus (= Copturus), Euzurus, Microzurus, Mnemyne Pascoe, 1880, and Paramnemyne, citing the “...structure of the mesosternum and procoxae, the small size of the tarsal claws, and form of the antennae...” as indicative of a closer relationship to Microzurus. The genera Cylindrocopturinus, Coturpus, and Turcopus were proposed by R.S. Anderson (1994) to be related to Eulechriops. As interpreted here, those genera, plus Macrolechriops and Copturomorpha, compose a group of lechriopines, the majority of which contain a combination of the following characteristics: unarmed and non-carinate hind femora, a rostral channel defined laterally by carina, and a second antennal funicular article that is subequal to or shorter than the first.

Species of Cylindrocopturinus have been collected on various species of Phoradendron on species of Quercus, Acacia Mill. (Fabaceae), Juniperus L. (Cupressaceae Gray), and Ipomoea L. (Convolvulaceae Juss.) (R.S. Anderson 1994).

Four (Hespenheide (1984) described one species, R.S. Anderson (1994) described two).

U.S.A.: AZ, Mexico, Honduras.

Eulechriops erythroleucus Faust, 1896 [by subsequent designation: Sleeper 1963: 215].

Masculine.

Figures 79–82. 

Lateral and dorsal habitus images of Lechriopini. 79a–b Eulechriops minutus [ASUHIC0024145] 80a–b Euzurus ornativentris [ARTSYS0000800]. 81a–b Hoplocopturus varipes [SSAC0001086] 82a–b Lechriops vestitus [SSAC0001114]. Scale bars = 2 mm.

Eulechriops is a large and variable genus that can be identified from related genera by the following combination of characters: antennal funicular article 2 is not longer than 1; the mesoventrite has a carinate channel that can terminate on the meso- or metaventrite, but when closed on the mesoventrite it is not pronounced and sharply carinate (as in Turcopus); the femora are not carinate and are ventrally unarmed; the profemora lack a striolate patch (as in Copturomorpha); the pronotum lacks a strongly convex, arcuate pronotal carina (as in Macrolechriops), and the hind legs are not sexually dimorphic (as in Cylindrocopturinus and Coturpus).

The genus was erected by Faust (1896: 91, in footnote) to accommodate Lechriops that have unarmed, non-carinate femora; Marshall (1922: 70) notes the lack of interdependence of these two characters. The type species was not originally designated by Faust and is given by Sleeper (1963) as the Venezuelan species E. erythroleucus, which was the first of three species described by Faust under his new genus – this was apparently overlooked in subsequent catalogs where the type species is listed as undesignated. Eulechriops is possibly the most diverse genus of Conoderine when considering the large number of undescribed species (Hespenheide 2007), as well as the most speciose genus of Conoderinae in the Caribbean region with sixteen species described by Hustache (1932a) from Guadeloupe.

Champion 1906b: 110 (for Central America), Hustache 1931: 285 (for Guadeloupe), Hespenheide 2003: 95 (for two U.S. species).

The genus is likely related to the genera Copturomorpha, Macrolechriops, Cylindrocopturinus, Turcopus, and Coturpus, most of which have a combination of the following characters: unarmed, non-carinate metafemora, a funicular article 2 that is not longer than article 1, and the rostral channel of the mesoventrite with relatively parallel longitudinal carinae. The species recognized as Eulechriops are unlikely to represent a monophyletic group (Hespenheide 2005b) and the species currently recognized in the genus can only be identified to Eulechriops by not having the distinguishing characters of the aforementioned related (and smaller) genera.

Mostly unknown. Two of the three U.S. species and related Mexican species (of the E. minutus species group of Hespenheide 2003) are associated with Quercus (Sleeper 1963, Hespenheide 2003). Several undetermined Central American species have been reared from Cecropia, Coussapoa Aubl. and Pourouma Aubl. (Urticaceae) (Jordal and Kirkendall 1998: 159, LaPierre 2002). Some South American species have been reared from Rubus L. (Rosaceae Juss.) (Hespenheide 2005), Manihot Mill. (Euphorbiaceae Juss.) (Monte 1938) and Gossypium L. (Malvaceae) (Barber 1926).

Fifty-seven species are known from the focal region (with one described by Hespenheide 2007 and one fossil species described by Poinar and Legalov 2013) and an additional 31 species are known from South America (Wibmer and O’Brien 1986: 264, with one more described by Hespenheide 2005b).

U.S.A., Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Dominican Republic (Poinar and Legalov 2013, fossil), Guadeloupe; South America.

Euzurus ornativentris Champion, 1906 [by original designation].

Masculine.

The single species of Euzurus can be identified by the posteriorly produced lobe of the pronotum concealing the scutellum from above, the distinctly structured mesoventrite (Fig. 8) which has lamellae extending anteriorly from the posterior modification, and the large tubercle on the metaventrite anterior to the anteroventral border of the metacoxa (Fig. 65). The second antennal funicular article is longer than the first, the eyes are vertical and separated, the metafemur is laterally bicarinate and ventrally toothed, and the tibial uncus is short and curved.

With the genera Copturus and Microzurus, Euzurus shares a concealed scutellum (but see “Phylogenetic relationships” section for Copturus), Type II sclerolepidia (Lyal et al. 2006: 229), and modification to the mesoventrite that does not have anteriorly extending carinae. The separated, vertical eyes are similar to some species of Cylindrocopturus and Poecilogaster (Figs 46, 49). Champion (1906b: 46) mentions a similar appearance to Macrocopturus (Eucopturus) Heller.

Unknown.

One.

Costa Rica, Panama.

Copturus armatus Gyllenhal, 1838 [by original designation].

Masculine.

Figures 55–66. 

Miscellaneous morphological structures. 55 Hoplocopturus sulphureus [ARTSYS0000801] with an overlay showing an elongate second funicular article that is more than twice the length of the first 56 Lissoderes subnudus [SSAC0001064] with an overlay showing a second funicular article that is subequal to the first, and seven distinct funicular articles 57 Philinna bicristata [ARTSYS0000799] showing a funiculus composed of six articles 58 Microzurus sp. [SSAC0001290] showing a short, slender fifth tarsal article and minute claws 59 Philides comans [ARTSYS0000804] with a broad tooth at the base of the tarsal claw 60 Left metafemoral apex of Piazurus trifoveatus [SSAC0001118] that is unarmed at the mesal and lateral faces 61 Left metafemoral apex of Cylindrocopturinus pictus [SSAC0001288] with a tooth only at the mesal face 62 Left metafemoral apex of Peltophorus polymitus seminiveus [SSAC0001117] showing a small tooth at both mesal and lateral faces 63 Left metafemoral apex of Hoplocopturus sulphureus [ARTSYS0000801] showing a small tooth at the lateral face and an acuminate process at the mesal face 64 Metaventrite of Pseudopiazurus centraliamericanus [SSAC0001291] showing deep, arcuate sulc 65 Lateral view of the metathorax of Euzurus ornativentris [ARTSYS0000796] showing a large tubercle in anterior to the metacoxa 66 Dorsal view of the profemora of Copturomimus caeruleotinctus [SSAC0001059] showing denuded, striolate regions.

As originally distinguished from the rest of the New World conoderines by Heller (1895) in his key to genera, Hoplocopturus, Mnemynurus, and the South American Balaninurus Heller, 1895 have an arcuate carina on the mesoventrite. This character separates Hoplocopturus and Mnemynurus from the Central American lechriopine genera that have an elongate second funicular article, carinate and ventrally toothed femora and modification to the mesoventrite, especially the often very similar looking species of Lechriops and Macrocopturus. From Mnemynurus caloderes Heller, 1895 (the only species of Mnemynurus at the time), Heller (1895) differentiated Hoplocopturus by the length of the rostrum and the corresponding rostral channel – in his key, Hoplocopturus species have a rostrum that does not extend beyong the mesocoxae, while Mnemynurus caloderes has a rostrum channel and rostrum that extends to the first abdominal ventrite. Champion (1906b) added several species to each genus, mentioning that “Hoplocopturus is connected to Mnemynurus by intermediate forms, and it can only be separated therefrom by the shorter rostrum” (Champion 1906b: 53). Most species of Mnemynurus described by Champion (except M. longispinis Champion, 1906) have a rostrum that extends at least to the posterior margin of the metaventrite, and species of Hoplocopturus (except H. javeti Champion, 1906 and H. nigripes Champion, 1906) have a shorter rostrum. Those species of Hoplocopturus with a rostrum that reaches near the middle of the metaventrite are additionally difficult to separate from Mnemynurus because they have a similar coloration to all described Mnemynurus (the “red-eyed fly” mimicry complex) and the region of the mesoventrite posterior to the arcuate carina is invaginated under the carina (not simply a semicircular depression as in other Hoplocopturus). Despite these exceptions, the following characters can be used to separate many of the species of Hoplocopturus from Mnemynurus: rostrum shorter (never reaching the posterior margin of the metaventrite) and the rostral apex cylindrical (apically flattened and dilated in Mnemynurus). The hind femora of both genera can be carinate or not and the femoral apex typically has an elongate spine at the mesal face.

Some species (e.g. H. varipes Champion, 1906) have a mesoventrite densely covered in scales and the distinguishing carina is difficult to see. Most species of Hoplocopturus are not clear members of a mimicry complex with the exception of a few species belonging to the “blue-thorax” complex and two described and several undescribed belonging to the “red-headed” fly-mimicking complex (Hespenheide 2005), but all described (and almost all observed undescribed) species of Mnemynurus belong to the “red-headed fly” complex. The species that have the “red-headed fly” coloration pattern are the most difficult to separate from Mnemynurus.

Champion 1906: 53 (for Central America).

Hoplocopturus is very similar to Mnemynurus and Balaninurus, sharing with those genera the arcuate carina on the mesoventrite. Some species are very similar looking to certain Lechriops and Macrocopturus (e.g. H. sherrywernerorum Hespenheide, 2009 and H. costatipennis Champion, 1906, respectively), but the structure of the mesoventrite easily separates the species of Hoplocopturus from those genera.

Some Hoplocopturus (e.g. H. varipes Champion, 1906) can be found on the upper and lower surface of the large leaves of Xanthosoma Schott (Araceae Juss.) [SSAC0001086], a plant family association with Araceae has also been made in Mnemynurus (see below). One species has been collected on treefalls of Sterculia L. (Malvaceae: Sterculioideae Burnett) [STRI_ENT_0082473, SSAC0001292].

Eighteen species are known from the focal region (with one more described by Hespenheide 2009) and an additional 12 species are known only in South America (Wibmer and O’Brien 1986: 270).

Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama; South America.

Rhynchaenus sciurus Fabricius, 1801 [by original designation].

Masculine.

Most species of Lechriops can be distinguished by the following combination of characters: the second antennal funicular article is longer than the first, the mesoventrite has a rostral channel that is bordered laterally by carinae, and the metafemora are carinate and ventrally toothed (Champion 1906b: 91). The anterior margin of the metaventrite is also usually excavated to receive the apex of the rostrum (Fig. 10; Champion 1906b: 91, Hespenheide 2009: 334), and the region of the mesoventrite lateral to the longitudinal carinae is often with dense multifid setae.

Many species have a white elytral sutural spot (as in Fig. 82b), but this is not exclusive to Lechriops (see Hespenheide 2009).

See Hespenheide 2003: 351 (for the seven U.S. species) and Champion 1906b: 91 (for Central America). Also Sleeper 1963: 210 (for U.S. species), Blatchley and Leng 1916: 418 (for Northeastern U.S. species, as Gelus) and LeConte and Horn 1876: 260 (for U.S. species, as Piazurus).

Some species look superficially very similar to species of Eulechriops, Macrocopturus, and Hoplocopturus, but the above combination of characters will separate most species. Champion (1906b: 91) considered Lechriops to be very close to Macrocopturus and in both genera he described species similar to the other genus (L. copturoides Champion, 1906 and Macrocopturus furfuraceus (Champion, 1906), the latter of which “forms a sort of connecting-link between Copturus [=Macrocopturus] and Lechriops” (Champion 1906b: 69)).

Some species in the U.S. and Mexico (the L. californicus species group of Hespenheide 2003) are associated with various species of conifers in the genera Pinus L. and Pseudotsuga Carrière (Pinaceae Spreng. ex Rudolphi). Some Central American species have been reared from petioles and stems of Cecropia and Coussapoa (Urticaceae) (Jordal and Kirkendall 1998: 159, LaPierre 2002). The Puerto Rican Lechriops psidii Marshall, 1922 is known to feed on guava fruits (Myrtaceae: Psidium guajava L.) (Marshall 1922: 70), but the placement of that species in Lechriops is suspect (though no specimens have been observed) due to the unmodified mesoventrite and lack of a femoral tooth.

Forty-nine species are known from the focal region [including two more described by Hespenheide 2003] and an additional 42 species are known exclusively from South America (Wibmer and O’Brien 1986: 263, including four more described by Rheinheimer 2011].

Canada, U.S.A., Mexico, Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, Puerto Rico, Guadeloupe; South America. A new species of Lechriops was recently described from India (Khairmode and Sathe 2015), though the position of the species in this genus or in the tribe Lechriopini is doubtful.

Copturus satyrus Gyllenhal, 1838 [by subsequent designation: Wibmer and O’Brien 1986: 20].

Masculine.

Figures 83–86. 

Lateral and dorsal habitus images of Lechriopini. 83a–b Macrocopturus lynceus [SSAC0001085] 84a–b Macrolechriops spinicoxis [ARTSYS0000529]. 85a–b Microzurus championi [ASUHIC0031507] 86a–b Microzygops nigrofasciatus [ARTSYS0000802]. Scale bars = 2 mm.

Most of the specimens of this very large and variable genus can be identified by the combination of a second funicular article that is longer than the first, an unmodified mesoventrite, a carinate and ventrally toothed hind femur, and the absence of a profemoral striolate patch. A few Central American species (and several more in South America – Heller’s subgenera Lamellocopturus and Cyphocopturus) have modification to the mesoventrite similar to Microzygops (e.g. Macrocopturus albidus (Champion, 1906)) but Champion did not assign them to one of Heller’s subgenera. Some smaller species can be difficult to distinguish from Lechriops but species of that genus in general tend to have a less spherical head (somewhat obliquely flattened above the rostral base) with comparatively smaller, more strongly acuminate eyes in addition to the modification to the mesoventrite.

All species included in this genus were treated as Copturus until Wibmer and O’Brien (1986: 17), see “Notes” section for Copturus above for a clarification of name use. Heller (1895) divided the genus Copturus into six subgenera or species groups (including Macrocopturus), reiterating Pascoe’s (1880: 494) comment on his own struggle with adequately constructing a subgeneric classification for this genus: “Copturus illustrates the difficulty of defining a large genus. Every character is liable to exception, not one appearing to have a generic value, although there is a common interresemblance which is not to be mistaken; the group, in fact, is a natural one, but which perhaps might, for the advantage of the systematist, be artificially divided into several genera.” Heller noted the probable superficiality of his subgenera.

Macrocopturus is the most widespread genus in the Caribbean, being the only genus of Conoderinae recorded from the Bahamas and Jamaica. Four different putative mimicry complexes are present in Macrocopturus as presently defined (Hespenheide 1995). Immature stages are described for Macrocopturus aguacatae (Kissinger, 1957) by Muñiz Vélez (1958) and M. burserophagus Muñiz-Vélez & Ordóñez-Reséndiz, 2010 by Muñiz-Vélez and Ordóñez-Reséndiz (2010).

Some of the described species are very similar in appearance to the following genera: Cylindrocopturus (e.g. the Mexican M. burserophagus which is placed in Macrocopturus because of the presence of a ventral femoral tooth), Copturomimus (which have a striolate region on the profemora), Lechriops, Hoplocopturus, and the South American genera Damurus Heller, 1895 (Champion 1906b: 69, in footnote) and Copturosomus (Champion 1906b: 69). Of those genera, Copturomimus, Lechriops, and Hoplocopturus are considered related in this paper, and Cylindrocopturus, which is currently in the Zygopini, is also probably a related lechriopine. The few observed specimens of South American Copturosomus are very similar to Macrocopturus and the genus Damurus was not observed in the course of this study, but was considered by Heller (1895: 55) to be related to Timorus (also currently in the Zygopini) which Champion (1906: 33) in turn considered related to Macrocopturus.

Champion 1906b: 69 (for Central America) and Heller 1895: 19 (for Central and South America).

Macrocopturus floridanus (Fall, 1906), known as “the mahogany notcher” (Morton 1987: 191) is wood-boring as a larva and as an adult feeds on foliage of mahogany (Meliaceae Juss.: Swietenia mahogany (L.) Jacq.) (Morton 1987). Other species are known from avocado (Lauraceae: Persea) (Kissinger 1957: 7, Muñiz V. 1965), Bursera citronella McVaugh & Rzed. (Burseraceae Kunth) (Muñiz-Vélez and Ordóñez-Reséndiz 2010), and Cecropia, Coussapoa, and Pourouma (Urticaceae) (LaPierre 2002).

Fifty-six species are known from the focal region, including one described by Hespenheide (1984), three by Zayas (1988), one by Muñiz-Vélez and Ordóñez-Reséndiz 2010, and one by Hespenheide (2017), and one species transferred from Archocopturus by Hespenheide (2005). An additional 101 species are known from South America (Wibmer and O’Brien 1986: 272, with two additional species described by Rheinheimer (2011)).

USA: FL, Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Bahamas, Cuba (Zayas 1988), Jamaica, Puerto Rico, Guadeloupe (Hespenheide 1984); South America.

Macrolechriops spinicoxis Champion, 1906 [by monotypy].

Masculine.

Macrolechriops belongs in the Eulechriops complex of genera with its short second funicular article, non-carinate and unarmed hind femora, and carinate and excavated mesoventrite, but can be distinguished (at least the Central American species) by the hump-like pronotal carina.

Champion reported the presence of a “...flattened, conical prominence on the intermediate, as well as on the anterior, coxae...” (1906b: 127) as being unique among the conoderines he examined. This character, however, has been observed in other species and genera, including some Eulechriops, reducing the diagnostic utility of that character among the putative relatives of Macrolechriops.

The specimen in Fig. 84 agrees with Champion’s description, which was based on “one worn specimen” (Champion 1906b: 127), but direct comparison of the specimen with the holotype is needed to confirm the identity as no other identified material of that species has been observed in the course of this study.

The South American species of Macrolechriops described by Hustache have a pronotum that is strongly convex but without a hump-like carina. This hump-like pronotal carina is known from other genera, such as Macrocopturus verrucosus (Champion, 1906), but none described or so far known in the genus Eulechriops or genera closely related to it, making it a useful character for separating the only currently known Central American species of Macrolechriops from its relatives. A very similar vestiture pattern and pronotal shape has been observed in other genera, most notably in a species of Copturomimus which has been seen in several collections incorrectly identified as Macrolechriops spinicoxis.

Champion (1906b: 126) notes the similarity with the South American Machaerocnemis Heller, 1895 and Copturosomus, but the genus is here considered part of the Eulechriops genus complex.

s. Unknown.

One species is known from the focal region and five additional species are known from South America (Wibmer and O’Brien 1986: 266, Rheinheimer 2011).

Mexico, Honduras [ARTSYS0000529]; South America.

Microzurus rhombus Heller, 1895 [by monotypy].

Masculine.

Microzurus can be differentiated from Copturus by the lack of a ventral tooth on the profemora, a thin fifth tarsomere with minute tarsal claws (Fig. 58), and costate elytral intervals. Champion (1906: 89) described two species that have shallow or absent modification to the mesoventrite – no material was observed of the species without modification (M. edentatus Champion, 1906), but the species would still be easily recognized as a Microzurus by the concealed scutellum, minute tarsal claws, and absent ventral tooth on the pro- and mesofemora. The second funicular article is not longer than the first and the hind femora are carinate and ventrally toothed. The observed species have a similarly apically laterally flanged receptacle of the mesoventrite as in Copturus, though it is usually much less prominent.

Hespenheide (1984) suggested a relationship between Microzurus and Cylindrocopturinus. Microzurus is here interpreted as closely related to Copturus, but the position of those two genera within the lechriopines is uncertain. See entry on Copturus.

Hespenheide (1984: 316) reported the possibility of seed-feeding based on label data. Costa-Lima (1956: 219) mentions South American species on fruits of Campomanesia Ruiz & Pav. and Psidium guajava (Myrtaceae).

Three species are known from the focal region and an additional four species are known from South America (Wibmer and O’Brien 1986: 266).

Mexico, Belize, El Salvador, Honduras, Panama; South America.

Microzygops nigrofasciatus Champion, 1906 [by original designation].

Masculine.

Microzygops can be distinguished from other lechriopine genera with an elongate second funicular article and modification to the mesoventrite by the following characters: the shape of the eyes, which are very large and contiguous in bottom 2/3 and widely separated in top 1/3 (Fig. 45), the distinct form of the mesoventrite (Fig. 11) which is similar only to the few species of Macrocopturus that have modification to the mesoventrite with an elevated posterior margin of the mesoventrite and faint, arcuate longitudinal carinae (as well as a deeply excavated anterior margin of the metaventrite), and the color pattern, which is putatively ant mimetic (Hespenheide 1995) but distinct from the other ant mimics with orange-brown ground color and transverse black fascia of the pronotum and elytra.

The metafemora are very elongate, extending well past the abdominal apex and lacking carina. The pro- and mesofemora are ventrally toothed, but the metafemur is unarmed in Microzygops nigrofasciatus though with a small tooth in the South American M. flavatus Rheinheimer, 2011 and one undescribed Central American species [SSAC0001210]. Other generic characters given by Champion (1906b: 46) include the “exserted head” and “cylindrical constricted prothorax”.

The species M. nigrofasciatus like the species of several other genera originally described as monotypic, is possibly a complex of several species – specimens identified to that species have been observed from Mexico to Peru and at a range of elevations.

The exserted head, elongate and slender hind legs, proportionately short and narrow third tarsomere, and linear carina of the vertex of the head, in combination, is only similar to Pseudolechriops and, to a lesser extent (excluding the head characters) Lissoderes, but the mesoventrite is distinct in each of those genera. Microzygops was among the genera moved from the Zygopini to the Lechriopini in Lyal et al. (2006), but the position of the genus within the Lechriopini is at present uncertain.

Unknown.

One species is known from the focal region and one additional species is known from French Guiana (Rheinheimer 2011: 68).

Mexico [ASUHIC0031512], Costa Rica [ASUHIC0086639], Panama; South America.

Mnemynurus caloderes Heller, 1895 [by monotypy].

Masculine.

Figures 87–90. 

Lateral and dorsal habitus images of Lechriopini. 87a–b Mnemynurus poeciloderes [ARTSYS0000803] 88a–b Paramnemyne decemcostata [ASUHIC0065104] 89a–b Poecilogaster brevis [ASUHIC0086631] 90a–b Pseudolechriops megacephalus [ASUHIC0086629]. Scale bars = 2 mm.

All of the described and most of the observed undescribed species of Mnemynurus are members of the red-headed fly mimicry complex (Hespenheide 1973, 1995), and are most difficult to distinguish from the similarly patterned species of Hoplocopturus and the South American Balaninurus which also share the distinctive inverted U-shaped carina on the mesoventrite. See “Diagnosis” for Hoplocopturus above for more information on the separation of the genera. The second funicular article is much longer than the first, the apex of rostrum is flattened and dilated, the hind femora are ventrally toothed and sometimes carinate, and the rostrum usually extends past the posterior border of metaventrite (though not in M. longispinis) and sometimes beyond the apex of the abdomen.

Mnemynurus, Paramnemyne, and the South American genus Mnemyne are the only New World genera having a rostral channel extending at least to the posterior half of the metaventrite. A distinct channel is found only in the larger Mnemynurus species and the metaventrites of other species are longitudinally depressed. Zayas (1988) described two species from Cuba, although based on the descriptions their placement in this genus is uncertain.

Champion 1906b: 49 (for Central America; M. caloderes in key = M. championi Heller, 1933 (Heller 1933: 150)), Heller 1932a: 5 (only to the three species described by Heller).

Most similar to the South American genus Balaninurus Heller, but generic limits between these two genera and Hoplocopturus need resolving – see Hoplocopturus above.

Species of Mnemynurus have been recorded as gall-inducing on young leaves of Philodendron Schott (Araceae) (Hanson et al. 2014: 503).

Nine species are known from the focal region including two described by Zayas (1988) and two additional species are known from South America (Wibmer and O’Brien 1986: 270).

Mexico, Honduras, Nicaragua, Costa Rica, Panama, Cuba [Zayas 1988]; South America.

Paramnemyne arcana Heller, 1895 [by subsequent designation: Rheinheimer 2011: 78].

Feminine.

Paramnemyne can be readily distinguished by the transverse carina near the posterior margin of the metaventrite, marking the end of the rostral channel (Fig. 12). The second antennal funicular article is several times longer than the first; the femora are not carinate, are ventrally toothed, and are unarmed at the apices; the mesopleura are small and non-ascending; and the eyes are completely separated with the greatest separation in the middle.

The unarmed femoral apices, small and non-ascending mesopleura, and piazurine type of mesoventrite suggest improper placement in the Lechriopini, however transferring the genus without observing the putative relatives of Paramnemyne, the South American genera Mnemyne (sec.Heller 1895: 11) and Paramnemynellus Hustache, 1932 (sec.Hustache 1932b: 207), would be remiss.