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Status of knowledge of the broad-nosed weevils of Colombia (Coleoptera, Curculionidae, Entiminae)

Broad-nosed weevils in the subfamily Entiminae (Coleoptera: Curculionidae) are highly diverse, not only in terms of number of species, but also in their sizes, forms and colours. There are eight tribes, 50 genera and 224 entimine species recorded from Colombia: seven genera and 142 species are considered endemic and only a handful of species, which are recognised as pests of Citrus or potatoes, are broadly known. The large diversity of this subfamily in the country is only superficially known and even though genus level identifications are generally achievable, species identification remains quite challenging, due in part to limited access to broadly-scattered basic information. Summaries of available information and bibliographic resources for each of the tribes represented in Colombia are offered, along with a checklist of the species of Entiminae recorded from the country, obtained from literature and a pictorial key for tribal recognition. New combinations are proposed for eight species of the genus Lanterius Alonso-Zarazaga & Lyal. Information on the distribution of entimine species in Colombia is compiled for the first time, including complete references to each original description and available taxonomic revisions. About a third of the species of Entiminae remain as recorded from the country without specific locality information. In addition, genus level distributional maps are presented, generated from data obtained from four Colombian entomological collections. Lastly, some challenges for entimine identification in Colombia, which likely extend throughout the Neotropical region, are briefly discussed. This contribution aims, in part, to facilitate and promote entimine research in northern South America. Neotropical Biology and Conservation 15(4): 583–674 (2020) doi: 10.3897/neotropical.15.59713 Copyright Jennifer C. Girón. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. REVIEW ARTICLE Jennifer C. Girón 584 Resumen Los gorgojos de pico corto en la subfamilia Entiminae (Coleoptera: Curculionidae) son altamente diversos, no sólo en términos del número de especies, sino además en tamaños, formas y colores. Existen ocho tribus, 50 géneros y 224 especies entiminos registradas en Colombia: siete géneros y 142 especies son consideradas endémicas, y sólo un puñado, las cuales son reconocidas como plagas de cítricos o papa, son ampliamente conocidas. La gran diversidad de esta subfamilia en el país sólo se conoce superficialmente y aunque identificar géneros es generalmente alcanzable, la identificación de especies sigue siendo un desafío, debido en parte al acceso limitado a información básica ampliamente dispersa. Se ofrecen resúmenes de información disponible y recursos bibliográficos para cada una de las tribus representadas en Colombia, junto con la lista de especies de Entiminae registradas para el país, obtenida a partir de la literatura, junto con una clave pictórica para el reconocimiento de tribus. Se proponen nuevas combinaciones para ocho especies del género Lanterius Alonso-Zarazaga & Lyal. Información sobre la distribución de las especies de Entiminae en Colombia se recopila por primera vez, incluyendo referencias completas a cada una de las descripciones originales y a las revisiones taxonómicas disponibles. Cerca de un tercio de las especies de Entiminae permanecen registradas para el país sin información específica de localidad. Adicionalmente, se presentan mapas de distribución a nivel de género, producidos a partir de datos obtenidos de cuatro colecciones entomológicas colombianas. Por último, algunas de las dificultades para la identificación de entiminos en Colombia, que probablemente se extienden a lo largo de la Región Neotropical, son brevemente discutidas. Esta contribución busca en parte facilitar y promover la investigación en Entiminae en el norte de Suramérica.

In taxonomic terms, the subfamily Entiminae has been recognised as such since Thompson (1992) and has been divided into as few as five tribes (Marvaldi 1997(Marvaldi , 1998 to as many as 55 tribes ; 54 according to Bouchard et al. (2011) is the most widely-accepted number of tribes. A summary of the taxonomic history of the Entiminae is provided by Marvaldi et al. (2014).

Biological generalities of entimines
Amongst curculionids, entimines are usually easy to recognise by the typical short and broad form of their rostrum, which is accompanied, in most cases, by conspicuous mandibular scars left by the breaking off of a deciduous process of the mandible at the time when the adult emerges from the pupal stage (Anderson and Howden 2002;see Fig. 1A). Entimines are phytophagous, trending to polyphagous and oligophagous; the larvae feed underground on the roots of plants, whereas the adults feed on the leaves (Marvaldi et al. 2014). Relatively few species of entimines (considering the whole diversity of the subfamily) are recognised as pests of economical importance; usually, a species only becomes a problem when populations are large. Especially relevant for Colombia are Compsus viridivittatus (Guérin-Méneville, 1855) (Guérin-Méneville 1855, 592), a Citrus pest in the tribe Eustylini (Cano et al. 2002a, b;Gallego et al. 2012, O'Brien andPeña 2012) and the 'Potato shooter' Leschenius vulcanorum (Kirsch, 1889) (Kirsch 1889, 17) in the tribe Naupactini (Canchala 1992;Peña 2001;Cortázar Gómez et al. 2012).
A thorough summary of the natural history and morphology (of both adults and larvae) of the Entiminae is provided by Marvaldi et al. (2014). Additional information regarding morphological features of the adults and the larvae can be found in Marvaldi and Lanteri (2005) and Marvaldi (1997Marvaldi ( , 1998, respectively. Most tribes are limited to particular biogeographic regions of the world (Marvaldi et al. 2014). Their distributions, in general, are more a reflection of habitat rather than host plant preferences (Anderson 1993(Anderson , 2018Marvaldi et al. 2014). The wide range of plants they feed on, paired with the strong association of species to particular habitats and geographic regions (e.g. Lanteri 1992; Anderson and Lanteri 2000;Franz and Girón 2009;Franz 2010b;Girón and Franz 2010;Franz 2011;Mazo-Vargas 2011, del Río andLanteri 2011a;Franz 2012, del Río et al. 2015), makes entimines a potential model group for biogeographic studies and the identification of areas of endemism. There is also evidence of altitudinal stratification of species (Girón and Franz 2010).
As a consequence of their diversity and abundance, entimines are commonly found in the field, including urbanised areas and are, therefore, frequently found in large numbers in biological collections (see Girón and Cardona-Duque 2018). There is also a trend to find large numbers of individuals of single economically-important species in cultivars, whereas only few specimens of several different species can be found associated with forested or less disturbed areas (pers. obs.). Furthermore, it is common that specimens in collections are only identified to family or, at best, sorted to subfamily, as is often the case in Colombia (see Girón and Cardona-Duque 2018). A right deciduous process still attached, left mandible with mandibular scar, maxillae fully covered by prementum (Naupactini) G mandibular scars well developed, maxillae partially covered by prementum (Anypotactus sp.) B, C frontal view of head indicating position of epistoma, nasal plate and mandibular scars: B Naupactini, with median furrow C Eustylini, with median fovea D-F lateral view of head and anterior section of prothorax: D Tanymecini, with anterior margin of prothorax straight and postocular setae clustered as a tuft E Eustylini, with anterior margin of prothorax slightly sinuate and without postocular setae F Lordopini, with anterior margin of prothorax strongly sinuate as to form welldeveloped postocular lobe, with postocular setae forming a fringe H-K profemora in frontal view (top end of drawings articulate to coxa, bottom end articulate to tibia): H Compsus sp., regularly shaped profemur I Eustylus sp., toothed profemur J Hadromeropsis sp. enlarged profemur K Anypotactus sp., clavate and toothed profemur L, M apical region of left metatibia (left margin on drawing is anterior (inner) in the beetle; right margin on drawing is posterior (outer) in the beetle), black triangles indicate posterior corner of metatibia: L Eustylini, posterior corner of metatibia rounded M Naupactini, posterior corner of metatibia angulate N-P anterior section of elytra in dorsal view: N shoulders absent O shoulders oblique, weakly developed P shoulders well developed.

Identification challenges
One of the most relevant references for Neotropical Broad-nosed weevils is the 'Biologia Centrali-Americana' by Sharp and Champion (1911), although it includes only scarce information on Colombian taxa. Even though many genera have been revised (see below under the treatment for each tribe), species identification of Neotropical and, especially northern South American, entimines remains highly challenging (e.g. Girón 2006). For Colombia, in particular, the fact that most type specimens are deposited in collections overseas, combined with the lack of identified material in national collections and compounded by a lack of specialists in the region, makes species identification a very difficult task at this time. In addition, one of the main difficulties for identifying entimine taxa is the limited access to information, which has been changing in the digital era. Many publications, including original descriptions and revisions are currently available online, some of them for free through the Biodiversity Heritage Library (https://www.biodiversitylibrary.org/). Nevertheless, some key publications with full references linking to those original descriptions (e.g. ) remain available only on paper and are essentially inaccessible to the broad community, especially in Latin American countries. An effort has been made in this paper to incorporate references and links to as many relevant publications as possible.
This contribution compiles and summarises the available information for the Entiminae recorded from Colombia. A general pictorial key to diagnose tribes is given. For each tribe represented in the country, a list of morphological characters for tribal recognition is offered, along with distributional information and bibliographic resources. A list of species is provided, including bibliographic records of distributional data within the country, where available. In addition, genus-level information has been recorded from specimens identified in Colombian national entomological collections, further providing information about distributions of entimines in the country.

Species list
The list of species of Entiminae recorded from Colombia, as well as general species distributions, were extracted from the annotated checklist of the weevils of South America by , the list of species of Entiminae by Morrone (1999), in addition to more recent revisions or new descriptions, where available. Tribal concepts and general distributions of genera are in accordance with Alonso-Zarazaga and . The full list of species is included here as Suppl. material 1 and available as a checklist via GBIF (Girón 2020, https://doi. org/10.15472/jdwfao); this online resource will be updated whenever new species are added to the Colombian fauna of Entiminae.
Only presence in Colombia was obtained from the main references Morrone 1999 and; therefore, original descriptions for each species were checked to obtain distributional records within the country. Localities taken from the 1800s' to mid-1900s' publications might not reflect current departments or municipalities (e.g. 'Bogotá' as a locality in 1857 does not strictly correspond to current 'Cundinamarca: Bogotá'). In those cases, locality is quoted as presented in the original description. Department was added for unambiguous localities (e.g. Antioquia was added to original localities recorded as 'Medellin'; Cundinamarca added for Fusagasugá; Meta added for Villavicencio, Boyacá added for Muzo). Localities in square brackets '[]' in the list of species indicate locality spelling in the original description. Type locality is recorded as 'Bogotá' for species described by Kirsch (1868) given the title of the paper: "Beiträge zur Käferfauna von Bogotà". A question mark '?' indicates that the locality record is doubtful and needs confirmation. Specific localities from countries other than Colombia are omitted here. For some of Schönherr's taxa, the page number corresponds to a column number in the publication and is indicated by 'col. ' (e.g. Entiminae Schönherr 1823, col. 1138.

Material examined
Specimens from entomological collections in Colombia were revised and distributional data at the generic level were recorded (Suppl. material 2). The insect collection abbreviations are adopted, for the most part, from Evenhuis (2020) as follows:

IAvH
Instituto Genus-level identifications were obtained using available keys (van Emden 1944a, b) and diagnoses (Sharp and Champion 1911) and confirmed by consulting experts in Curculionidae (Dr. Robert Anderson, Canadian Museum of Natural History, Ottawa, Canada and Dr. Charles O'Brien, USA), who have access to reference collections. The recognition section presented here for each tribe applies to entimines seen in Colombian collections and might not reflect the entire variation across tribes and genera. In most cases, there is more than one morphospecies with-in each genus at each collection. Species were not identified. There remain specimens recognised as entimines, but not identified to tribe. Relevant information for entimine tribes represented in Colombia was compiled and is presented here. A general pictorial key is provided here just as a way to narrow down identifications (Fig. 2). Preliminary identifications using this key should be checked against the lists of characters offered for each tribe for confirmation.

Figures
Line drawings were produced by tracing over photographs into drawing software. Figures are grouped to show morphological variation or to keep relevant tribal information together. Maps are based on data from collections. The record density map (Fig. 3) was created using R version 3.5.2 (R Core Team 2018), interfaced through RStudio version 1.1.463 (RStudio Team 2016) with the R package 'tmap' (Tennekes 2018). Distribution maps in Figs 3-10 were created using SimpleMappr (Shorthouse 2010); each map is accompanied by sketches of the general appearance of the head in dorsal view of some representative specimens of the tribe as seen in Colombian collections.
Anypotactines can be confused with some small Naupactini, from which they can be recognised by the shape of the apical region of the metatibia, which is rectangular and fringed by flat spines along ventral and posterior margins in Naupactini (Fig. 1M, rounded in Anypotactini, see Fig. 1L). The general appearance of some Anypotactini resembles members of Tanymecini ( Fig. 1D, with postocular setae) and can be differentiated, most of the time, by the lack of postocular setae in Anypotactini.
Diversity. The tribe contains 11 extant genera with 81 species described, all distributed in the New World O'Brien 1986a, 44, Alonso-Zarazaga andLyal 1999, 145). Seven genera represented by 11 species have been recorded from Colombia. Hypsometopus Kirsch, 1868(Kirsch 1868, represented by a single species, is considered endemic to the country; seven additional anypotactine species are considered endemic as well. General distribution. The tribe ranges from south-western USA to Argentina and Chile and some Caribbean islands, including the following countries: Argentina, Bolivia, Brazil, Chile, Colombia, Ecuador, Peru, Paraguay, Uruguay, Venezuela; Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama; Cuba, Dominica, Granada, Guadeloupe, Puerto Rico, Saint Vincent; SW USA (Texas) (Alonso-Zarazaga and . Most genera are distributed in Central America and northern South America, but the largest genus, Hyphantus Germar, 1824 (Germar 1824, 334;45 species;see Vaurie 1963), is distributed in southern Brazil, Argentina, Paraguay and Uruguay.
Bibliographic resources. The best resource available for the tribe as a whole is the Biologia Centrali-Americana (Sharp and Champion 1911, 215, as ' Anypotactina'). It includes a brief diagnosis of the tribe and diagnoses of the Central American genera and species (with some keys). There is a key in van Emden (1944b, 506) to identify the tribe, as well as a key to genera (van Emden 1944b, 510). The only genus of the tribe that has been revised is Hyphantus (Vaurie 1963), but it is not represented in Colombia.
Natural history. Except for a few plant-association records for Hyphantus (Vaurie 1963) and records of an unidentified species of Bothynodontes on Alnus jorullensis (Betulaceae) in Caldas and Baccharis sp. (Asteraceae) in Tolima (Bustillo and Villegas Isaza 1986), the natural history of Anypotactini remains largely unknown.
Remarks. A large proportion of anypotactine records in Colombia are concentrated in the mountainous regions (Fig. 4A). Some specimens amongst the undetermined Anypotactini from Caldas (Manizales, Río Blanco; MUSENUV) likely belong to an undescribed genus (Dr. Robert Anderson, pers. comm.). Specimens recorded from lowlands, especially from the north coast, share affinities with material seen from Central America at ASUCOB. One species of Anypotactus was included in a morphologybased phylogenetic analysis (Girón and Franz 2010), resulting as sister to Apodrosus Marshall, 1922(Marshall 1922aPolydrusini, exclusively Caribbean); no anypotactines have been included in molecular-based phylogenetic analyses to date. Recognition. Medium to large (approx. 7-45 mm), often very robust; scale coverage either nearly uniform and dense or forming patches across surface, highly variable in colouration including brown, white, purple and iridescent tones of green and blue; head (including rostrum) subrectangular, usually longer than wide ( Members of Entimini are generally easily distinguishable by their usually large size and the general bulky shape of their bodies [Cydianerus Schönherr, 1840(Schönherr 1840a, Entimus, Polyteles Germar, 1829 (Germar 1829, 358)] or the characteristic shape of their heads with narrow frons and rostrum with lateral margins apically diverging [Cydianerus, Rhigus Schönherr, 1823(Schönherr 1823, col. 1138. A more detailed diagnosis is offered by Vaurie (1951).

Tribe Entimini Schönherr, 1823
Diversity. The tribe contains seven Neotropical genera with 46 species described to date, two of them fossil O'Brien 1986a, 99, Gaiger 2001;Morrone 2002;Vanin and Gaiger 2005;. Three genera, represented by four species, have been recorded from Colombia, one of them fossil .
General distribution. The tribe ranges from Mexico to Argentina, with no representatives in the Caribbean islands. Entimini can be found in the following countries: Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Uruguay, Venezuela;Costa Rica, El Salvador, Honduras, Mexico, Nicaragua, and Panama (Alonso-Zarazaga and Lyal 1999). All genera are mainly represented in Brazil.

Remarks.
Even though members of Entimini are amongst the largest and most striking weevils in the subfamily, they are extremely rare in Colombian collections; only a few specimens of Rhigus have been found in the revised collections along with a few other unidentified Entimini. Romo and Morrone (2011) analysed biogeographic trends in the tribe. With the exception of the life cycle for Entimus nobilis (Olivier, 1790) (Olivier 1790, 525, Bruch 1932, the natural history of members of Entimini remains largely unknown. Plant associations have only been recorded for one species of Phaedropus (Vanin 1983) and one species of Rhigus (Gaiger 2001). There is one fossil species of Cydianerus recently described from Colombia ) and one fossil species of Entimus described from the USA (Scudder 1876). There is a recent study on the morphological variation and sexual dimorphism in Cydianerus latruncularius (Perty, 1832) (Perty 1832, 70, Regueira et al. 2020). Rhigus speciosus has been recently recorded from Colombia  and Peru (Gillett and Barr 2020). The type species for Entimus, Entimus imperialis Forster, 1771 (Forster 1771, 34), has been the focus of studies regarding the structural colouration of its scales which, as in other entimines, contain three-dimensional photonic crystals with diamond-type structure (e.g. Wilts et al. 2012). Polyteles Germar, 1829 (Germar 1829, 358) has not been recorded from Colombia, but its presence in the south of the country is likely, given that the genus is distributed in Argentina, Bolivia, Brazil, Ecuador, Paraguay, Peru and Uruguay. Fig. 6 Recognition. Small weevils (approx. 3-8 mm); scale coverage usually light to dark brown, forming patterns on dorsal surface; iridescent areas with blue or green scales or reddish or yellowish stripes may be present in some genera; setae along surface may also be present; surface of body smooth and even; head (including rostrum) subrectangular, usually nearly as long as wide (Fig. 6B, C); rostrum robust, usually nearly parallel-sided (Fig. 6B, C); nasal plate (see Fig. 1B, C) usually well developed; anterior margin of prothorax in lateral view strongly sinuate forming conspicuous broad postocular lobe (Fig. 6D); elytral shoulders moderately to well-developed (e.g. Fig. 1O, P); legs relatively stout, femora never toothed [except in Eurysaces Schönherr, 1840(Schönherr 1840b, not present in Colombia].

Tribe Eudiagogini LeConte, 1874
Eudiagogines are easily recognisable by their small size, well-developed postocular lobes and overall robustness. They could potentially be confused with anypotactines because of their size, but their postocular lobe sets eudiagogines apart. They can resemble some small Entimini, but eudiagogines have a comparatively shorter and stouter rostrum. Additional characters to define the tribe can be found in Lacordaire (1863, 384, in French).
Remarks. Promecops contains over 60% of the species and is also the most widespread genus of the tribe; some species are considered economically important in soybean cultivars in Argentina (Lázaro et al. 1997(Lázaro et al. , 1998 and Brazil (Rocha Barreto and Cavalet 2016). One species of Colecerus has been associated with cultivated Pecan trees in Mexico (Soto-Hernández and Barros-Barrios 2018). In North America, Eudiagogus has been associated with legumes in the genus Sesbania (Warner 1979); Eudiagogus has not been recorded from Colombia, but its presence in the south of the country is likely, given that the genus is distributed in Argentina, Bolivia, Brazil, Ecuador, Paraguay, Peru, Uruguay, Costa Rica, Honduras, Mexico and USA. Promecops is the only eudiagogine genus recorded from the revised Colombian collections; other eudiagogines were located, but remain undetermined (Suppl. material 2, Fig. 6A). Fig. 7 Recognition. Medium to large weevils (approx. 10-25 mm); scale coverage highly variable in presence, density and colouration; iridescent scales, erect setae or waxy secretions are frequently present; surface smooth and even or strongly sculptured and irregular; head (including rostrum; Fig. 7C-F) subrectangular, nearly as long or longer than wide; eyes small to mid-sized, slightly dorsally positioned; frons usually as wide as or narrower than interantennal distance, often bearing median fovea (see Fig. 1C); rostrum nearly parallel-sided or broadened apically; dorsal surface of rostrum with variable elevations or depressions, including longitudinal carinae or oblique fossae; antennal scrobe generally fully visible in dorsal view ( Fig. 7C-F); nasal plate (see Fig. 1B, C) usually well developed, either depressed, flat or elevated regarding surface of rostrum; anterior margin of prothorax in lateral view straight, seldom slightly sinuate (Fig. 1E), never forming conspicuous postocular lobe; postocular setae may be present, if so, forming a fringe instead of a tuft (compare Fig. 1D  cordaire, 1863 (Lacordaire 1863, 130), reduced in some Compsus and Exorides Pascoe, 1881 (Pascoe 1881, 43) (see Fig. 1N-P); tubercles and apical projections may be present on elytra; femora usually not toothed ( Fig. 1H; except in some Eustylus Schönherr, 1842(Schönherr 1842, Fig. 1I).

Tribe Eustylini Lacordaire, 1863
Eustylines are relatively easy to recognise amongst South American entimines by their size, frequently iridescent/bright colourations and the shape of their head ( Fig. 7C-F). They may be confused with similarly-coloured Naupactini, but the shape of the head, eyes, nasal plate and apex of metatibia [compare Fig. 1L Faust (1893, 16, in German). Marshall (1922b) discusses several classification issues with Compsus, Exophthalmus Schönherr, 1823 (Schönherr 1823(Schönherr , col. 1140) and allies, describes new genera and species and presents a key to identify species of Exorides (p. 203). Hustache (1938a, in French) revised part of South American Compsus; O'Brien and Peña (2012) translated to English and modified Hustache's key and provided identification and records for two species of Compsus recorded as Citrus pests in Colombia. Franz (2010a) re-described several type species in Eustylini and presented a morphology-based phylogeny of the so called "Exophthalmus genus complex", which involves several eustyline genera, especially Central American and Caribbean groups. Girón and Chamorro (2020) defined the "Compsus genus complex" and discussed affinities amongst species in this group. Some eustylines were included in phylogenetic analyses emphasising South American taxa by Marvaldi et al. (2018).
Remarks. The circumscription of Eustylini has been problematic since Lacordaire (1863), in French (some as 'Cyphides' p. 107, some as 'Geonémides' p. 125 and some as 'Eustylides' p. 205). Different genera, currently placed in Eustylini, have also been grouped together with members of Phyllobiini and Naupactini through time; most genera were treated as incertae sedis in the Coleopterorum Catalogus (Lona 1938: 508-525, 530-532). Kuschel (in Wibmer and O'Brien 1986a) brought together the majority of the genera that now compose Eustylini into a single tribe; the tribe is treated as 'Eustylina' by Morrone (1999). Some additions to the tribe have been made with transfers from other tribes, justified by a morphology-based phylogeny presented by Franz (2012). Marvaldi et al. (2018) made the most recent addition by placing Galapagonotus Lanteri, 2000 (Anderson andLanteri 2000, 3) in Eustylini; they indicate that Coconotus Lanteri, 2000 (Anderson andLanteri 2000, 6) might also be an eustyline, but this remains to be confirmed, as it also shares features with the geonemine genus Lachnopus Schönherr, 1840 (Schönherr 1840b, 380), particularly with species in the luctuosus species-group (see . Recent phylogenetic analyses (Franz 2012;Zhang et al. 2017;Marvaldi et al. 2018) indicate close relationships of Eustylini with taxa currently placed in Geonemini Gistel, 1856(Gistel 1856, which is primarily Caribbean and Central American. The generic limits between some eustyline genera are not clearly defined; this is particularly true for Compsus, Oxyderces Schönherr, 1823(Schönherr 1823, col. 1140, Exorides and Xestogaster Marshall, 1922(Marshall 1922b, all well represented in Colombia. This highlights the pressing need for taxonomic and phylogenetic revisions in the tribe, including (ideally recent) Colombian material that is already deposited in national collections. This particular group is part of the "Compsus genus complex" (Girón and Chamorro 2020); the authors compare several species of Compsus, Oxyderces and Exophthalmus which, by their colouration, may be confused with the golden-headed weevil, Compsus auricephalus (Say, 1824) (Say 1824), which is the only species of Compsus to occur in North America.
Members of Eustylini are amongst the most commonly found entimines in Colombian biological collections; Compsus is the most frequently observed and collected genus, followed by Exorides and perhaps Eustylus. Species of Exophthalmus are rare and seem to be restricted to the Pacific Region. Species of Compsus have been recorded as Citrus pests (Cano et al. 2002a, b;Gallego et al. 2012, O'Brien andPeña 2012) in Colombia and sugar cane in Venezuela (Kuschel 1955a). The Colombian endemic Oxyderces viridipes (Boheman in Schönherr 1840b, 179) has been intercepted at ports in the USA on Hydrangea flowers (Hydrangeaceae; see Girón and Chamorro 2020). In nature, eustylines commonly rest with their prolegs extended towards the front of the body. Fig. 8 Recognition. Medium-sized weevils (approx. 10-20 mm); scale coverage variable in presence, density, pattern and colouration; iridescent scales and setae may be present; surface usually even (not coarsely sculptured); head (including rostrum, Fig. 8B, C) subrectangular to conical, usually longer than wide; eyes usually large and elongated in lateral view, weakly to moderately projecting from surface of head; frons nearly as wide as or wider than interantennal distance, often bearing median fovea; rostrum nearly parallel-sided, broadened apically (Fig. 8B) or with lateral margins apically converging (Fig. 8C); dorsal surface of rostrum either flat or with one or three longitudinal carinae; antennal scrobe generally only visible at apical region in dorsal view; nasal plate (see Fig. 1B, C) usually well developed, flat or depressed regarding surface of rostrum; anterior margin of prothorax in lateral view strongly sinuate, forming conspicuous postocular lobe (Fig. 8D); postocular setae present, forming a fringe instead of a tuft (Fig. 8D); elytral shoulders absent to well-developed (see Fig. 1N-P); tubercles may be present on elytra; legs relatively slender.

Tribe Lordopini Schönherr, 1823
Amongst entimines with well-developed postocular lobe, lordopines are recognisable because of their elongated rostrum (Fig. 8B, D) or their conical heads with apically-converging lateral margins of the rostrum (Fig. 8C). They may be confused with mid-sized Entimini, but the bulky shape of the body of most Entimini and the overall shape of the head differentiate them (compare Fig. 5B, C with Fig. 8B, C).
Diversity. There are 44 genera with 291 species of Lordopini described (Wibmer and O'Brien 1986a). Eleven genera, represented by 33 species, have been recorded from Colombia, four and 25, respectively, being endemic.
General distribution. Lordopini is a strictly Neotropical tribe, ranging from Mexico to Paraguay, including some of the Lesser Antilles. Lordopies have been recorded from the following countries: Argentina, Bolivia, Brazil, Colombia, Ecuador Bibliographic resources. A general diagnosis for Lordopini can be found in Lacordaire (1863, 260, as 'Hypsonotides' , in French). Only two genera appear in the Biologia Centrali-Americana (Sharp and Champion 1911, 300, under 'Entimina'). Kessel (1932, 1935, 1937 revised several genera and provided keys to genera and species, focusing on Brazilian fauna, but these volumes are not yet available online and are rather difficult to find in libraries. Remarks. Lordopini is one of the least-studied entimine tribes in the region and needs systematic revision: 18 genera are Brazilian endemics; more than one third of the species belong to Hypsonotus Germar, 1824 (Germar 1824, 367, 115 spp.) and one fifth belongs to the genus Lordops Schönherr, 1823 (Schönherr 1823(Schönherr , col. 1142. From the remaining genera, 17 are monotypic and 11 are represented by only two species. All of these facts might indicate problems with current generic definitions. Furthermore, there are no phylogenetic analyses, including lordopines, to date. There are records of an undetermined species of Hypsonotus attacking peanuts in Brazil (Araujo et al. 1977); other than that, the natural history of the Lordopini remains largely unknown. Fig. 9 Recognition. Small to medium weevils (approx. 3.5-35 mm); scale coverage highly variable in presence, density and colouration; iridescent scales, erect setae or waxy secretions are frequently present; surface usually even; head (including rostrum; Fig. 9C-F) subrectangular to trapezoid, nearly as long or shorter than wide; eyes rounded to oval in lateral view, small to large, laterally positioned and moderately to strongly projected from surface of head; frons usually as wide as or wider than interantennal distance, often bearing median longitudinal furrow extending through rostrum; rostrum with lateral margins usually apically converging or nearly parallelsided, sometimes slightly broadened apically (Fig. 9C-F); dorsal surface of rostrum usually flat (excepting median furrow; broadly anteriorly depressed in Platyomus Sahlberg, 1823 (Sahlberg 1823, 29) and relatives, for example, Fig. 9F); antennal scrobe generally not broadly visible in dorsal view; nasal plate (see Fig. 1B, C) of variable development, usually flat to depressed regarding surface of rostrum; anterior margin of prothorax in lateral view straight, never forming postocular lobe; postocular setae absent; elytral shoulders absent to well-developed (see Fig. 1N-P); tubercles may be present on elytra; profemora sometimes enlarged (see Fig. 1J); apical region of metatibia rectangular, fringed by spines along ventral and posterior margins (Fig. 1M).

Tribe Naupactini Gistel, 1856
Naupactines are relatively easy to recognise amongst South American entimines by the overall shape of their head, the usual presence of a median longitudinal furrow along the head and rostrum and the characteristic apex of their metatibia. They may be confused with similarly-coloured Eustylini, but the overall shape of the head (compare Fig. 7C-F with Fig. 9C-F) and the apex of the metatibia differentiate them (compare Fig. 7C-F with Fig. 9C-F). Small naupactines may be confused with anypotactines, from which they can be differentiated by the metatibial apex. From Tanymecini, naupactines can be distinguished by the lack of postocular setae in Naupactini and the shape of the metatibial apex.
Diversity. Naupactini contains 67 genera with over 500 species described (Lanteri and del Río 2016a). Fourteen genera, represented by 36 species have been recorded from Colombia, two genera and 22 species being endemic.
General distribution. In the New World, Naupactini ranges from the USA to Argentina and Chile, including some of the Caribbean islands. Naupactines have been recorded from the following countries: Argentina, Bolivia, Brazil, Chile (including Isla de Pascua, Juan Fernández Is.), Colombia, Ecuador (including Galapagos Is.), French Guiana, Guyana, Paraguay, Peru, Uruguay, Surinam, Trinidad, Venezuela; Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama; USA (Alonso-Zarazaga and Lyal 1999). Its highest diversity is concentrated in Argentina, Bolivia, Brazil, Paraguay and Uruguay, which coincides with areas where they have been extensively studied.
Remarks. Naupactini is the largest tribe of New World entimines and one of the best known, mainly because of the revisionary and phylogenetic efforts of Dr. Analía Lanteri, Dr. Adriana Marvaldi, Dr. Guadalupe del Río and collaborators, based in Argentina. Different naupactine clades are specific to certain biogeographic regions in South America (see Lanteri 1992;del Río and Lanteri 2011a), with flightlessness and parthenogenesis being frequent in certain systems (Lanteri and Normark 1995;Guzmán et al. 2012;Lanteri et al. 2013;Lanteri and del Río 2016a). A detailed study of the female genitalia and oviposition habits in Naupactini, in a phylogenetic context, can be found in Lanteri and del Río (2008).
Most revisionary efforts to date have focused on fauna from Argentina, Brazil and surrounding countries. In northern South America, naupactines from Venezuela were studied by Bordón [1991, genus Macrostylus Boheman, 1840(Schönherr 1840a, genus Naupactus Dejean, 1821(Dejean 1821, 24 spp. described as new]; none of these species has been recorded from Colombia to date. Other than that, the naupactine fauna from northern South America, Central America and the Caribbean is still in need of revision. Several naupactine species are economically important (e.g. Lanteri et al. 2002Lanteri et al. , 2013; Leschenius vulcanorum (Kirsch, 1889) (Kirsch 1889, 17) has been recorded in Colombia as the 'Potato shooter' or 'tiroteador de la papa' (Canchala 1992;Peña 2001;Cortázar Gómez et al. 2012).
Members of Naupactini are very common in Colombian entomological collections, especially the genera Lanterius Alonso-Zarazaga &Lyal 1999, 164), Mimographus Schönherr, 1847 (Schönherr 1847, 34) and allies. Platyomus was recorded for the first time for Colombia by Girón (2007a; for Risaralda and Valle del Cauca) and then by Cardona-Duque et al. (2018; for Antioquia); these records might be erroneously identified and, instead, the specimens cited could be representatives of Chamaelops Kirsch, 1868 (Kirsch 1868, 235), which is fairly closely related to Platyomus (Lanteri and del Río 2016a); Chamaelops has previously been recorded for Colombia, considered endemic and includes two described species. Galapaganus Lanteri, 1992(Lanteri 1992 and Pantomorus Schönherr, 1840(Schönherr 1840a were also recorded by Girón (2007a; both from Valle del Cauca) and are presented here (see Suppl. material 2; including records from Bolívar for Galapaganus and from Casanare and Meta for Pantomorus), but these records need to be confirmed.

Tribe Premnotrypini Kuschel, 1956
Premnotrypines may not be easily recognisable as entimines, as their rostrum is relatively slender and usually directed towards the body when mounted; the reduction or absence of mandibular scars would make them pass as other miscellaneous Curculionidae. Amongst entimines, they can be recognised by their small size and well-deve loped postocular lobe, fringed by postocular setae. By their size, premnotrypines can be confused with anypotactines or small tanymecines, but can be recognised by the pre sence of a well-developed postocular lobe, lacking in both Anypotactini and Tanymecini.
Diversity. There are only three genera in Premnotrypini with 28 species (Kuschel 1956;Wibmer and O'Brien 1986a, 88). There is only one species recorded from Colombia.
General distribution. Premnotrypini is an exclusively Andean group, distributed from Colombia to Bolivia and northern Chile. Their diversity is concentrated in Bolivia and Peru, at elevations over 3000 m a.s.l. No specimens of this tribe were identified in the collections revised for this work.
Bibliographic resources. The tribal diagnosis, key to genera, descriptions of two of the genera and keys to all species can be found in Kuschel (1956, in Spanish).
Remarks. The genus Premnotrypes is associated with potato cultivars and contains economically-important species commonly known as the "Andean potato weevils" (see Alcázar and Cisneros 1998). There are many studies in the natural history and control of pest species (e.g. Alcalá and Alcázar 1976;Calvache 1987;Dueñas 1989), but no phylogenetic studies in Premnotrypini have been published to date. Fig. 10A-C Recognition. Small to large weevils (approx. 4-20 mm); scale coverage variable in presence, density and colouration, often dense and uniform, with colouration patterns formed by scales on dorsal surface; surface usually even, sometimes granulate; head (including rostrum; Fig. 10B, C) usually subrectangular and nearly as long as wide; eyes laterally positioned and weakly to strongly projected from surface of head; frons usually as wide as interantennal distance, bearing median fovea or small (narrow and short) longitudinal furrow that may extend along rostrum; rostrum subquadrate, usually with lateral margins parallel, sometimes slightly constricted basad of antennal insertion; dorsal surface of rostrum flat to slightly depressed; antennal scrobe only visible along anterior section in dorsal view; epistoma usually well-developed; nasal plate (see Fig. 1B, C) of variable development, flat to depressed regarding surface of rostrum; anterior margin of prothorax in lateral view straight (Fig. 1D), never forming well-developed postocular lobe; postocular setae usually present, clearly visible, grouped as a tuft, usually positioned near ventral margin of prothorax (Fig. 1D); elytral shoulders absent to well-developed (see Fig. 1N-P); tubercles may be present on elytra and elytral apices may be projected; profemora frequently enlarged (see Fig. 1J).

Tribe Tanymecini Lacordaire, 1863
Amongst Neotropical entimines, members of Tanymecini are easily recognisable by the presence of postocular setae, although not all tanymecines have this feature and not all the species with postocular setae are tanymecines. The postocular setae in Tanymecini are usually clustered, forming a tuft as opposed to distributed along the margin forming a fringe (compare Fig. 1D with Fig. 1F). Some tanymecines can be potentially confused with Naupactini because of their flat frons, median furrow and laterally-positioned eyes, but the presence of postocular setae in Tanymecini sets them apart easily; they can also be differentiated by the particular shape of the apex of the metatibia in Naupactini, which is rectangular and fringed by spines along the ventral and posterior margins (Fig. 1M; compare with rounded margins in tanymecines, Fig. 1L).
General distribution. Tanymecines are distributed throughout the Americas and the Caribbean islands, including the following countries: Argentina, Bolivia, Brazil, Chile, Colombia, Ecuador, Peru, Paraguay, Uruguay, Venezuela;Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama;Bahamas, Cuba, Dominica, Dominican Republic, Grand Cayman, Grenada, Guadeloupe, Haiti, Jamaica, Saint Vincent;C, E Canada, USA (Alonso-Zarazaga and Lyal 1999). Their highest diversity is concentrated in Central America and northern South America.
A key to genera recorded from Mexico was published by Cortés-Hernández and Morrone (2019), which includes most genera represented in Colombia. Howden's revision of the species of Pandeleteius of Colombia and Venezuela (Howden 1976) considerably increased the number of species known from Colombia and includes a key to species.

Remarks.
Tanymecini is one of the best-known tribes of entimines of the Americas, thanks to the works of Anne Howden. Tanymecines, especially Pandeleteius, are quite common in Colombia. In nature, tanymecines are commonly seen with their prolegs resting perpendicular to the body axis, a type of behaviour shared with some naupactines.

List of valid species of Entiminae recorded from Colombia
Genera and species in bold letters indicate endemic taxa. Fossil taxa are indicated by (+). Localities in bold letters indicate type locality. Localities marked with an asterisk are records taken from ASUCOB. A double asterisk means that identitications need to be corroborated. A question mark (?) in the distribution indicates that the locality record is doubtful and needs confirmation. Type localities from countries other than Colombia are omitted. Taxon names include author, year and page number of the original description (e.g. Anypotactus Schönherr 1840b, 299); for some of Schönherr's taxa, the page number corresponds to a column number in the publication and is indicated by 'col. ' (e.g. Entiminae Schönherr 1823Schönherr , col. 1138. See other locality remarks in the methods section.

Additional comments
This paper, as a compilation of information, constitutes the first step towards a better understanding of the biodiversity of entimines as a whole in Colombia and northern South America. It also highlights the areas and groups where a lot of work has been done and groups in need of revisionary work.
Due to its geographic location and attributes, Colombia harbours elements of different faunal components of the Americas: Andean, Amazon, Caribbean, Pacific and Orinoco (Ministerio del Medio Ambiente, Departamento Nacional de Planeación, Instituto Alexander von Humboldt 1996). Colombia is in a key position to contribute to the understanding of distributional, ecological and evolutionary patterns of Entiminae across the Americas, but remains as one of the most prominent knowledge gaps in the region for this subfamily and plenty of other insect taxa.
Except for economically-important species, there is a general lack of knowledge about the natural history of Colombian entimines (e.g. host plants, life cycles, natural enemies), which makes it very difficult to understand some of the patterns observed, for instance, the abundance vs. scarcity of some taxa. On the other hand, most entimines in Colombian collections are vegetation dwellers, whereas leaf litter, a highly diverse microhabitat for weevils (e.g. Anderson 2010), is essentially unexplored.
Most revisionary studies in northern South America have yielded numerous new species (Hustache 1938a;Howden 1976;Bordón 1991Bordón , 1997. With the notable exception of Howden (1976), not many taxonomic revisions of Neotropical entimines have included Colombian specimens; many of the specimens studied by Howden were collected directly by her and her colleagues in the early 1970s (Howden 1976). In the few recent revisions including Colombian material, examined specimens are housed in international collections and collected by early researchers in the times of "Nova Granada" (e.g. del Río and Lanteri 2007a, for Melanocyphus). This highlights a major issue for advancing the knowledge of Colombian entomofauna, in general: access to Colombian specimens by researchers is greatly limited. This fact, in turn, highlights other problems: (1) type material or even specimens correctly identified to species are extremely scarce (or non-existent) in national collections; (2) national experts (in Curculionidae), who would potentially be able to study type material abroad, are even more scarce, considering the high diversity of the group in the country (see Girón and Cardona-Duque 2018); (3) international experts essentially have no access to recent Colombian specimens, unless they visit Colombian national collections, which may not always be feasible; and, even if international researchers are able to visit, (4) national regulations make it very difficult to borrow specimens to take them out of the country for study. All of these issues combined, call for creating or strengthening international collaborations and adapting regulations to allow specimen exchange.
The good news is that digitisation efforts in libraries, and both in national and international biological collections, are contributing to improved access to specimens, data and expertise. Platforms, such as Biodiversity Heritage Library (https://www. biodiversitylibrary.org/), provide access to publications of original descriptions, museums are digitising their collections (entering data and imaging specimens, including types; for example, The Natural History Museum in London, (https:// data.nhm.ac.uk/search/entomology) and SiB Colombia (https://sibcolombia.net/; https://colecciones.biodiversidad.co/) is providing access to specimen data from national collections. In addition, platforms, like iNaturalist (https://www.inaturalist. org/home), are continuously generating records for Colombian species. The greatest limitation to learning and understanding Colombian biodiversity is the availability of national experts who are able to study specimens both in national and international collections. Identifiying entimines in Colombia is still a challenging task. Some of the specimens recorded here remain identified only to subfamily (Fig. 11) and are presented here in the hope that those can be revised and determined in the future, especially those from scarcely-explored regions.
Even though this work does not include all the entomological collections in the country and many remain to be revised, each collection revised here harbours a very unique diversity. It is evident that there is collecting bias towards the Andean region, with all other regions, but specially Amazon, Caribbean, Orinoco and the Islands (both Caribbean and Pacific), which are potentially as rich in diversity of species, poorly sampled sampled (see maps in Figs 2-11). The distribution of many species within the country is still to be determined: about one third of the species remain simply recorded from "Colombia" and about another third have been described from specimens collected in 'Bogotá' of the 1800s to early 1900s. Studying entimines in national collections and collecting them in poorly sampled regions will likely produce new records and taxa new to science all across the country. georeferencing all entimine records, revising and editing earlier versions of this manuscript and overall keeping me convinced that getting this compilation of information out in the world is valuable. Analía Lanteri has provided invaluable and critical feedback on several versions of this contribution over the years; thanks to her, the content of this document is much more accurate and comprehensive. The late Charles O'Brien and Nico Franz contributed their expertise, access to specimens and literature and encouragement at several points during the development of this work. Juan José Morrone also provided feedback and encouragement as a reviewer of the very first version of this manuscript, back in 2007. Alexander Ortiz facilitated the initial production of line-drawings. Ivon Babativa assisted databasing specimens at IAvH. My main motivation in producing this manuscript is to make information accessible and easy to find. It has taken me years to become familiar with literature about Colombian entimines and nowadays, I have the bibliographic resources, but living overseas, no access to Colombian specimens. I would like young researchers in Colombia to be able to quickly find information and to study entimine specimens right away! Many of the resources referenced here are available via The Biodiversity Heritage Library (https://www.biodiversitylibrary.org/), for which I'm also very grateful.