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Systematic revision and cladistic analysis of Phacepholis Horn (Coleoptera: Curculionidae)

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... 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). ...
... 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. ...
... 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. ...
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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.
... Phacepholis Horn, 1876 was resurrected and revised by Lanteri (1990). At that time, it included five species mainly distributed throughout the Great Plains of the USA: Phacepholis elegans Horn, 1876; Phacepholis candidus Horn, 1876; Phacepholis obscurus Horn, 1876; Phacepholis viridis (Champion, 1911); and Phacepholis planitiatus (Buchanan, 1939). ...
... Previously, Phacepholis was treated as a subgenus or a synonym of Pantomorus Schoenherr, 1840 (Dalla Torre et al. 1936;Buchanan 1939;Blackwelder 1947;O'Brien and Wibmer 1982). The results of a phylogenetic analysis including most species of the Pantomorus-Naupactus complex from North and Central America (Rosas et al. 2011b) supported the hypothesis formulated by Lanteri (1990) that Phacepholis is an independent lineage of this complex, probably related to a group of Mexican-Central American species, which comprises the type species of Athetetes Pascoe, 1886 (Athetetes globicollis Pascoe, 1886) and most species assigned by Sharp (1891) to his Pantomorus group II. As a consequence of these results, Rosas et al. (2011b) considered Phacepholis as a senior synonym of Athetetes, enlarged the concept and the geographical range of the genus, and established new combinations for its species. ...
... pronotum more convex, particularly in males, lacking scales arranged in clusters surrounding setae, but usually with scattered large punctures on the sides; profemora broader (usually more than 1.5X as wide as metafemora); protibiae longer and with more denticles (9-16 instead of 7-9); metatibiae usually with a row of 5-6 denticles on the inner side, along half or distal third in males; and spermatheca subglobose instead of globose. Lanteri (1990) recognized the Pantomorus globicollis species-group as distinct and assigned it to Pantomorus "at least until the revision of the Central American species of the Pantomorus-Naupactus complex is completed". In that paper, the group included 10 species: Pantomorus globicollis (Pascoe, 1886) Champion, 1911. ...
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Phacepholis Horn is a lineage of the Pantomorus-Naupactus complex (Curculionidae: Entiminae: Naupactini) that evolved in North and Central America independently from the species traditionally assigned to Pantomorus Schoenherr. The genus includes 13 species, five distributed in North America and eight in Mexico and Central America. The Mexican-Central American species are Phacepholis globicollis (Pascoe), Phacepholis brevipes (Sharp), Phacepholis trituberculatus (Champion), Phacepholis albicans (Sharp), Phacepholis viridicans (Sharp), Phacepholis strabo (Sharp), Phacepholis howdenae Lanteri and del Río, new species (type locality: Zacapa, Guatemala) and Phacepholis burkei Lanteri and del Río, new species (type locality: Guerrero, Mexico). Phacepholis annectens (Sharp) is a junior synonym of P. brevipes, new synonymy. Lectotypes are designated for Pantomorus brevipes Sharp, Pantomorus annectens Sharp, Pantomorus trituberculatus Champion, Pantomorus albicans Sharp, and Pantomorus viridicans Sharp. Among the eight species of the P. globicollis species-group, six species are endemic to Mexico and two, P. strabo and P. howdenae, range into Central America. The Neotropical Phacepholis mainly differ from the Nearctic species by being larger, with longer antennae, the more convex male pronotum, and the spermathecae subglobose rather than globose. We provide descriptions or redescriptions of the eight species of Phacepholis from Mexico and Central America, a dichotomous key for their identification, habitus photographs, line drawings of male and female genitalia, and distribution maps.
... The taxonomic works of later authors (Kuschel, 1945(Kuschel, , 1949(Kuschel, , 1950Hustache, 1947;Voss, 1954;Bordón, 1991Bordón, , 1997 contributed to knowledge the diversity of the tribe Naupactini in the Neotropics, and the revisions and phylogenetic analyses published by other specialists have shed light on the relationships of several Neotropical genera and species (Lanteri, 1990a(Lanteri, ,1990d(Lanteri, , 1992(Lanteri, , 1995Lanteri & O'Brien, 1990;Lanteri & Morrone, 1991Lanteri & Díaz, 1994;Normark & Lanteri, 1998;Sequeira et al., 2000Sequeira et al., , 2008aSequeira et al., ,2008bScataglini et al., 2005;del Río et al., 2006Rosas-Echeverría et al., 2011a;Lanteri & del Río, 2016). Nevertheless, several problems still remain unresolved. ...
... Based on the hypothesis of nonmonophyly of Pantomorus sensu lato, some generic names considered as junior synonyms of Pantomorus in the old catalogues, e.g. Atrichonotus, Aramigus and Phacepholis, have been revalidated and treated as independent (Lanteri, 1990a;Lanteri & O'Brien, 1990;Lanteri & Díaz, 1994). ...
... Naupactines with these combinations of characters are usually widespread in treeless or almost treeless environments, e.g. P. elegans in the Great Plains of North America (Lanteri, 1990a); P. albosignatus in the Mexican Plateau (Rosas-Echeverría et al., 2011b); and the species of Aramigus, Atrichonotus, Eurymetopus, Parapantomous, N. cervinus and N. leucoloma in the steppes and prairies of South America (and in similar environments of other continents where they have been introduced). ...
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We examined the type material of 15 species of Naupactus Dejean and related genera (Curculionidae: Entiminae), most of them described by Alphonse Hustache (Museum National d'Histoire Naturelle, Paris) and Carl Boheman (Naturhisto-riska Riksmuseet, Stockholm). The information associated with the types was compared with the original descriptions, and large series of specimens were examined in order to analyze the variation of each species throughout its range (sexual dimorphism, other kinds of polymorphism or geographic variation). Based on the results of this study we establish seven new combinations, one new placement and nine new synonymies, we make 21 lectotype designations and provide three new country and state records. Alceis curtus (Boh.), Lanterius leucophaeus (Boh.), L. sparsus (Boh.), L. sellatus (Boh.), Parapantomorus carinirostris (Hust.), Symmathetes signatus (Blanchard) and Teratopactus acerbus (Boh.) are new combinations and T. sulphureoviridis Hust. is a new placement. The following names are senior synonyms of those between parentheses: Lanterius inermis (Hust.) (syn. Macrostylus ocellatus Lanteri); Naupactus auricinctus Boh. Hust.); and Trichonaupactus densior Hust. (syn. T. rexus Hust.). For each species we provide the following information: synonyms, available data on type specimens (sex, body length, geographic location, kind of type, and repository), complete geographic distribution, pictures of types or specimens compared with types, an explanation about the reasons that justify the nomenclatural acts.
... This tribe may not be monophyletic in its present definition because it includes some genera from Africa, New Guinea, North America and an extinct genus from the Baltic Amber [3], which probably do not belong to this tribe. However, all the Neotropical genera and a few ranging also in the Nearctic region (e.g., Phacepholis Horn, Ericydeus Pascoe) [4,5] are likely to have a more recent common ancestor, as suggested by previous phylogenetic analyses based on morphological and combined data [2]. ...
... Aramigus (South America) and Phacepholis (Central and North America), considered subjective synonyms of Pantomorus in some classifications (e.g., [13,51] and later revalidated [4,52], are monophyletic genera, which is in agreement with previous phylogenetic analyses [16,47]. They are grouped neither together nor with other Pantomorus, thus supporting the hypothesis that Pantomorus sensu Wibmer and O'Brien [13] is not monophyletic. ...
... Pantomorus sensu lato (including species from different areas of North, Central and South America) is an example of a non-monophyletic genus diversified in new adaptive zones or marginal areas within the range of this complex. Other groups undergoing diversification in marginal areas would have acquired exclusive synapomorphies, allowing the recognition of monophyletic genera, e.g., Phacepholis would have diversified along the western coast of Central America and the Great Plains of North America, and it is recognized by the particular shape of the spermatheca and the presence of a series of small denticles on the second ventrite of the male [4,16,20]; Aramigus and Eurymetopus have acquired several synapomophies in the female genitalia (particular shape of spermatheca, sternite VIII or ovipositor) and would have diversified in grasslands and steppes of South America [18,52,57]; and Galapaganus, which displays synapomorphies in the male genitalia (setae around the ostium), would have diversified along the western coast of South America and the Galapagos Islands [24,53,54,58,59]. ...
Article
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Naupactus (Curculionidae: Entiminae) is the most speciose weevil genus of the tribe Naupactini. The main objective of this work is to recognize species groups within Naupactus and to analyze the relationships between this and other Neotropical genera. For this purpose, we compiled a combined data matrix of 60 terminal units corresponding to 40 species for which we recorded 812 molecular and morphological characters (763 and 49 respectively), which were analyzed by Maximum Parsimony and Bayesian analyses. The single tree obtained from each analysis was rooted with Cyrtomon inhalatus. The species of Naupactus were recovered as different monophyletic groups, some of them closer to other genera of Naupactini (Lanterius, Teratopactus, Pantomorus and Parapantomorus) than to species of the same genus. We conclude that Naupactus is non-monophyletic, even though most species can be recognized based on a particular combination of morphological characters, which are probably symplesiomorphic. To be consistent with the cladistic principles, some genera diversified in marginal areas of the Pantomorus-Naupactus complex should be synonymized with Naupactus; however, these nomenclatural changes may not ensure a generic definition based on synapomorphies. We prefer to be conservative about the current classification until more evidence is available. The only nomenclatural amendments proposed herein are the transference of Naupactus inermis Hustache to Lanterius and of N. setarius to Symmathetes.
... Most of the earliest authors that applied Halffter's ideas to the distributional analyses of plant and animal taxa worked under a dispersalist framework (e.g., Evans 1966;Matthews 1966;Ball 1968;Rapoport 1968;Udvardy 1969;Reyes-Castillo 1970;Martins 1971;Mateu 1974;Reichardt 1977;Axelrod 1979;MacVean and Schuster 1981;. In addition, other analyses placed in a dispersal-vicariance framework have used cladograms to analyze the dispersal of particular taxa in the Mexican Transition Zone (e.g., Halffter 1988, 1990;Lanteri 1990;Daza et al. 2009;. ...
Chapter
The biota of the Mexican Transition Zone was assembled through the successive dispersal of four cenocrons from North and South America and their incorporation to the Paleoamerican biota, which was the original North American (Holarctic) biota that extended in Mexico in the Jurassic-Cretaceous. The Mexican Plateau cenocron dispersed to southern North America from South America (Gondwana) in the Late Cretaceous-Paleocene. The Mountain Mesoamerican cenocron dispersed from South America to the mountain forests of Central America and southern Mexico in the Oligocene-Miocene and then northward in the Pliocene. The Nearctic cenocron dispersed from northern North America to the mountains of the Mexican Transition Zone in the Miocene-Pliocene. Finally, in the Pleistocene the Typical Neotropical cenocron dispersed from the Neotropical region, being represented by genera widespread in South America. Based on the successive assembly of these cenocrons, three horobiotas are distinguished: Paleogene horobiota (original Paleoamerican biota plus Mexican Plateau cenocron), Neogene horobiota (Paleoamerican biota plus Mexican Plateau, Mountain Mesoamerican, and Nearctic cenocrons), and Quaternary horobiota (Paleoamerican biota plus Mexican Plateau, Mountain Mesoamerican, Nearctic and Typical Neotropical cenocrons)
... Most of the earliest authors that applied Halffter's ideas to the distributional analyses of plant and animal taxa worked under a dispersalist framework (e.g., Evans 1966;Matthews 1966;Ball 1968;Rapoport 1968;Udvardy 1969;Reyes-Castillo 1970;Martins 1971;Mateu 1974;Reichardt 1977;Axelrod 1979;MacVean and Schuster 1981;. In addition, other analyses placed in a dispersal-vicariance framework have used cladograms to analyze the dispersal of particular taxa in the Mexican Transition Zone (e.g., Halffter 1988, 1990;Lanteri 1990;Daza et al. 2009;. ...
Chapter
Several authors have considered that the complex area where the Neotropical and Nearctic biotas overlap corresponds to a transition zone. In the strict sense that is followed in this book, the Mexican Transition Zone includes the highlands of Mexico, Guatemala, Honduras, El Salvador, and Nicaragua north of Lake Nicaragua, whereas northern Mexico, the United States and Canada belong to the Nearctic region, and the lowlands of the Pacific coast and the Gulf of Mexico, the Yucatán Peninsula, and Central America belong to the Neotropical region. In a series of contributions, Gonzalo Halffter provided a coherent theory that explains how cenocrons that evolved in different geographic areas assembled in the Mexican Transition Zone. I review herein the historical development of Halffter’s theory, including the characterization of the dispersal or distributional patterns recognized by this author. These distributional patterns are considered to represent cenocrons, namely, sets of taxa that share the same biogeographic history and constitute identifiable subsets within a biota by their common biotic origin and evolutionary history. The biotic assembly of the Mexican Transition Zone is summarized into five stages, from the Jurassic-Cretaceous to the Pleistocene.
... Most of the earliest authors that applied Halffter's ideas to the distributional analyses of plant and animal taxa worked under a dispersalist framework (e.g., Evans 1966;Matthews 1966;Ball 1968;Rapoport 1968;Udvardy 1969;Reyes-Castillo 1970;Martins 1971;Mateu 1974;Reichardt 1977;Axelrod 1979;MacVean and Schuster 1981;. In addition, other analyses placed in a dispersal-vicariance framework have used cladograms to analyze the dispersal of particular taxa in the Mexican Transition Zone (e.g., Halffter 1988, 1990;Lanteri 1990;Daza et al. 2009;. ...
Chapter
The Mexican Transition Zone is the area where the Neotropical and Nearctic regions overlap. In its strict sense followed in this book, it corresponds to the moderate- to high-elevation highlands of Mexico, Guatemala, Honduras, El Salvador, and Nicaragua. This area is considered a transition zone between the Nearctic and Neotropical regions, so from the perspective of biogeographic regionalization, it should be assigned simultaneously to both regions. Within the Mexican Transition Zone, I recognize 5 biogeographic provinces and 22 districts. The Sierra Madre Occidental province is situated in western Mexico at elevations between 200 and 3000 m, with most of the area above 2000 m; it includes the Apachian and Durangoan districts. The Sierra Madre Oriental province is situated in eastern Mexico at elevations above 1500 m; it includes two subprovinces: Austral-Oriental (with the Saltillo-Parras and Potosí districts) and Hidalgoan (with the Sierra Gorda and Zacualtipán districts). The Transmexican Volcanic Belt province is situated in central Mexico, at elevations above 1800 m; it includes two subprovinces: West (with the Otomí and Tarascan districts) and East (with the Aztec and Orizaba-Zempoaltepec districts). The Sierra Madre del Sur province comprises south central Mexico at elevations above 1000 m; it includes three subprovinces: Western Sierra Madre del Sur (with the Jaliscian and Jaliscian-Manantlán districts), Central Sierra Madre del Sur subprovince (Michoacán district), and Eastern Sierra Madre del Sur (with the Guerreran and Oaxacan Highlands districts). The Chiapas Highlands province comprises southern Mexico, Guatemala, Honduras, El Salvador, and Nicaragua, basically corresponding to the Sierra Madre de Chiapas, from 500 to 2000 m altitude; it includes the Sierra Madrean, Comitanian, Lacandonian, Soconusco, Guatemalan Highland, and Nicaraguan Montane districts.
... Most of the earliest authors that applied Halffter's ideas to the distributional analyses of plant and animal taxa worked under a dispersalist framework (e.g., Evans 1966;Matthews 1966;Ball 1968;Rapoport 1968;Udvardy 1969;Reyes-Castillo 1970;Martins 1971;Mateu 1974;Reichardt 1977;Axelrod 1979;MacVean and Schuster 1981;. In addition, other analyses placed in a dispersal-vicariance framework have used cladograms to analyze the dispersal of particular taxa in the Mexican Transition Zone (e.g., Halffter 1988, 1990;Lanteri 1990;Daza et al. 2009;. ...
Chapter
Most of the authors involved in the theoretical development of evolutionary biogeography assume that dispersalism, panbiogeography, cladistic biogeography, and phylogeography represent alternative approaches. Instead, I consider that different biogeographic methods may be used to answer different questions, which are different steps of an integrative biogeographic analysis. This stepwise approach comprises five steps, each corresponding to particular questions and methods. Track analysis and methods for identifying areas of endemism are used initially to identify biotas (graphically represented on maps as generalized tracks or areas of endemism), which represent hypotheses of primary biogeographic homology and are the basic units of evolutionary biogeography. Then, cladistic biogeography uses available phylogenetic data to test the historical relationships between these biotas (secondary biogeographic homology). Based on the results of these analyses, a biogeographic regionalization is achieved. Intraspecific phylogeography, molecular dating, and fossils are incorporated to help identify the different cenocrons (set of taxa that share the same biogeographic history, which constitute identifiable subsets within a biota by their common biotic origin and evolutionary history) that became assembled in a biota. Finally, the geological and biological knowledge available is integrated to construct a geobiotic scenario that helps explain the way different dispersal and vicariance events contributed to biotic assembly and how the cenocrons dispersed to the biota analyzed. I present the concepts implied in these steps and some of the methods that may be applied to answer particular biogeographic questions and discuss how they can be integrated to explain biotic assembly within an integrative framework.
... Most of the earliest authors that applied Halffter's ideas to the distributional analyses of plant and animal taxa worked under a dispersalist framework (e.g., Evans 1966;Matthews 1966;Ball 1968;Rapoport 1968;Udvardy 1969;Reyes-Castillo 1970;Martins 1971;Mateu 1974;Reichardt 1977;Axelrod 1979;MacVean and Schuster 1981;. In addition, other analyses placed in a dispersal-vicariance framework have used cladograms to analyze the dispersal of particular taxa in the Mexican Transition Zone (e.g., Halffter 1988, 1990;Lanteri 1990;Daza et al. 2009;. ...
Chapter
A biogeographic transition zone is a geographical area of overlap, with a gradient of replacement and partial segregation between different biotas (sets of taxa sharing a similar geographic distribution as a product of a common history). It is an area where physical features and environmental conditions allow the mixture and co-occurrence of species belonging to two or more biotas, but also constrain their distribution further into one another. The biogeographic affinities of the taxa assigned to these biotas are the most fundamental information considered to analyze accurately biogeographic transition zones. Ecological biogeographers have plotted the frequency of different distribution patterns on maps, detecting gradual changes in their relative contribution to a given area and identifying the most heterogeneous places in terms of distributional patterns as transition zones. Evolutionary biogeographers have found transition zones particularly interesting for analyzing causal connections between evolutionary and geological processes at large spatial and temporal scales. Biogeographic transition zones constitute natural laboratories for investigating evolutionary and ecological principles shaping biotic assembly. Additionally, they represent places where different evolutionary lineages coexist, having important implications for conservation, particularly when they also exhibit high diversity.
... Most of the earliest authors that applied Halffter's ideas to the distributional analyses of plant and animal taxa worked under a dispersalist framework (e.g., Evans 1966;Matthews 1966;Ball 1968;Rapoport 1968;Udvardy 1969;Reyes-Castillo 1970;Martins 1971;Mateu 1974;Reichardt 1977;Axelrod 1979;MacVean and Schuster 1981;. In addition, other analyses placed in a dispersal-vicariance framework have used cladograms to analyze the dispersal of particular taxa in the Mexican Transition Zone (e.g., Halffter 1988, 1990;Lanteri 1990;Daza et al. 2009;. ...
Article
This book presents an evolutionary biogeographic analysis of the Mexican Transition Zone, which is situated in the overlap of the Nearctic and Neotropical regions. It includes a comprehensive review of previous track, cladistic and molecular biogeographic analyses and is illustrated with full color maps and vegetation photographs of the respective areas covered. Given its scope, the book will be of interest to students and researchers whose work involves systematic and biogeographic analyses of plant and animal taxa of the Mexican Transition Zone or other transition zones of the world, and to ecologists working in biodiversity conservation, who will be able to appreciate the evolutionary relevance of the Mexican Transition Zone for establishing conservation areas.
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