HESPEROPHANINA

HESPEROPHANINA

Piezosecus Martins & Galileo, 2003 View in CoL ( Figs. 1 View Figure 1 B-1G, 2A-2B, 2H, 3A-3C)

Piezosecus Martins & Galileo, 2003: 225 View in CoL .

Remarks: Given the complexities inherent in taxonomic classification within Cerambycidae tribes, defining features of these taxa often pose challenges, particularly due to the reliance on genera restricted to specific geographic regions. To enhance clarity and avoid excessive cross-referencing, we have consolidated the primary definitions of Hesperophanini here. This task is complicated by the limited availability of some works, many of which are not accessible online or in English. Therefore, we have included translations of relevant descriptions from various sources, when necessary, to provide a comprehensive overview of Hesperophanini without requiring the reader to consult multiple references. Currently, establishing a precise definition for Hesperophanini is not feasible without a comprehensive revision. However, it is possible to delineate the differences between Hesperophanini , as currently defined, and Piezocerini .

Mulsant (1839) defined Hesperophanini as follows (translated): Prothorax without lateral tubercles, subdepressed or not very convex in some species, while globose in others. Maxillary palpi sometimes poorly developed. Eyes usually slightly notched. Femora compressed and not abruptly bulged into a club in those who have a slightly convex prothorax, sometimes clavate among those with a globular one. Elytra with sutural angle unarmed. Body usually elongated. Mulsant included the following genera: Asemum Eschscholtz, 1830 (currently included in Asemini ), Criocephalus Mulsant, 1839 (currently a junior synonym of Arhopalus Audinet-Serville, 1834 , and included in Asemini ), Solenophorus Mulsant, 1839 (currently a junior synonym of Stromatium Audinet-Serville, 1834 ),and Hesperophanes Dejean, 1835 .This original definition of Hesperophanini is problematic as it also included genera belonging to the tribe Asemini .

The key to the genera of Hesperophanini from Mulsant (1839) already made it possible to separate the two tribes: eyes very slightly notched, not surrounding the base of the antennae, leading to Asemum and Criocephalus ; eyes deeply notched, leading to Solenophorus and Hesperophanes . However, some other details of the original description do not align with the current species included in Hesperophanini . For example, the sutural angle of elytra may be slightly projected (e.g., some species of Stromatium and all of Anatinomma Bates, 1892 ) or may not be slightly projected (e.g., Austranoplium Chemsak & Linsley, 1963 ); and the prothorax may have lateral tubercle (e.g., Corupella Martins & Napp, 2007 and Cerasphorus Audinet-Serville,1834 ) or lack them (e.g., Hesperophanes ).

Lacordaire (1868) defined Hesperophanini as follows (translated): Ligula membranous, bilobed, or notched. Palpi short, with the maxillary palpi usually slightly longer than the labial palpi; apical palpomere triangular. Mandible short (except in males of Gnatholea ), arched, and acute apically. Head most often not very protruding; antennal tubercles weakly indented in most species; genae extremely short (except in a few Zoodes ). Antennae with more or less bristling, fine setae, rarely spiny, longer than the body, at least in males. Eyes coarsely faceted, voluminous, close together above, deeply notched; low- er eye lobes very large, strongly protruding above the antennal tubercles frontally. Prothorax with or without lateral tubercles, pronotum often tuberculate. Scutellum small. Elytra basally wider than the prothorax, more or less elongated. Legs of variable length; procoxae transversely oval or subglobose, more or less angular laterally; pro- and mesocoxal cavities open laterally. Mesoventral process curved backward, notched apically. Prosternal process rarely very narrow. Body usually elongated.While this definition remains one of the more precise for species in various regions worldwide, it poses certain difficulties. For example, in Ochrus Lacordaire, 1869 (originally described in Oemini ), the maxillary palpi are distinctly longer than the labial palpomeres, and some genera initially included by him are now excluded from Hesperophanini .

According to Gahan (1906) on Hesperophanini :"Gula without mentigerous process, except in Hesperophanes , in which it is very short; ligula membranous; eyes large, deeply emarginate; antennae ciliated, longer than the body in the ♂ [male]. Prothorax unarmed at the sides. Elytra rather long, more or less parallel-sided. Front coxae subglobular, more or less angulate at the side; their acetabula open posteriorly; the intercoxal process either very little or not at all dilated at the end. Acetabula of middle coxae extended to the epimera. First abdominal sternite not longer than the second except in the middle and at the sides. Wing-venation reduced by the disappearance either of vein Cu 2 or of the posterior branch of Cu 1, probably the latter; Cu 2 in that case having lost its connection with A 1 appears simply as a branch of Cu 1.″ This description agrees well only to specimens from the region studied by Gahan (1906). For example, the prothorax may or may not have distinct lateral tubercles and the abdominal ventrite 1 may be distinctly longer than ventrite 2 (e.g., Malcho Mondaca & Beéche, 2022).

Linsley (1962) defined Hesperophanini as follows: "Head moderately short; antennae ciliated, longer than body in male, segments simple or rarely spinose at apex, second segment short; eyes large, coarsely faceted, deeply emarginate; ligula membranous; palpi usually unequal in length. Pronotum rounded or tuberculate at sides; prosternum with intercoxal process not dilated behind coxae; anterior coxae subglobular, their cavities scarcely to broadly angulate externally, open posteriorly; intermediate coxal cavities usually widely open to epimera. Elytral apices rounded, usually unarmed. Legs with femora sometimes clavate.″ This is a description based on genera of Hesperophanini from North America. However, only a few features do not agree with genera from other regions, such as the shape of the elytral apex. It is worth noting that Hesperophanini sensu Linsley (1962) encompassed Eburiini , Graciliini , and Bothriospilini .

Finally, Martins (1999) briefly described Hesperophanini as follows (translated):Upper eye lobes present, separated by a distance equal to twice the width of one upper lobe. Lower eye lobes large, occupying entire side of head. Galea and lacinia developed; apical palpomeres securiform. Mentum as wide as twice its length, with transverse sulcus centrally. Ligula emarginate. Antennae filiform, unarmed. Scape subcylindrical, without basal depression. Antennomere III not longitudinally sulcate. Prothorax often without lateral tubercles. Prosternal process arched, without lateral projections. Procoxal cavities angulated laterally, open posteriorly. Mesoventral process with lateral projections, notched apically. Metanepisternum without glandular opening. Base of epipleura without spicule. Procoxae with slightly distinct tab. Femora not linear, unarmed apically. Parameres separated, distinct. Digestive tube of larvae with crop. This description applies only to South American genera and, even so, with exceptions. For example, the distance between the upper eye lobes may be much shorter than twice the width of one upper lobe (e.g., Daramus (Daramus) Fairmaire, 1892 , and Hespereburia Tavakilian & Monné, 1991 ; the latter pointed out in Martins & Galileo, 1999); antennomere III may be slightly dorsally carinate, as in Catoptronotum Zajciw, 1959 , and Hesperophanoschema Zajciw, 1970 , or even distinctly carinate, as in Liostola Zajciw, 1962 ; and antennae may not be filiform (e.g., Daramus ( Daramus )).

According to Linsley (1963), Piezocerini has the mesocoxal cavities closed laterally, while Martins (2003) reported them as open. However, our observations confirm that they are indeed closed ( Figs. 2 View Figure 2 F-2G).

While some genera within Hesperophanini exhibit the basal flagellomeres bicarinate dorsally, the carina is not strongly marked and not keeled as observed in Piezocerini . Additionally, in Piezocerini , the ventral surface of the basal antennomeres is often also longitudinally carinate, at least near the inner surface, a feature not observed in Hesperophanini . Furthermore, differentiation between Piezocerini and Hesperophanini can be based on at least two other features: the shape of the procoxal and mesocoxal cavities. The procoxal cavities in Hesperophanini ( Fig. 2B View Figure 2 ) are open laterally, sometimes strongly so, while they are closed in Piezocerini ( Figs. 2 View Figure 2 C-2E). In Piezocerini , at most, the procoxal cavities are open close to coxa, and the remaining surface is distinctly closed ( Fig. 2C View Figure 2 ). The mesocoxal cavities are variable laterally in the genera currently included in Hesperophanini : they may be closed (e.g., Liostola Zajciw, 1962 ); open (e.g., Turcmenigena Melgunov,1894 ); or imperfectly closed (e.g., Trichoferus Wollaston, 1854 ). Conversely, in Piezocerini , while they also show some variability, they are never perfectly open laterally ( Figs. 2 View Figure 2 F-2G).

We examined the mesocoxal cavities in 14 genera of Piezocerini . They are distinctly closed in: Acruspex Martins, 1976 ; Alienosternus Martins, 1976 ; Cicatrizocera Martins, 1976 ; Colynthaea Thomson, 1878 ; Gorybia Pascoe, 1866 ; Haruspex Thomson, 1864 ( Fig. 2F View Figure 2 ); Hemilissa Pascoe, 1858 ( Fig. 2G View Figure 2 ); Pharcidodes Martins, 1976 ; Piezasteria Martins, 1976 ; Piezocera Audinet-Serville, 1834 ; and Pseudocolynthaea Martins, 1976 . They are imperfectly closed in: Piezarina Martins, 1976 ; Thyellocerus Martins, 1976 ; and Zelliboria Lane, 1951 . Although we have not examined specimens of Migmocera Martins, 1976 , the figure included in the original description of the genus shows the mesocoxal cavity distinctly closed laterally. We do not know the shape of the mesocoxal cavities in Piezogenista Martins, 1976 . Based on figures of Migorybia santossilvai García, Botero & Martinez, 2019 , they are closed or, at most, imperfectly closed.

Linsley & Chemsak (1984) provided a clear explanation of the shape of the mesocoxal cavities: "There has been much confusion in the interpretation of this character, and we hope that our definition will aid in stabilization of usage. Figure 1 View Figure 1 diagrammatically illustrates the three conditions most commonly found. The upper and lower figures clearly indicate the open and closed situations.The middle figure shows the type of condition that has caused most of the past disagreement. In spite of the fact that a small opening exists between the coxal cavity and epimeron, the epimeron is not in direct contact with the cavity, and we consider this as being closed [herein, named imperfectly closed].Therefore, we define the cavities as open when the epimeron is directly between the two sclerites and in direct contact with the cavity.″ Since Piezosecus lacks distinct longitudinal carinae on both the dorsal and ventral sides of its antennae and possesses open lateral pro- and mesocoxal cavities, a feature absent in Piezocerini , we propose its transfer to Hesperophanini .

Sama (2008) divided Hesperophanini into two subtribes. Hesperophanini ( Daramina ) was defined as follows: "Similar to Hesperophanini [sic, Hesperophanina ] but mandibles without a fringe of hair along the inner edge; palpi unequal, maxillary palpi very long, 1 st segment hardly shorter than 2 nd; last segment of maxillary and labial palpi securiform, strongly dilated at apex, chiefly in male; ligula reduced, deeply bilobed, without lateral lobes; prosternal process subtriangular in front, laminiform between coxae, coxal cavities widely angulate laterally; mesonotum without stridulatory plate, with median endocarina; mesocoxal cavities widely open externally to epimera. Metendosternite with lateral arms elongate, longer than lateral laminae, which are short, moderately enlarged, truncate apically, divided by a deep notch. Male genitalia: ventral arc (IX sternite) fork shaped; dorsal arc (IX tergite) absent. Larva conspicuously elongate, with dorsal ampullae protruding.″ At least some of these features are present in American Hesperophanini , including in Piezosecus . However,based especially on the length and shape of the maxillary palpomeres, we are including Piezosecus in Hesperophanini ( Hesperophanina ).

According to Martins & Galileo (2003) (translated): Piezosecus belongs to the Piezocerina subtribe and is separated from all genera with open anterior procoxal cavities by the prosternal and mesosternal processes, which are markedly acuminated toward the apex. However, as defined and illustrated in figure 2 from Martins & Galileo (2003) (see Fig. 2H View Figure 2 ), the shape of the pro- and mesosternal processes is not a distinguishing feature. In fact, the shape and width vary within the type series ( Figs. 3 View Figure 3 A-3C).

Material examined: BRAZIL, Minas Gerais: Parque Estadual Rio Doce, 1 male ( MZSP 61420 View Materials ), 25.IX-13.X.2013, L. Migliore leg. ( MZSP). Espírito Santo: Córrego do Itá , 2 paratypes male ( MZSP 61418 View Materials ; MZSP 61419 View Materials ), X.1954, W. Zikán leg. ( MZSP) ; Santa Teresa (19°55′47.80″S, 40°44′50.90″W), [attracted to light], 1 male ( MZSP 61424 View Materials ), 2020-2021, F.Z. Madalon leg. ( MZSP). São Paulo: Araras, holotype male ( MZSP 61272 View Materials ), 13.X.1981, S.M. Nunes leg. ( MZSP) GoogleMaps .