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THE SPECIES OF EUCHEILOID PERICALINA: CLASSIFICATION AND EVOLUTIONARY CONSIDERATIONS (COLEOPTERA: CARABIDAE: LEBIINI)

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TL;DR

This study classifies the endemic New World eucheiloid complex of pericaline Lebiini, describing three genera and 16 species, including seven new species and one subspecies, and proposes new synonymies. A phylogeny based on 60 characters indicates Hansus as the sister group to Inna and Eucheila, with evolutionary changes mainly in size, head, reproductive, and surface features, driven by habitat and predator avoidance. The diversification primarily occurred in tropical South America, with five lineages invading Middle America during the Cenozoic.

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Abstract The endemic New World eucheiloid complex is distinguished from the other complexes (thyreopteroid, eurycoleoid, somotrichoid, and pericaloid) of pericaline Lebiini by securiform labial palpomere 3, serrate-setose margins of the pronotum, and flattened stylomere 2 of the ovipositor. The geographical range of eucheiloids extends from northern Argentina to southernmost United States (Brownsville, Texas). All taxa are believed to be arboreal. A key distinguishes among the three genera and 16 species, and each taxon is characterized in terms of structural features, habitat, and geographical distribution. The genera are:Hansus, new genus (generitype—H. reichardti, new species);InnaPutzeys (generitype—Inna punctataPutzeys =Polystichus boyeri(Solier)); andEucheilaDejean (generitype—Euchyla flavilabris(Dejean)). Seven new species and one new subspecies are described:Hansus reichardti(type locality—Guyana, EssequiboR.. Morrabali Ck.);Inna palpalis,I. atrata arbor, andI. inpa(type locality—Brazil, Amazonas, near Manaus);I. purpurea(type locality—Brazil, Amazonas, 60 km n. Manaus);I. splendens(type locality—Venezuela, Aragua, Rancho Grande);Eucheila adisi(type locality—Brazil, Amazonas, 60 km n. Manaus); andE. cordova(type locality—México, Veracruz, Cordova). The following new synonymies of specific names are proposed (valid names listed first):Inna costulataChaudoir, 1872 =I. granulataChaudoir, 1872;Inna boyeri(Solier, 1835) =I. punctataPutzeys, 1863, andI. texanaSchaeffer, 1910. A reconstructed phylogeny of the eucheiloids, based on analysis of 60 character states included in 33 characters, shows thatHansusis sister group of the ancestral stock ofInna+Eucheila. Evolutionary modifications have affected principally body size, head and mouthparts (feeding and associated adaptations), reproductive structures, and features of the body surface thought to be associated with avoidance of predators (concealment and flash coloration). Diversification may also have involved differentiation by habitat (type of forest occupied). The principal theater of evolution has been tropical South America, with incursions both southward and northward. Middle America has been invaded by five lineages, probably during Cenozoic time, from Middle Tertiary to the Quaternary. Of these lineages, three are represented in Middle America by endemic species:E. cordova,I. planipennisBates, andI. nevermanniLiebke. Two lineages are represented by species whose ranges extend northward from South America:I. costulataChaudoir andI. boyeri(Solier).

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Differences in vessel shape and decoration in the earliest ceramic complexes of Middle and South America permit the recognition of two generalized ceramic traditions: an earlier one featuring large rounded jars or tecomates, and a later one emphasizing flange-rim bowls and broad-line incised decoration. The earliest sites of the tecomate tradition are shell middens, suggesting primary diffusion in a nonagricultural context. Sites of the flange-rim tradition are associated with tropical forest vegetation, and its initial spread may be coupled with that of slash-and-burn agriculture. The tecomate tradition diffused between 3000 and 1500 B.C. from a source on the coast of Ecuador. The flange-rim tradition spread rapidly between 1200 and 1000 B.C., probably from a center on the Caribbean coast of Colombia. 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In the case of many early ceramic complexes in the New World, geographical distribution is incompletely known and chronological controls are poor, preventing conclusive demonstration that similarities are the result of common origin rather than independent invention. Examination of the environment in which the cultural complexes occur may shed light on the cor' This article is a revision of Especulaciones sobre rutas tempranas de difusion de la ceramica entre Sur y Mesoamerica (Hombre y Cultura. tomo 1. no. 3. pp. 1-15. Panama. 1964). We are grateful to the editor, Reina Torres de Arauz, for permission to publish an English version. 20 BIOTROPICA 1(1): 20-27. 1969 This content downloaded from 157.55.39.254 on Mon, 05 Sep 2016 06:05:01 UTC All use subject to http://about.jstor.org/terms rect interpretation of such traits. Culture is the primary means by which man adapts to his physical environment, and the main difference between man and other animals is the substitution of this cultural means of adaptation for biological ones (e.g., Mead 1964). Of the major categories of culture-technological, social, and ideological-technology, which includes subsistence techniques, is most intimately related to the environment. All environments are not equally suitable for hunting, for shellfish gathering, or for agriculture. To the extent that environments offer different potentialities for human subsistence exploitation, either in terms of wild foods or in terms of agricultural productivity, they limit the level of complexity attainable by cultures occupying them. O'n the other hand, an environmental setting with high potential for cultural exploitation will not necessarily lead to realization of this potential. 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Usnea is a highly diverse lichen genus, currently including ca. 350 species that are widely distributed from polar to tropical regions. The exceptional plasticity of morphological traits towards environmental parameters causes difficulties in the circumscription of species, and taxonomic revisions are needed, especially in tropical regions. In this taxonomic treatment of Usnea in tropical South America (Bolivia, Brazil, Colombia, Ecuador, incl. the Galapagos Islands, Paraguay, Peru and Venezuela), we identified 51 species, representing ca. half of the total diversity of species occurring in this region. Out of these 51 species, 16 are so far endemic to South America, whereas several species have a wide distribution, sometimes across several continents. Twelve species were newly described and 15 species were newly reported for South America, indicating that the diversity of Neotropical Usnea species is particularly high and that numerous discoveries are yet to be made. In addition to taxonomic studies, the phylogeny of Usnea was investigated using the nuclear ribosomal genes ITS rDNA and nuLSU, as well as two protein-coding genes RPB1 and MCM7. The concatenated dataset revealed that Usnea is subdivided into four highly-supported clades, corresponding to the traditionally circumscribed subgenera Eumitria, Dolichousnea, Neuropogon and Usnea. However, phenotypic characters that have been used to characterize these subgenera are often homoplasious in the phylogeny and a parallel evolution of these characters is suggested. On the other hand, the majority of species described from phenotypic characters are monophyletic in our phylogeny. We conclude that the combinations of phenotypic characters are suitable discriminators for the delimitation of species. On the other hand, most of these characters have evolved several times independently during the diversification of the genus Usnea and are therefore inadequate to describe generic subdivisions.

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In this paper we present a preliminary vertical modal decomposition of FGGE level IIIb data for the tropical sector over South America during the summer period 29 January-16 February. The vertical modes are defined by a linearized σ-coordinate primitive equation model as in Kasahara and Puri. The average flow during the analysed period is separated into 9 blocks covering tropical and subtropical South America and the average energetics is computed as a function of the vertical mode index. It is shown that the external mode is dominant at higher latitudes and, as the equator is approached, an internal mode with equivalent depth of the order of 250 m becomes dominant. The basic characteristics of the tropical upper circulation over South America, such as the Bolivian High and the deep trough off the NE coast of Brazil, are clearly depicted by the 4th internal mode. The flow at higher latitudes in the southern hemisphere is represented by a combination of both external and 4th internal modes with a dominant external mode at higher northern latitudes. The transient behaviour of the energetics over the continental blocks is correlated with the convective activity over the region. DOI: 10.1111/j.1600-0870.1985.tb00280.x

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