Cladistics and redescription of Hybreoleptops Kuschel (Coleoptera: Curculionidae: Entiminae) with the description of two new species from the Central Chilean subregion

Hybreoleptops Kuschel was established in a key to Chilean “Leptopini” for four species previously included in other genera. Those species are: H. tuberculifer (Boheman), H. aureosignatus (Blanchard), H. vestitus (Blanchard), and H. xanthomelas (Fairmaire and Germain). The genus is endemic to southern South America, inhabiting the southernmost part of Santiago Province (Central Chilean subregion) and Maule province, and the northernmost part of Valdivian province (Subantarctic subregion). All the species are recorded only from Chile, except H. tuberculifer which has been recorded both for Chile and Argentina. We undertake a cladistic analysis of the genus based on adult external morphology and genitalia. In the cladistic analysis two terminal taxa correspond to new species. The resulting cladogram confirms that these new species must be assigned to Hybreoleptops and lead us to redescribe the genus. Accordingly, we also describe the two new species, H. santiagensis Pérez and Posadas n. sp. and H. juanjosei Pérez and Posadas n. sp., and provide a key to the species of the genus.

The original description of Hybreoleptops by Kuschel (1949) was inserted in a key to the Chilean ''Leptopini''. The main characters used to define the genus were: enclosed corbels; stria 9 and 10 notably close to each other in the basal third; mentum glabrum; rostrum slightly wide, always forward directed; almost straight scrobes, directed toward lower edge of eye and notably visible in dorsal view; intervals 3 and 5 with one or two tubercles, tubercle on interval 5 bigger than the one on interval 3.
Hybreoleptops is endemic to southern South America. According to the biogeographical scheme proposed by Morrone (2001), Hybreoleptops inhabits the southernmost part of Santiago province (Central Chilean subregion) and the northernmost part of the Subantarctic subregion (Maule and Valdivian provinces). Figure 1 shows the distribution of the four known species of Hybreoleptops and the two new species described herein.
The main objectives of this paper are: (1) to undertake a cladistic analysis of Hybreoleptops; (2) to complement the original description of this genus; and (3) to describe two new species of this genus. Additionally, a key to the species is provided.
Measurements were made with an ocular micrometer in a stereoscopic microscope. Body length was measured dorsally, along the midline, from elytral apex to fore margin of prothorax. Photographs were made with a digital camera attached to a stereoscopic microscope and drawings were made with a camera lucida. Label data are cited in full for type specimens, each line separated by a slash and enclosing information from each label with square brackets.
The species were analysed cladistically based on morphological characters. Analysis of the data matrix (Table I) was carried out using an exhaustive search under equal weights in Hennig86 (Farris 1988) using the ie* command. Three taxa were used as outgroups: Leptopius Oke and the two species included in Megalometides Kuschel (M. cacicus Kuschel and M. discors Kuschel). The ingroup consisted of six species: H. aureosignatus (Figure 2A (Nixon 1999(Nixon /2000 was used for examination of character distribution.

Cladistic analysis
Data matrix analysis under equal weights results in a single most parsimonious cladogram (L583; CI569; RI573). The cladogram and the optimization of characters are illustrated in Figure 4 (only those characters with unambiguous optimization were illustrated). According to this cladogram the two new species belong to Hybreoleptops.
Scrobes widely visible in dorsal view, scales on distal half. Suprascrobal sulcus slightly developed. Mandibular scars subcircular and well-developed, located at apex of a cylindrical pedicel; scar on left mandible larger than that on right mandible. Funicular article 1 shorter than article 2 (articles 1/2 ratio: 0.5-0.8). Hind margin of prothorax and fore margin of elytra densely covered by multifid setae. Interval 3 with one declivital tubercle. Interval 5 with or without declivital tubercle (if present, larger than declivital tubercle on interval 3). Ventrites 1-4 completely covered by subcircular scales. Ventrite 5 mostly covered by subcircular or seta-like scales, some Hybreoleptops species could exhibit a middle triangular area lacking scales. Hind tibiae showing an enclosed corbel delimiting a flat lenticular area which could be smooth and bare or setose. Males with pro, meso, and metatibiae with well-developed mucro; females with pro, meso, and metatibiae with slightly developed mucro; metatibial mucro occasionally absent in some females. Female sternum VIII ovate, apex incised.

Etymology
The name refers to species distribution in the Santiago biogeographic province.

Diagnosis
This species is recognized by the following combination of characters: strongly evident, subcircular mandibular scars with a conical protuberance placed at apex of a cylindrical and well-developed pedicel. Dorsal rostrum surface with strong lateral convexities. Scape reaching middle line of eye when resting in scrobe. Prothorax showing anterior impression. Interval 7 protruding at base, resulting in protuberant humeri. Interval 3 with rounded, low, declivital tubercle; this tubercle slightly elongated on anterior-posterior axis, completely covered by white subcircular scales and exhibiting numerous translucid and costate seta-like scales. Ventrite 5 middle area lacking scales (integument exposed); both lateral areas of ventrite 5 completely covered with seta-like scales and setae. Lenticular area of metatibial corbel covered with setae.

Description of holotype
General habitus as in Figure 5. Body length 9.8 mm. General body shape ovate. Integument mostly covered by scales; black in exposed areas (e.g. rostrum apex). Body vestiture mainly constituted of subcircular scales, which cover most of the integument; subcircular scales ornamented with longitudinal ribs which are only visible up to 6100, in some areas as ventrite 5 vestiture constituted by white seta-like scales. Dorsal general colour of vestiture mixed, mainly of white scales, alternating with grey scales and sparse black scales. Ventral surface vestiture mainly of white scales. Additionally, tubercle on interval 3, abundant decumbent and translucid seta-like scales, especially at its apex. Base of interval 3, abundant decumbent and translucid seta-like scales. Dorsal surface of rostrum and prothorax, most elytra intervals, femora, and ventrites showing sparse decumbent and translucid seta-like scales. All these seta-like scales ornamented with longitudinal ribs visible up to 660. Each mandible with a well-developed, subcircular mandibular scar placed at apex of a cylindrical and tall pedicel which exhibits a conical protuberance; pedicel showing several long setae and white seta-like scales on its external side. Mandibular scar differing from all scars on other Hybreoleptops species (placed at apex of a short, cylindrical pedicel and lack conical protuberance). Rostrum 2.1 times longer than wide; dorsal surface convex with central longitudinal sulcus containing a very slightly developed central carina. Suprascrobal sulcus visible at each side of rostrum. Frons slightly convex, elongate fovea. Scrobes incurved, directed downward to lower margin of eyes. As in all species of the genus, distal half of scrobe presenting several scales (subcircular and seta-like). Eyes ovate and flat, with major axis dorso-ventrally orientated. Antennal scape reaching middle line of eye when resting in scrobe. Funicular articles conical; article 2 1.4 times longer than 1; article 3 shorter than 1 but longer than articles 4-7; articles 4-7 subequal in size. Club ovate, short, inflated. Prothorax subquadrangular, 1.25 times longer than its maximum width; presenting anterior impression and longitudinal sulcus; impressions ornating both sides of sulcus. Prothorax base showing a brush of multifid setae. Postocular lobes present. Scutellum rounded and small. Elytra 1.3 times longer than its maximum width; its anterior margin covered with a dense brush of multifid setae. Interval 7 convex at its base, resulting in protuberant humeri. Intervals 3, 5, and 9 strongly convex (specially at apical third of elytra). Interval 7 slightly convex; remaining intervals flat. Declivital tubercle on interval 3 low, rounded, and slightly enlarged following anteriorposterior axis of body. Declivital tubercle completely covered by white subcircular scales and apex presenting numerous, translucid and costate seta-like scales. Declivital area on interval 5 with similar vestiture. Intervals 3 and 9 fused at end resulting in a highly evident and elongated convex rib; this rib, extending diagonally to elytral apex. Elytral apex slightly acuminate, with abundant setae. Ventrites 1-4 completely covered with white subcircular scales, exhibiting sparse translucid seta-like scales and setae. Ventrite 5 with two broad lateral stripes completely covered by white seta-like scales; central area almost black because of the exposure of integument. Venter joints between ventrites 2 and 3, 3 and 4, and 4 and 5 profound and evident. Fore and hind edge of ventrite 1 and fore edge of ventrite 2 with multifid setae, specially abundant in fore edge of ventrite 1 and sparse on hind edge of ventrite 1 and fore edge of ventrite 2. Ventrites 3+4 3.2 times shorter than ventrites 1+2. Apex of tibiae 1 and 2 with comb of enlarged setae, comb continuing around all tibial apices, briefly interrupted on dorsal edge of apex. Tibia 3 with enclosed corbel surrounded by setae, defining an internal and external comb (following nomenclature by Thompson 1992). Lenticular area limited by these combs, flat and extremely setose. All tibiae slightly mucronate (at least in females). Female genitalia as in Figure 6.

Distribution
Hybreoleptops santiagensis is known only from its type locality (Chile, Curicó : El Coigo). This locality is located in the Santiago province which belongs to the Central Chilean subregion (see Figure 1), according to the biogeographic scheme proposed by Morrone (2001).

Etymology
The name refers to Juan José Morrone, our professor and friend, who has contributed enormously to the knowledge of the Andean weevil fauna.

Diagnosis
This species is recognized by the combination of the following characters: ovate to subcircular, flat mandibular scar, placed at apex of a cylindrical and well-developed pedicel; rostrum with slightly developed central carina; scape reaching posterior margin of eyes when resting in scrobe; intervals 5-7 convex at bases, resulting in protuberant humeri; interval 3 with rounded, low, declivital tubercle; slightly elongate on anterior-posterior axis, and covered mostly by white subcircular scales and numerous apical, translucid, and costate seta-like scales (ornamentation visible up to 660); ventrite 5 with central area smooth (integument exposed); both lateral areas of ventrite 5 covered by white, subcircular scales; tibia 3 with enclosed corbel, lenticular area covered by abundant setae.

Description
General habitus as in Figure 7. Body length 8.85 mm in female and 6.95 mm in male. General body shape ovate. Integument mostly covered by scales; black in exposed areas (e.g. rostrum apex). Vestiture mainly of subcircular scales, ornamented with longitudinal ribs visible up to 6100. Vestiture colour mixed; dorsal surface of body mainly of an irregular mix of white and iridescent pink scales. However, prothorax exhibiting more uniform distribution of colours as follows: two white longitudinal stripes at each side of prothorax median sulcus; each of these followed by a longitudinal pink iridescent stripe; each followed by a new white stripe. Basal area of interval 3 completely covered by white subcircular scales and decumbent, costate seta-like scales (ornamentation visible up to 660), this vestiture covering interval 3 from base to 4th punctuation and visible as a lighter stripe on elytra. Seta-like scales abundant on declivital tubercle on interval 3 and sparse on dorsal surface of rostrum and prothorax, most elytra intervals, femora, and ventrites. White scales covering venter area both in male and female. Mandibular scar present, strongly developed, subcircular to ovate in shape, lacking conical protuberance; placed at apex of a well-developed cylindrical pedicel; pedicel showing several long setae and white seta-like scales on its external side. Rostrum two times longer than wide, poorly developed central carina. Suprascrobal sulcus present. Frons slightly convex, elongated fovea (less evident in female). Scrobes incurved, directed downward to lower margin of eyes; as in all species of genus, distal half of scrobe presenting several subcircular scales. Eyes ovate and flat, longer axis dorso-ventrally orientated. Antennal scape reaching posterior margin of eye when resting in scrobe. Funicular articles conical; article 2 longer than 1 (approximately two times longer); articles 3-7 sub-equal in size and slightly shorter than article 1. Club ovate, short, inflated. Prothorax subquadrangular, 1.2 times longer than its maximum width in female and male; lacking anterior impression, with strongly developed median sulcus on dorsal surface, irregular impressions ornating both sides of sulcus. Postocular lobes present. Prothorax base with a brush of multifid setae. Scutellum visible and round. Elytra 1.5 times longer than maximum width in female and 1.6 times in male. A brush of multifid setae densely covering the anterior margin of elytra. Intervals 5-7 notably protruding at basis, resulting in protruding humeri. Intervals 3 and 5 strongly convex, especially at apical third. Intervals 7 and 9 just slightly convex. Short and rounded declivital tubercle on interval 3; this tubercle slightly elongated following the anterior-posterior axis and mostly covered by white subcircular scales with numerous translucid, costate seta-like scales at apex. Similar vestiture on interval 3 base and on declivital area of interval 5. Intervals 3 and 9 fused at end, resulting in an inconspicuous rib continuing toward the elytral apex. Elytral apex slightly acuminate and with abundant setae. Venter area completely covered by subcircular scales, excepting middle area of ventrite 5 lacking scales. Venter joints between ventrites 2 and 3, 3 and 4, and 4 and 5 notably deep. Ventrites 3+4, 3.5 times shorter than ventrites 1+2 in female and 3.2 times shorter in male. Apex of tibiae 1 and 2 with comb of enlarged setae, continuing around tibial apex, but briefly interrupted in dorsal edge of apices. Tibia 3 with enclosed corbel, surrounded by setae, defining an internal and external comb (following nomenclature by Thompson 1992). Lenticular area limited by combs, flat and setose. All tibiae slightly mucronate in female and strongly mucronate in male. Female genitalia as in Figure 8. Male genitalia as in Figure 9.  4. Scape reaching posterior margin of eye when resting in scrobe; prothoracic median sulcus absent (if present, discontinuous); base of intervals 3 and 4 with iridescent pink stripe which ends at the fourth or fifth puncture of the corresponding striae; base of interval 9 with a similar stripe; both declivital tubercle and the area between them coloured equal to basal area of intervals 3 and 4; basal area of interval 3 showing abundant costate seta-like scales . . . . H. aureosignatus -Scape reaching middle line of eye when resting in scrobe; prothoracic median sulcus starting at prothorax apex; base of intervals 3 and 4 lacking irridescent pink stripe; interval 9 lacking an iridescent pink stripe; both declivital tubercle and area between them not of iridescent pink colour; basal area of interval 3 not showing abundant costate seta-like scales . . . . . . . . . . . . . . . 5 5. Prothoracic median sulcus not reaching base; prothorax with anterior impression; femora integument completely covered by subcircular scales; vestiture uniform gold; elytra and prothorax lacking lateral white stripes; interval 5 with slightly developed tubercle behind declivital tubercle . . . . . . . . H. vestitus -Prothoracic median sulcus reaching base; prothorax lacking anterior impression; femora integument reddish and lacking scales; elytra and prothorax with lateral white stripes; remaining areas of prothorax and elytra black; interval 5 without slightly developed tubercle behind declivital tubercle . . . . .