The paralobe: A new diagnostic and synapomorphic character for the genera Paulipalpina Gnaspini & Peck, 1996 and Parapaulipalpina Gnaspini, 1996 (Leiodidae: Cholevinae: Ptomaphagini).

In insect taxa with homogeneous external morphology, genital structures often emerge as essential traits for interspecific differentiation. In the tribe Ptomaphagini (Coleoptera, Leiodidae, Cholevinae), precise identification often depends on analyzing the male genital morphology, even at the genus level. Here, we present a new character for diagnosing the genera Paulipalpina Gnaspini & Peck, 1996 and Parapaulipalpina Gnaspini, 1996. This feature, which we dub 'paralobe', is a projection arising from the internal surface of the right lobe of the aedeagal apex. Based on its absence in other beetles, including other Ptomaphagini, we recognize it as a putative synapomorphy for those genera. The recognition of this previously overlooked structure adds important information for understanding the sequence of changes that occurred in the male genitalia among the genera of Ptomaphagini.

Updated catalog and generic keys of the Leiodidae (Insecta: Coleoptera) of the Neotropical region ("Latin America": Mexico, the West Indies, and Central and South America).

This paper presents an updated catalog of all taxa of Leiodidae (s.lat.) reported from the Neotropical Region. Keys are presented for the identification of all subfamilies, tribes, and 62 described genera. Three undescribed genera are included in the keys. A total of 600 valid named species are listed, with type localities, type depositories, synonyms, distributions, and biologies where known, and some unnamed species as recorded in the literature. Many species remain to be described. In this work we formally establish no new synonyms and no new combinations although we may indicate the existence of these; but we add new records for described species, and we make spelling corrections of scientific names, when appropriate. A brief review of distribution patterns is given. The fauna has been derived partly from some Nearctic elements that have penetrated as far south as Bolivia. A few genera in the Neotropical element have penetrated the Nearctic Region as far north as the northern U.S.A. or southeastern Canada. Most of the Neotropical genera are autochthonous. In southern South America there is a diverse Neo-Austral fauna with clear "Transantarctic" relationships to Australia and New Zealand and weakly to southern Africa. Some genera variously occur on other continents.

Adelopsis diabolica, a new species of Ptomaphagini (Coleoptera: Leiodidae: Cholevinae) from São Paulo State, Brazil.

Adelopsis diabolica, new species (Coleoptera: Leiodidae: Cholevinae: Ptomaphagini) is described from Boracéia Biological Station, São Paulo State, Brazil. The apex of the male aedeagus is uniquely armed and diagnostic. Females have slightly expanded protarsomeres (less expanded than those of males), a very rare feature in the family.

Redescription of the 'older Adelopsis' species (Coleoptera: Leiodidae: Cholevinae: Ptomaphagini) based on the analysis of type specimens.

As preparation for a revision of the Neotropical genera Adelopsis Portevin, 1907, Paulipalpina Gnaspini and Peck, 1996, and Parapaulipalpina Gnaspini, 1996, we review and redescribe the earlier named species and code characters of their genitalia. These characters are then used to redefine species groups. We review the following 22 "older" species: Adelopsis ascutellaris (Murray, 1856) (male lectotype here designated); Adelopsis aspera Jeannel, 1936; Adelopsis asperoides Szymczakowski, 1963; Adelopsis azzalii Szymczakowski, 1975 (here raised to specific status-previously as Adelopsis brunnea azzalii); Adelopsis bellator Szymczakowski, 1968; Adelopsis benardi (Portevin, 1923); Adelopsis brasiliensis Jeannel, 1936; Adelopsis brevicollis Szymczakowski, 1975 (here raised to specific status-previously as Adelopsis brunnea brevicollis); Adelopsis bruchi (Pic, 1926) (male lectotype here designated); Adelopsis darwini Jeannel, 1936; Adelopsis grouvellei Jeannel, 1936; Adelopsis heterocera Portevin, 1907 (the type species of Adelopsis, here considered a junior synonym of Adelopsis ruficollis (Portevin, 1903)); Adelopsis insolita Szymczakowski, 1961; Adelopsis luculenta Szymczakowski, 1963; Adelopsis orcina Szymczakowski, 1975 (here raised to specific status-previously as Adelopsis brunnea orcina); Adelopsis ovalis Jeannel, 1936; Adelopsis pteromoria Szymczakowski, 1975 (here raised to specific status-previously as Adelopsis brunnea pteromoria); Adelopsis ruficollis (Portevin, 1903) (male lectotype here designated) (here considered a senior synonym of Adelopsis heterocera Portevin, 1907); Adelopsis triangulifer Szymczakowski, 1961; Parapaulipalpina filicornis (Jeannel, 1936); Paulipalpina dispar (Portevin, 1903) (male lectotype here designated); and Paulipalpina simoni (Portevin, 1903). The status of Adelopsis brunnea Jeannel, 1936 is discussed, and is restricted to a single, nominal subspecies. Specific status is restablished for Adelopsis linaresi Szymczakowski, 1969 (previously as Adelopsis brunnea linaresi). The status of Paulipalpina claudicans (Szymczakowski, 1980) is discussed. To correct published misidentifications in museum collections, we also describe the following nine new species: Adelopsis claudina sp. n., Adelopsis mrazi sp. n., Adelopsis szymczakowskii sp. n., Adelopsis waclawi sp. n., and Paulipalpina jeanneli sp. n. (all from Brazil), Paulipalpina consuelo sp. n. from Peru, Adelopsis portevini sp. n. and Paulipalpina aragua sp. n. from Venezuela, and Paulipalpina coatepec sp. n. from Mexico. Iutururuca Gnaspini, 1993, described as a subgenus of Adelopsis, is here considered a junior synonym of Adelopsis Portevin, 1907, which is defined as having no subgenera.

Morphology-based phylogenetic reconstruction of Cholevinae (Coleoptera: Leiodidae): a new view on higher-level relationships.

The current state of knowledge of the suprageneric relationships in Cholevinae is either derived from informal evaluations of putative synapomorphies or based on molecular studies with limited taxonomic sampling. Here we assessed the higher-level relationships in this subfamily based on a phylogenetic analysis of 97 morphological characters scored for 93 terminals, representing all tribes. Both parsimony and Bayesian analyses were used. The monophyletic origin of Cholevinae was corroborated, except for the unexpected inclusion of Leptinus in the implied weighting analysis. Eucatopini + Oritocatopini were retrieved as basal branches in the evolution of Cholevinae. The monophyletic origin of all remaining Cholevinae was confirmed, which is consistent with molecular evidence. Anemadini was non-monophyletic, in accordance with earlier hypotheses. Cholevini was rendered non-monophyletic by the uncertain inclusion of Prionochaeta and the consistent exclusion of Cholevinus. A close affinity of Ptomaphagini to Sciaphyini and Leptodirini was suggested, although the position of Sciaphyes remains uncertain. The phylogenetic hypothesis of Cholevinae provided here is the most comprehensive presently available. The list of characters shows that a substantial part of the data was obtained from the ventral side. This is a strong argument for a detailed pictorial documentation of the ventral body parts in taxonomic descriptions, in contrast to the common practice of only illustrating the dorsal habitus of the beetles.

Pretarsal structures in Leiodidae and Agyrtidae (Coleoptera, Staphylinoidea).

We analysed pretarsal characters of 87 species of Leiodidae (including 10 cholevines and representatives of all tribes and ca. 60% of the genera of non-cholevines), five species of Agyrtidae, and nine representatives of outgroup taxa (Hydraenidae, Staphylinidae, Hydrophilidae, and Histeridae) using scanning electron microscopy. We focused our observations on the architecture of the empodium (including the sclerites and associated setae), the shape and composition of the medial projection of the distal margin of the terminal tarsomere, and the armature of the claws, which were considered a promising source of information for delimiting supraspecific taxa in our previous study. We identified several diagnostic features and recognize potential synapomorphies at the tribal, subtribal and generic levels. The internal systematic arrangement and/or even the monophyletic status of most of the subfamilies of Leiodidae (Camiarinae, Catopocerinae, Leiodinae, and Platypsyllinae) are challenged. We identified potential synapomorphies for Camiarinae (Camiarini and Agyrtodini) and Leiodinae. The non-monophyly of Cholevinae is possible because part of the tribe (Anemadini, Eucatopini, and Oritocatopini) shares potentially apomorphic features with Leiodinae (e.g., a triangular medial projection with a diagonal row of conical spines), whereas another part (Leptodirini and Ptomaphagini) shares a potentially apomorphic feature with Coloninae and Platypsyllinae (a typical medial projection with two distinct triangular projections).

Show me your tenent setae and I tell you who you are - Telling the story of a neglected character complex with phylogenetic signals using Leiodidae (Coleoptera) as a case study.

The tarsal setae in 97 species of Leiodidae and eight outgroups were examined using SEM imaging and dissections. Modified adhesive setae present in males are referred to as "male tenent setae" (MTS). In most cases, dilated tarsomeres were associated with MTS, which were always present on the protarsi and sometimes the mesotarsi. MTS are reported for the first time on the mesotarsi of Leptodirini and on the metatarsi in two genera of Sogdini. Contrary to reports in the literature, the reduction in the number of the MTS bearing mesotarsomeres is considered a derived condition. Both sexes of Leptinus (Platypsyllinae) have modified setae (referred to as tenent setae in the literature), probably related to their specialised association with mammals, and a patch of MTS was recognized for the first time among those modified setae among males. Four main types of MTS are recognised: (1) a plesiomorphic discoidal type that has a shaft with a round cross-section and maintains a similar diameter throughout its length until forming the expanded discoidal terminal plate; (2) a minidiscoidal type, similar to discoidal but with a relatively small terminal plate, found in Cholevinae; (3) a conical type, present in Leiodinae (excluding Estadiini) where the shaft increases in diameter until forming the terminal plate; and (4) a spatulate type, where an even wider terminal plate has a lateral projection, derived from the conical form and synapomorphic for the leiodine tribes Pseudoliodini, Scotocryptini, and possibly Agathidiini.

Two new synonymies at the generic level in Neotropical Ptomaphagini (Coleoptera, Leiodidae, Cholevinae).

We here recognize the genus Excelsiorella (s. str.) Salgado, 2008 as a junior synonym of the genus Adelopsis Portevin, 1907, and Viruana Salgado, 2013, a subgenus of Excelsiorella, as a junior synonym of the genus Parapaulipalpina Gnaspini, 1996. As a consequence, we propose the following new combinations: Adelopsis latissima (Salgado, 2008) n. comb. (from Excelsiorella (s. str.)) and Parapaulipalpina tambopata (Salgado, 2013) n. comb. (from Excelsiorella (Viruana)).

Embryonic development of Ampheres leucopheus and Iporangaia pustulosa (Arachnida: Opiliones: Gonyleptidae).

The first studies concerning the embryonic development of harvestmen started in the late 19th century, and focused mostly on holarctic species, and only three species of the suborder Laniatores (the largest, among the four suborders considered presently) were studied. Moreover, the last studies on embryology of harvestmen were made during the late 1970s. This study focused on the embryonic development of Ampheres leucopheus (Gonyleptidae, Caelopyginae) and Iporangaia pustulosa (Gonyleptidae, Progonyleptoidellinae). The embryonic development was followed in the field, by taking daily photographs of different eggs during about 2 months. When laid, eggs of A. leucopheus and I. pustulosa have approximately 1.13 and 1.30 mm in diameter, respectively, and the second is embedded in a large amount of mucus. The eggs grow, mainly due to water absorption at the beginning of the process, and they reach a diameter of about 1.35 and 1.59 mm, respectively, close to hatching. It took, respectively, 29-56 days and 35-66 days from egg laying to hatching. For the description of the embryonic development, we use photographs from the field, SEM micrographs, and histological analysis. This allowed us, for instance, to document the progression of structures and pigmentation directly from live embryos in the field, and to record microstructures, such as the presence of perforations in the cuticle of the embryo in the place where eyes are developing. Yet, contrary to what was expected in the literature, we record an egg tooth in one of the studied laniatoreans.

Sexually dimorphic tegumental gland openings in Laniatores (Arachnida, Opiliones), with new data on 23 species.

Sexually dimorphic glands often release sexual pheromones both in vertebrates and invertebrates. Species of Laniatores (Arachnida, Opiliones) seem to depend on chemical communication but few studies have addressed this topic. In this study, we review the literature for the Phalangida and present new data for 23 species of Laniatores. In 16 taxa, we found previously undescribed sexually dimorphic glandular openings on the femur, patella, metatarsus, and tarsus of legs I and metatarsus of legs III and IV. For the other species, we provide scanning electron micrographs of previously undescribed sexually dimorphic setae and pegs located on swollen regions of the legs. We also list additional species in which males have swollen regions on the legs, including the tibia, metatarsus, and tarsus of legs I, trochanter and tibia of legs II, femur, metatarsus, and tarsus of legs III, and metatarsus and tarsus of legs IV. The function and biological role of the secretions released by these glands are discussed.

Comparative study of the defensive behavior and morphology of the gland opening area among harvestmen (Arachnida, Opiliones, Gonyleptidae) under a phylogenetic perspective.

The arachnids of the order Opiliones (harvestmen) have a pair of scent glands opening at the sides of the body, the substances of which are used in defense. Several types of behavioral, morphological and chemical defensive mechanisms have been assigned to the order as a whole, although some of these tactics were restricted to particular groups. Only around 25 species have been studied from this perspective so far. In the present paper, we analyzed 33 species (mostly from the largest harvestmen family, the Gonyleptidae) aiming at recognizing the usefulness of the defensive characters in taxonomy and evolutionary biology. We observed the morphology of the gland opening (ozopore) area and the defensive behavior, and their relationship, and mapped these traits on an available phylogenetic hypothesis of relationship within Gonyleptidae. As outgroups, we analyzed Cosmetidae and Stygnidae. Combining the observed behavioral characters of the emission of defensive secretion (near the ozopore, with liquid displacement through an integumentary groove, or in form of a jet) with the morphological types of the gland opening (direction of the integumentary dome that surrounds the gland opening, presence of two openings and the relationship between their sizes, and presence of a V-shaped cut at the anterior opening), we recognized eight patterns. In addition, we could examine the evolution of such traits within Gonyleptidae.