Iheringia, Série Zoologia

The definition of Tapeinini is discussed. A new genus with a new species, Wappesicus wappesi is described from Bolivia in Acanthoderini. Tapeina rudifrons Marinoni, 1972 is illustrated and its geographic distribution updated.

Santos-Silva & Nascimento (2018) called attention to the chaotic taxonomic status of some genera in Lamiinae (Cerambycidae) and to the problematic delimitation of its tribes, especially Acanthoderini and Desmiphorini.Through the study of both, new and already described species, it is possible to see the fragility of the limits between those two tribes, as defined by some authors (e.g.Lacordaire, 1872;Linsley & Chemsak, 1985).
Herein, the definition of Tapeinini Thomson, 1857, and the problematic status of Acanthoderini and Desmiphorini are discussed.New taxa and new distributional records are also provided.

MATERIALS AND METHODS
Photographs were taken in the MZSP with a Canon EOS Rebel T3i DSLR camera, Canon MP-E 65mm f/2.8 1-5X macro lens, controlled by Zerene Stacker AutoMontage software.Measurements were taken in ''mm'' using a measuring ocular Hensoldt/Wetzlar -Mess 10 in the Leica MZ6 stereomicroscope, also used in the study of the specimens.

RESULTS
Tapeinites Thomson, 1857:41. Tapeinitae;Thomson, 1860:73.Tapeininae; Pascoe, 1864:9.Tapeinini; Bates, 1881:191.According to Marinoni (1972) (translated): "Thomson, Bates, and Breuning related Tapeinini with Desmiphorini, while Serville, Laporte and Lacordaire indicated its relationship with Enicodini."However, this statement is not entirely true.Thomson (1857) only reported that (translated): "it would be difficult, in the present state of the classification of Cerambycids, to assign to the Tapeinites a place other than among the Lamiites."Thomson (1860:349) did not provide a clear relation between Tapeinini and Desmiphorini, however, the author demonstrates a certain similarity of both with old-world groups (translated): "It is incontestable that Enicodes attaches the Tmesisternites to the Saperdites by Desmiphorites, and especially by the Tapeinites… It can therefore be said that the Tmesisternites must be placed between the Saperdites and the true Lamiites."In the same work, Thomson (1860:71) by discussing on Desmiphoritae stated (translated): "Tapeinitis: 1, frons in male unarmed; 2, prothorax laterally spinose, different."Thomson (1864Thomson ( -1865) ) did not provide details on his insinuated relationship between Tapeinini and Desmiphorini.
According to Bates (1866): "This group was placed provisionally under the Saperditae, in the synopsis previously given of the Lamiaires.A more accurate examination has convinced me that it has closer affinities with the members of the Desmiphoritae.The Tapeinae, in fact, appear to be abnormally flattened forms of Exocentrinae."Currently, Exocentrini is a synonym of Pogonocherini.Monné (2018) listed Exocentrini in the synonymy of Acanthocinini (however he neglected to remove the old synonymy with this tribe) and Pogonocherini, Bouchard et al. (2011) listed it in the synonymy of Pogonocherini, and Tavakilian & Chevillotte (2018) considered Exocentrini as a distinct tribe (apparently, ignoring the synonymy proposed by Sama, 2008).Subsequently, Bates (1881) changed his opinion: "I am now inclined to doubt that Tapeina has any close affinity with the Exocentrini, the anterior acetabular sutures being less tightly closed than they are in the true members of that group; and I think their true place is near Eupogonius and allies in the Desmiphorini."Breuning (1950)  Still according to Marinoni (1972) (translated): "All the characters that are common to Tapeina and Peritapnia, also are common to the genera of Acanthoderini, such as: … mesocoxal cavities closed."However, in the redescription of Tapeina, he reported (translated): "Mesepimeron enters the coxal cavity."But this latter information is incorrect, because the mesocoxal cavities are closed, although the posterior angle of mesoventrite does not touch the anterior angle of the metaventrite (this shape of mesocoxal cavity more common in Desmiphorini than in Acanthoderini).
The main problem to correctly allocate Peritapnia Horn, 1894, Tapeina, and Wappesicus gen.nov., is to understand the definition of Acanthoderini, Desmiphorini (including Estolini), and Pogonocherini, without mentioning several other tribes not occurring in the American continent.Linsley & Chemsak (1985) provided a key separating these three tribes.However, nearly all features used to separate them are only useful when studying North American genera, which makes the definitions of the tribes both incomplete and inaccurate.In that key, the first step where these tribes were separated is the alternative of couplet "13": "Antennae with scape clavate", leading to Acanthoderini; "Antennae with scape cylindrical or conical", leading to Desmiphorini, Pogonocherini, and Estolini.This is problematic as there are genera in Acanthoderini with the scape not or only very slightly clavate, such as Exalphus Restelo et al., 2001 andPycnomorphidiellus Tavakilian &Peñaherrera-Leiva, 2003; or genera with the shape of the scape very variable, from distinctly clavate to distinctly not clavate, as for example in Oreodera Audinet-Serville, 1835.In the same way, Pogonocherini were separated from Estolini (currently equal to Desmiphorini) as follows: "Body short and convex, long flying hairs often present, if elongate and flying hairs absent, pronotum laterally unarmed", leading to Pogonocherini; "Body elongate and parallel-sided, flying hairs absent; pronotum tuberculate laterally", leading to Estolini.A study of the genera currently included in Desmiphorini indicates that the body shape, presence or absence of long setae and/ or tubercle on sides of prothorax are useless as characters to separate the Desmiphorini from Pogonocherini, except for North America genera.The same problem occurs with nearly all tribes of Lamiinae because, their definitions are usually based on local genera.
When defining Acanthoderini, Linsley & Chemsak (1985) affirmed: "This tribe may be recognized by the clavate scape of the antennae, prominent lateral tubercles of the pronotum, short stout body form, almost separated eyes, clavate femora and by the fringed front tarsi of the males of most genera."The lateral tubercles of the prothorax may be very small, as in Amblysaphes Bates, 1885, or only a rounded protuberance as in Mundeu Martins & Galileo, 2008; and the body shape, and distance between upper and lower eye lobes, and femoral shape are all noticeably variable in Acanthoderini.Sama (2008) reported: "The tribe Exocentrini Pascoe, 1864 is usually regarded as a synonym of Acanthocinini (Aurivillius, 1923, Breuning, 1962).Exocentrus differ from palaearctic Acanthocinini by the characteristic shape of the last segment of the abdomen of females: the last two tergites are internally divided by a longitudinal, cartilaginous apodeme, beginning as a narrow lamina from the middle of the 4 th sternite and ending at the apex of the 5th where it is enlarged laterally.As written above an analogous structure exists in Ceroplesis and Crossotus (Ceroplesini) as well as in other genera of Laminae [sic] such as Idactus Pascoe, 1864 (Ancylonotini), Sophronica (Apodasyini) and Pogonocherus (Pogonocherini), and never in Acanthocinini.Exocentrus and closely related genera and subgenera must be consequently transferred to the tribe Pogonocherini.According to Švácha (pers.comm.), this systematic change is supported by the immature stages morphology."We do not know if this feature is really absent in all Acanthocinini but, probably it is another variable character.Even larval features, often indicated as differential for Acanthocinini, have been revealed as only generic variations (Sônia Casari, pers.comm.).We agree with Sama (2008) that the existence of Exocentrini is incorrect, but a reliable feature that allows separating Pogonocherini, Desmiphorini and Acanthoderini has not been found.According to Nascimento et al. (2019): "Lacordaire (1872) included Desmiphorini among the tribes with mesocoxal cavities open laterally, and Pogonocherini, Acanthocinini and Acanthoderini among the tribes with mesocoxal cavities closed laterally.However, the mesocoxal cavities in all genera of Desmiphorini examined by us are exactly as in the Pogonocherini and Acanthocinini: closed laterally (mesoventrite not touching metaventrite)."These authors concluded that all genera currently placed in Pogonocherini may be included in Desmiphorini and/or Acanthoderini.
The main arguments of Marinoni (1972) to include Peritapnia in Acanthoderini and to keep Tapeinini as a distinct tribe are (translated): "… Peritapnia has in common with Acanthoderini and not with Tapeina: the anterior border of the first abdominal sternum narrowly inserted between the posterior coxae and scape piriform.Plus, the intercoxal processes is narrow, convex and in a plane much higher than the anterior portion of the respective sternum."However, there are several problems with these statements.Firstly, the abdominal process is not narrow in Peritapnia (Figs 2, 6) as suggested.Rather, it is or may be very similar to that of Tapeina (Figs 10,14).As previously mentioned, the scape in Acanthoderini is very variable and frequently, it is not piriform.Although the prosternal process and mesoventral process are wide in Tapeina (always wider than coxal cavity), it is more variable in Peritapnia, and although it is always narrower than coxal cavity, it may be distinctly narrower or wider than half of the coxal cavity.Furthermore, the central area of the prosternal process in the new genus described here is slightly narrower than procoxal cavity, and the narrowest area of the mesoventral process is as wide as mesocoxal cavity.The "plane" of the intercoxal processes is variable in Peritapnia, but can be similar to that in Tapeina.For example, Chemsak & Linsley (1978) reported on their new species P. minima: "… intercoxal process [prosternal process] almost plane…" Finally, the new genus has features of Tapeina and Peritapnia: scape similar to Peritapnia; lateral tubercles of the prothorax acute at apex as in Peritapnia (rounded in Tapeina); intercoxal processes similar to Tapeina; meso-and metafemora nearly fusiform as in Tapeina (pedunculateclavate in Peritapnia); and abdominal process more similar to Peritapnia (slighter wider in Tapeina, but not so wide as suggested by Marinoni, 1972).
Based on the lack of reliable features, and especially by the existence of Wappesicus gen.nov., which clearly have intermediate features between Tapeina and Peritapnia, would be evident the synonymy, based on the characters currently proposed for Acanthoderini and Tapeinini.However, it is possible that Tapeina (Figs 9-18), Peritapnia (Figs 1-8), and the new genus  belong to Desmiphorini because the imperfectly closed mesocoxal cavities (mesoventrite not touching metaventrite), is much more common in this tribe than in Acanthoderini.Nevertheless, as indicated, this feature is variable in the species (not only genera) currently placed in those tribes.Thus, as it was not possible to know what defines and separates Acanthoderini and Desmiphorini, it is not possible to establish a synonymy with security.However, the understanding of the true separation between Acanthoderini and Desmiphorini will only be possible after an embracing study of all genera including in them, not only base in genera of a particular area of the World, as has been done, but including genera from several places.We are being conservative and including the new genus in Acanthoderini, due to the current inclusion of Peritapnia in this tribe.
Description, female.Body moderate-sized, flattened.Head not retractile; frons transverse, not forming transverse plate, slightly convex close to clypeus, V-shaped, centrally depressed toward vertex.Gena distinctly shorter than lower eye lobe.Eyes coarsely faceted; distance between upper eye lobes distinctly larger than twice width of one upper eye lobe.Outer side of mandibles with large triangular depression, with its apex surpassing middle of mandible.Last maxillary and labial palpomeres gradually narrowed toward apex.Antennal tubercles not forming plate laterally on scape ball.Antennae filiform, shorter than body; scape longer than antennomere III, slightly piriform, not reaching middle of prothorax; antennomeres III and IV subequal in shape and length.Prothorax transverse, with distinct, wide posterior constriction; sides strongly divergent from anterolateral angles to acute lateral tubercle, then strongly convergent toward base of posterior constriction.Procoxal cavities closed posteriorly.Prosternum slightly transversely sulcate near base of prosternal process.Prosternal process wide, with narrowest area slightly narrow than procoxal cavity, very slightly more elevated than surface of prosternum, strongly widened from middle toward apex, which is laterally lobeshaped, and with posterior margin widely concave; posterior half inverted V-shaped depressed centrally.Mesoventral process wide, moderately narrowed centrally (narrowest area about as wide as procoxal cavity).Mesocoxal cavities closed, but with apex of mesoventrite distinctly not touching apex of metaventrite.Scutellum proportionally small, posteriorly rounded.Elytra slightly longer than 1.5 time humeral width; parallel-sided from humerus to slightly after middle, then gradually, round narrowed toward sutural angle.Femora fusiform.Abdominal ventrite V strongly inverted, V-shaped and centrally depressed, with distal margin of this area narrowly elevated (plate-shaped).
Wappesicus wappesi Santos-Silva & Nascimento, sp.nov.Description, female.Head dark mostly reddish brown; narrow area surrounding eyes, distal area of genae, distal area of postclypeus, and median groove on frons dark brown; gulamentum reddish brown, gradually lighter toward posterior area; antennae dark reddish brown; mouthparts mostly light reddish brown, with palpomeres mostly yellowish brown; mandibles dark reddish brown on basal half, black on posterior half.Pronotum mostly dark reddish brown, narrowly dark brown anteriorly and posteriorly.Prosternum and prosternal process light reddish brown, slightly darker in sides of prosternum, with margins darkened.Central area of mesoventrite and mesoventral process light reddish brown, and sides of mesoventrite, mesanepisternum and mesepimeron dark reddish brown (margins of the ventrites narrowly darkened).Metaventrite mostly light reddish brown, gradually darker toward sides.Scutellum reddish brown centrobasally, brown laterally.Elytra mostly dark brown, with part of sutural margin dark reddish brown.Femora mostly dark reddish brown, with darkened irregular areas.Tibiae dark brown basally, gradually dark reddish brown toward apex.Tarsi mostly dark reddish brown, with darkened irregular areas.Abdominal ventrites mostly dark reddish brown, slightly darker laterally, and dark brown on distal area of ventrites I-IV.
Thorax.Pronotum moderately tumid in anterior 2/3 (except narrow area close to anterior margin), with one wide, subcircular, slightly elevated gibbosity each side, and one elongate gibbosity centrally, nearly fused with lateral ones; minutely, abundantly punctate, with coarser punctures interspersed (absent in lateral gibbosities), except smooth central gibbosity; with yellowish-brown pubescence not obscuring integument, except glabrous central gibbosity; with long, erect, thick, dark setae emerging from part of coarser punctures (Fig. 20).Sides of prothorax with yellowish-brown pubescence not obscuring integument, with sparse, long, erect dark setae (shorter than in pronotum).Prosternum minutely, sparsely punctate centrally, slightly striatepunctate laterally; with sparse, short, decumbent yellowish pubescence, slightly more abundant laterally, with long, erect, dark setae interspersed.Prosternal process with yellowish pubescence as on central area of prosternum, with long, erect, thick dark setae basally and close to lateral margins, not reaching distal area.Ventral surface of mesothorax with yellowish pubescence not obscuring integument, with sparse, long, erect, thick dark setae in sides of mesoventrite and lateral margins of mesoventral process.Ventral surface of metathorax with yellowish pubescence not obscuring integument, sparser toward central area of metaventrite; with sparse, long, erect, thick dark setae in sides and posterocentral area of metaventrite, and sparse, long, erect yellowish setae laterally (Fig. 21).Scutellum with sparse yellowish-brown setae, forming fringe with denser setae along margins (longer toward centrodistal apex).Elytra coarsely, abundantly punctate throughout; with short, decumbent yellowish-brown pubescence not obscuring integument, with abundant, long, erect, thick dark setae throughout (Fig. 22); erect setae not emerging from the punctures.Legs: femora and tibiae with abundant yellowish pubescence not obscuring integument, with long, erect, thick, dark setae interspersed.

Tapeina rudifrons
This species was originally described from Brazil (Goiás, Mato Grosso do Sul, Espírito Santo, São Paulo), Paraguay (Concepción), and Argentina (Salta, Formosa, Santiago del Estero).Di Iorio (2004) reported the species for the Paraguayan department of Alto Paraguay; Wappes et al. (2006) recorded the species in the Bolivian department of Santa Cruz; Wappes et al. (2013) listed it in the Bolivian department of Tarija; and Martins et al. (2011) recorded it in the Brazilian state of Maranhão.Originally, Marinoni (1972) listed some paratypes as being from the Brazilian state of Mato Grosso, which was true at that time.However, currently, the places where those paratypes were collected are in the Brazilian state of Mato Grosso do Sul.
We are taking this opportunity to illustrate a paratype female (Fig. 17), and another female (Fig. 18) collected in Bolivia.Although Marinoni (1972) had drawings of females, only a frontal view photograph of the head of a female paratype has been published.