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Taxonomy of Saldula: revised genus and species group definitions, and a new species of the pallipes group from Tunisia (Heteroptera: Saldidae)
Abstract
A revised diagnosis of the genus Saldula Van Duzee based upon synapomorphic characters of male genitalia and hemelytral pigmentation patterns is given. According to this concept ca. 55% of the species World-wide currently assigned to Saldula will have to be transferred to other genera. Several new characters are employed in the definition of species and species groups. The species attributable to the pallipes species group are listed and supplemented by a brief overview of the taxonomic relationships of the Old World species. Saldula luteola sp. n. is described from coastal salt marshes on the Island of Djerba, Tunisia. It mainly appears in extremely depigmented morphs unparalleled elsewhere in the Saldidae and likely constitutes a sister-species of S. sardoa Filippi. The conspicuous intraspecific variation in body size and hemelytral pigmentation in S. palustris (Douglas), the closest relative of the former species, is depicted and briefly discussed. The descriptive accounts are accompanied by numerous photographic illustrations of somatic and genital characters.
... Cobben realizó varios trabajos relacionados con morfología (Cobben, 1957(Cobben, , 1961(Cobben, , 1968) y clasificación (Cobben, 1959). Los trabajos más recientes pertenecen a J. T. Polhemus (1966Polhemus ( , 1967aPolhemus ( , b, 1968Polhemus ( , 1969Polhemus ( , 1972Polhemus ( , 1985bPolhemus ( , 1991Polhemus ( , 1993 algunos de ellos en colaboración (Chapman & Polhemus, 1965;Cobben & Polhemus, 1966;Polhemus & Evans, 1969;McKinnon & Polhemus, 1986;Lindskog & Polhemus, 1992; Drake & Hoberlandt (1950, mundial), Schuh et al. (1987, mundial), Uhler (1877), Cobben (1980, islas Hawaii), Polhemus (1988, neártico), Lindskog (1995, paleártico), Cassis & Gross (1995, australiano), Cobben (1987a, b, africano), Polhemus (1981, africano), Polhemus & Polhemus (1991, Madagascar), Drake (1961b, Micronesia), Polhemus (1985a, centroamericano), Froeschner (1981, Froeschner (1999, Panamá) y Protić (2009, Serbia). En Argentina, Berg en su serie "Hemiptera Argentina" (1878-1884) trata a las especies de Heteroptera conocidas para el país. ...
... Parámeros generalmente con processus sensualis muy desarrollado. Esclerito medio del endosoma con proyecciones laminares anteriomedianas (Lindskog & Polhemus, 1992). Este género es el más grande de la subfamilia, incluye aproximadamente 60 especies y ha sido dividido en cinco grupos de especies (Reuter, 1912;Lindskog & Polhemus, 1992): pallipes, saltatoria, opacula, orbiculata y notera. ...
... Esclerito medio del endosoma con proyecciones laminares anteriomedianas (Lindskog & Polhemus, 1992). Este género es el más grande de la subfamilia, incluye aproximadamente 60 especies y ha sido dividido en cinco grupos de especies (Reuter, 1912;Lindskog & Polhemus, 1992): pallipes, saltatoria, opacula, orbiculata y notera. En la Argentina han sido registradas S. coxalis, S. differata y S. penningtoni (Fig. 1). ...
SALDIDAE Resumen Se presenta una sinopsis del conocimiento de la familia Saldidae en la Argentina. Se exponen aspectos mor-fológicos, biológicos y filogenéticos, y se compendian los trabajos más importantes que tratan a la familia, así como catálogos taxonómicos y regionales que la in-cluyen. Se brindan claves para subfamilias y géneros sudamericanos y especies argentinas, además de bre-ves diagnosis de los géneros registrados. Se incluye un listado de las especies de la Argentina. Abstract We present an outline of the knowledge of the family Saldidae in Argentina. We include morphological, biological, and phylogenetic aspects; we also list the major works about the family, including taxonomic and regional catalogs. We present keys to separate sub-families, South American genera, and Argentinean species ; we also provide a short diagnosis of the genera. We include a list of the species from Argentina.
... tic; Neotropical; Oceanian (Fiji, FrenchPolynesia, Guam, New Caledonia, Palau, Samoa, Solomon Islands); Oriental; Palearctic; Panamanian; Saharo-Arabian; Sino-Japanese. Remark: Saldula (sensu lato) is a large, nearly worldwide polyphyletic taxon that has often served as a 'wastebasket genus' to classify species that could not readily fit elsewhere.Lindskog & Polhemus (1992) estimated that nearly half of known Saldidae were contained in Saldula and that nearly half of Saldula species could eventually be transferred to other genera. To this date, almost a third of Saldidae species remain in Saldula (sensu lato), most of which remaining to be considered within the monophyletic concept of Saldula (sensu strict ...
... Remark: Saldula (sensu lato) is a large, nearly worldwide polyphyletic taxon that has often served as a 'wastebasket genus' to classify species that could not readily fit elsewhere.Lindskog & Polhemus (1992) estimated that nearly half of known Saldidae were contained in Saldula and that nearly half of Saldula species could eventually be transferred to other genera. To this date, almost a third of Saldidae species remain in Saldula (sensu lato), most of which remaining to be considered within the monophyletic concept of Saldula (sensu stricto) put forward byLindskog & Polhemus (1992). Afrotropical; Palearctic; Saharo-Arabian; Sino-Japanese. ...
A supplement to the Saldidae section of the “Catalog and bibliography of the Leptopodomorpha (Heteroptera)” published by Schuh, Galil and Polhemus (1987: Bull. Amer. Mus. Nat. Hist. 323), is provided. A total of 30 genera and 307 species-group taxa (298 species, 9 subspecies) belonging to 3 tribes and 2 subfamilies of extant Saldidae, is recorded for the world. Changes to the 1987 catalogue are documented. The synonymy and type locality of taxa described between 1987 and 2018 are given. Under each genus, species, and subspecies, geographic distribution is broadly categorised following the terrestrial zoogeographic realms of Holt et al. (2013: Science 339). A summary of the geographic distribution of Saldidae genera is provided as well as faunistic lists of species-group taxa by zoogeographic realms.
... The proposition that the New Zealand saldids described to date are not congeneric with Saldula, is not new (Polhemus 1985a-b;Lindskog & Polhemus 1992;Schuh & Polhemus 2009) but little could be done until now to revise this classification for lack of sufficient and well-documented material. Extensive fieldwork by the authors since 1992 has yielded large amount of new saldid material, including long series of specimens from several populations and a wealth of new information on geographic distribution and biology. ...
... The genus Saldula (sensu lato) is a large, nearly worldwide, polyphyletic taxon that has often served as a 'wastebasket genus' to classify species that could not readily fit elsewhere. Lindskog & Polhemus (1992) estimated that nearly half of known Saldidae were contained in Saldula and that nearly half of Saldula species could eventually be transferred to other genera. These authors were the first to suggest a more restricted taxonomic definition and description of Saldula (sensu stricto), using a monophyletic concept based on two synapomorphies (median cell of corium with eye spot; median endosomal sclerite of male with anteromedial laminar projections). ...
Zemacrosaldula, new genus, is described with Salda australis White, 1876, as type species, resulting in the following new combination Zemacrosaldula australis (White, 1876). Three new species are described: Zemacrosaldula kapekape new species, Z. whakarunga new species, Z. pangare new species. A revision of the taxonomy of all taxa is presented. Species are keyed. Morphological descriptions are provided together with illustrations emphasising the most significant diagnostic features of external morphology and male genitalia. Information is given on synonymy, type specimens, material examined, geographic distribution and biology
... Similarly to Zemacrosaldula species, the taxa under study do not fit Lindskog & Polhemus's (1992) monophyletic concept for Saldula (sensu stricto) or the taxonomic definition of other Saldidae genera. ...
Aoteasalda new genus, is described with Saldula maculipennis Cobben, 1961, as type species, resulting in the following new combination Aoteasalda maculipennis (Cobben, 1961). Kiwisaldula new genus, is described with Saldula parvula Cobben, 1961, as type species. The following new combinations are made: Kiwisaldula parvula (Cobben, 1961); Kiwisaldula butleri (White, 1878); Kiwisaldula laelaps (White, 1878); Kiwisaldula stoneri (Drake & Hoberlandt, 1950). Two species are described as new: Kiwisaldula manawatawhi new species, Kiwisaldula porangahau new species. A lectotype is designated for Salda laelaps White, 1878. The holotype of Salda butleri White, 1878, and the type series of Saldula trivialis Cobben, 1961, and Saldula maculipennis Cobben, 1961, are documented. A new synonymy is established in the genus Zemacrosaldula (valid name listed after equal sign): Saldula trivialis Cobben, 1961 = Zemacrosaldula australis (White, 1876). A revision of the taxonomy of Aoteasalda and Kiwisaldula species occurring on New Zealand’s North Island and nearby offshore islands, is presented. Morphological descriptions are provided, with illustrations emphasising the most significant diagnostic features of the external morphology and male genitalia. Information is given on synonymy, type data, material examined, geographic distribution, and biology. Species of Kiwisaldula and genera of Saldidae recognised for New Zealand, are keyed.
... Recently, Zhang et al. ( 2014 ) found the same relationships among the extant leptopodomorphan families, placing the fossil Palaeoleptidae as sister group to Omaniidae + Leptopodidae, and the fossil Archegocimicidae more closely related to Aepophilidae. Other phylogenetic contributions to the Leptopodomorpha are the cladistic revisions of Saldula Van Duzee and of Pseudosaldula Cobben (Lindskog and Polhemus 1992 ;Schuh and Polhemus 2009 ). ...
Abstract A review of Heteroptera classification, biogeography, and phylogeography is presented. The use of molecular data significantly expanded the knowledge of phylogenetic relationships among and within heteropteran infraorders. However, taxa historically less studied continue to receive little attention. Promising, new molecular approaches with increased genetic markers and broader taxon sampling, as well as new morphological approaches (e.g., microtomography), are the future for more stable classifications and a better comprehension of the heteropteran evolutionary history, but their application is still incipient. A non-exhaustive overview of studies about Neotropical heteropteran biogeography is made and discussed, including those about intercontinental connections and regional distribution patterns.
The most comprehensible studies, and more promising area, seem to be the one
focused on distribution patterns, especially employing macroecological methods,
and trying to elucidate what are the major factors responsible for the distribution of
the group in the Neotropics. Finally, we present an overview of phylogeographic
studies involving Neotropical Heteroptera. It is clear that the best biogeographic
and phyllogeographic studied groups are those with medical and economical importance
(e.g., Reduviidae and Pentatomidae).
The genus Pseudosaldula Cobben, which is restricted to the Andean Subregion of South America, is revised. Fourteen valid species are recognized, nine of them being described as new and 10 previously published names are treated as junior synonyms based on the examination of approximately 3500 specimens from Colombia, Ecuador, Peru, Bolivia, Argentina, and Chile. All taxa are described or redescribed. A key to the species is provided. Color habitus illustrations, distributional maps, and detailed measurements are provided for all species. Scanning electron micrographs of the vestiture, parameres, parandria, face, and pretarsus are provided for representative species, as are color views of the face and the nymphs. The concept of a postclypeus in the Saldidae is questioned and the term transverse swelling, as coined by Parsons, is applied in discussing distinctive aspects of facial morphology in Pseudosaldula. A previously unreported, presumably glandular, pore is documented on the parameres in the Saldinae in the form of a cavernous pit with internal digitiform processes. A phylogenetic analysis based on morphological character data documents the monophyly of Pseudosaldula. Characters treated as synapomorphic for Pseudosaldula are five cells in the membrane of the forewing, the incomplete connection of the transverse swelling across the posterior margin of the clypeus, and the straight connection across the posterior margin of the parandria; nymphal coloration is also distinctive, although treated as ambiguous because this character was not scored for all species. DNA sequence data from the 16S rDNA region of the mitochondrion and H3 nuclear region were acquired for 13 Pseudosaldula spp. and five outgroup taxa. The combined analysis of morphological and sequence data consistently treated Pseudosaldula as paraphyletic. These results are interpreted as the result of inadequate sampling of both taxa and gene regions, in light of the fact that the patterns of distribution become transpacific, as opposed to a monophyletic group in the Andean Region. Not unexpectedly, several morphological characters documenting the monophyly of Pseudosaldula show greater homoplasy in the combined analysis than when analyzing morphological data alone. Therefore, the results of the morphological cladistic analysis are further used to examine distributional patterns in the group. Five areas of endemism are recognized: northern Andes, northern Peru, Puna, central Chile, and subantarctic; the boundaries of these areas show substantial correspondence with those proposed for other groups of insects.
Heteroptera, or true bugs, are part of the most successful radiation of nonholometabolous insects. Twenty-five years after the first review on the influence of cladistics on systematic research in Heteroptera, we summarize progress, problems, and future directions in the field. The few hypotheses on infraordinal relationships conflict on crucial points. Understanding relationships within Gerromorpha, Nepomorpha, Leptopodomorpha, Cimicomorpha, and Pentatomomorpha is improving, but progress within Enicocephalomorpha and Dipsocoromorpha is lagging behind. Nonetheless, the classifications of several superfamily-level taxa within the Pentatomomorpha, such as Aradoidea, Coreoidea, and Pyrrhocoroidea, are still unaffected by cladistic studies. Progress in comparative morphology is slow and drastically impedes our understanding of the evolution of major clades. Molecular systematics has dramatically contributed to accelerating the generation and testing of hypotheses. Given the fascinating natural history of true bugs and their status as model organisms for evolutionary studies, integration of cladistic analyses in a broader biogeographic and evolutionary context deserves increased attention.
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