A revision of genus Atalophlebioides (Ephemeroptera: Leptophlebiidae)

Abstract

Atalophlebioides Phillips is redescribed as a monotypic genus endemic to New Zealand. All life stages of A. cromwelli (Phillips) are described, and a lectotype is designated. The relationships of the genus and the ecology of A. cromwelli are discussed.     

A New Species of Thraulodes (Ephemeroptera: Leptophlebiidae) from Mexico

A new species of Thraulodes from Mexico is described from male and female imagines. The male imagines of this species have very characteristic genitalia that allow its easy separation from the other known species of the genus.     

Phylogeny and tribal classification of Sterrhinae with emphasis on delimiting Scopulini (Lepidoptera: Geometridae)

Abstract. The phylogenetic relationships of tribes of the geometrid subfamily Sterrhinae (Lepidoptera) were studied, with special emphasis on finding delimiting characters for the tribe Scopulini. Two cladistic analyses were conducted for fifty‐nine species representing all previously recognized Sterrhinae tribes and covering the geographical range of the subfamily. In the first analysis, twelve putative synapomorphies of Scopulini, taken from the literature, were coded for actual specimens in order to test their ability to support the monophyly of the group. The resulting strict consensus cladogram was totally unresolved. In the second analysis, the twelve characters were combined with additional information from the morphology and ecology of adults and immature stages. Analysis of these ninety‐six characters resulted in a well‐resolved cladogram. The tribes were found to be monophyletic, except Cosymbiini and Rhodostrophiini. There are two main lineages within Sterrhinae: Cosymbiini + Rhodometrini + Timandrini and Rhodostrophiini + Cyllopodini + Sterrhini + Scopulini. Aletini and Problepsini lay within the concept of Scopulini. The association of the included Larentiinae taxa with the Cosymbiini + Rhodometrini + Timandrini lineage questions the monophyly of Sterrhinae. A majority of the recovered synapomorphic characters had been recognized previously, but several new phylogenetically informative characters were found, especially from the thorax. No unique characters diagnosing the tribe Scopulini were found, but many homoplastic synapomorphic features were found which diagnose parts of it. All recognized Sterrhinae genera are assigned tentatively to tribes and problematic cases are discussed.     

Three new genera of Leptophlebiidae (Ephemeroptera) from New Zealand

Abstract

Three new genera—Austroclima, Mauiulus, and Cryophlebia—are established for species of Leptophlebiidae from New Zealand. The following new combinations are included: Austroclima sepia (Phillips) and Cryophlebia aucklandensis (Peters). Atalophlebioides sepia is redescribed as Austroclima sepia and a neotype is designated. Two new species, Austroclima jollyae and Mauiulus luma, are described. All life stages are described, and the relationships of each genus are discussed. Keys are given to male and female imagos, subimagos, and nymphs of each species, and to all New Zealand genera with species previously placed in Atalophlebioides.     

New genera and species of Leptophlebiidae (Ephemeroptera) from New Zealand

Abstract

New genera Isothraulus, Arachnocolus, and Penniketellus are established for three species of leptophlebiid mayfly from New Zealand. Each genus is monotypic and endemic to New Zealand. Isothraulus and Arachnocolus are known only from the northern North Island, and Penniketellus is known only from the Arthur's Pass area of the central South Island. The male and female imago, nymph, and egg of Isothraulus abditus n.sp., the male imago, male subimago, and nymph of Arachnocolus phillipsi n.sp., and the male and female imago, female subimago, and egg of Penniketellus insolitus n.sp. are described. The relationships of each genus and the ecology of nymphs of each species are discussed.     

A new species of Ulmeritoides from Brazil (Ephemeroptera: Leptophlebiidae)

The leptophlebiid mayfly genus Ulmeritoides is composed of nine species, three of which have been recorded in Brazil: U. patagianus, U. uruguayensis, and U. misionensis. In the present paper, a new species of this genus, Ulmeritoides oepa sp.nov., is described and illustrated based on fifteen larvae collected in Uraricoera River, state of Roraima, northern Brazil. The species was compared to U. misionensis, U. tifferae, and U. guanacaste, the other species of Ulmeritoides known from the larval stage. U. opea sp.nov. can be distinguished from these species by the following combination of characteristics: 1) medial denticle on anteromedian emargination of labrum much larger than others; 2) anterior tibiae slightly stained black toward the apex; 3) medial femora with few dorsal spines, similar to short thick needles; 4) hind femora with numerous dorsal spines, similar to short thick needles; 5) medial femora with no medial black markings, apical brown markings. The new species seems to be more closely related to U. misionensis, another species in which the anteromedian emargination of the labrum has a median denticle much larger than the others. The studied specimens were deposited in the Invertebrates Collection of the Amazonian Research National Institute, Manaus, state of Amazonas, Brazil, and in the Entomological Collection of the Department of Zoology, Federal University of Rio Janeiro, Rio de Janeiro, state of Rio de Janeiro, Brazil.     

Phylogeny of Ephemeroptera (mayflies) based on molecular evidence

This study represents the first molecular phylogeny for the Order Ephemeroptera. The analyses included 31 of the 37 families, representing approximately 24% of the genera. Fifteen families were supported as being monophyletic, five families were supported as nonmonophyletic, and 11 families were only represented by one species, and monophyly was not testable. The suborders Furcatergalia and Carapacea were supported as monophyletic while Setisura and Pisciforma were not supported as monophyletic. The superfamilies Ephemerelloidea and Caenoidea were supported as monophyletic while Baetoidea, Siphlonuroidea, Ephemeroidea, and Heptagenioidea were not. Baetidae was recovered as sister to the remaining clades. The mayfly gill to wing origin hypothesis was not supported nor refuted by these data. Mandibular tusks were supported as having at least one loss in Behningiidae and, together with the burrowing lifestyle, possibly two origins. The fishlike body form was supported as plesiomorphic for mayflies with multiple secondary losses. Topological sensitivity analysis was used as a tool to examine patterns concerning the stability of relationships across a parameter landscape, providing additional information that may not have been acquired otherwise.     

Morphology and evolution of the wing bullae in South American Leptophlebiidae (Ephemeroptera)

Insects were the first animals to take to the skies, and have been flying for over 320 million years. The order Ephemeroptera is, or at least is part of, the most early-diverging lineage of extant winged insects. The extant species present a very short adult life span, mainly dedicated to reproduction and dispersal of eggs. Mating and egg-laying behavior depend on flight. Wings are structures to fly and as such face a number of physical and physiological challenges. The convex curvature along the anterior–posterior axis of the wing generates a camber that must be carefully regulated. One of the most interesting ways of wing bending is provided by the bullae, which have been defined as short sections of flexible chitin, where the flexion lines cross veins. Although the bullae have been frequently used as taxonomic characters, there is no study focused on their morphology, although their prevalence on the wings of mayflies strongly suggests a role in flight. In order to identify evolutionary trends of these structures within Ephemeroptera, we constructed a matrix with comparative anatomy data of the bullae from whole mounts of the wings of 300 specimens belonging to 70 species of several mayfly families, as well as scanning microscopy samples of selected specimens. We also surveyed the number of bullae and their distribution in the wings of the different species within the South American Leptophlebiidae clade. We optimized the characters onto the latest published phylogeny for Leptophlebiidae.     

Observations on Neotropical Rainpools (Bolivia) with emphasis on Chironomidae (Diptera)

Nine rainpools in the savanna of northern Bolivia were examined using a random, semi‐quantitative sampling method. Both the macrophyte and insect assemblages showed that the longevity of a pool tended to be the dominant influence on the colonizing biota in the Tropics. Twenty‐four families of insects were recorded, including seven Heteroptera, five Odonata and five Diptera. The highly ephemeral (A‐)pools, which were dominated by beetles, differed from longer‐lived waters (B‐ and C‐pools). The composition of the insect assemblages in the B‐ and C‐pools formed a continuum, and the Chironomidae became more numerous and diverse with increasing water body longevity and habitat complexity. Out of the 20 recorded chironomid genera, three uncommon larval types are presented (Chironomini Gen X, "Tanytarsus”; (b), "Tanytarsus”; (dark L.o.). The study area is among the least researched in Bolivia, so many insects, and the chironomids in particular, are suspected as being newly discovered species.     

Three New Species of Thraulodes (Ephemeroptera: Leptophlebiidae: Atalophlebiinae) from Venezuela

Three new species of Thraulodes from Venezuela are described. One species is known from male and female imagines, and the other two only from male imagines. With this, the known Thraulodes species for Venezuela are four. A key for the male imagines of the known species of Thraulodes from Venezuela is provided.     

A remarkable new genus of Atalophlebiinae (Ephemeroptera: Leptophlebiidae) from the Neotropics

An unusual species of Leptophlebiidae is described based on males, females, and nymphs from Rio de Janeiro State, Brazil. As a consequence of its distinct characteristics on all stages, a new genus is established. The new genus can be distinguished from other South American Leptophlebiid genera mostly by: Adults: vein MA of fore wings asymmetrical; hind wings with costal projection well developed, Sc ending at cross vein near costal projection; tarsal claws dissimilar, one blunt other acute; projections of styliger plate forming two well developed lobes with rounded apex, ventrally obstructing view of the penes; penes fused on basal half, each lobe with a ventral furrow and a long and slender spine directed anteriorly. Nymphs: Head prognathous, wider than labrum; labrum with prominent median emargination, with three subtle crenulations; body flattened; hind wings pads present; tarsal claws with over 20 denticles, subapical denticle much larger than remaining denticles; gills long and narrow, present on abdominal segments I-VII; posterolateral projections present on abdominal segments VIII-IX. Phylogenetic analyses conducted based on a previously published data matrix that included other South America leptophlebiid genera placed Poranga nessimiani gen. nov. et sp. nov. as sister to Bessierus + Perissophlebioides. In most analyses this clade was recovered within the Farrodes complex. Nymphs of the new taxa are particularly similar to Bessierus, whereas adults share the very acute costal projection on hind wings seen in non-dipterous members of the Farrodes complex.     

Phylogeny and classification of the Xeromelissinae (Hymenoptera: Apoidea, Colletidae) with special emphasis on the genus Chilicola

Abstract A phylogenetic analysis and classification are provided for the bee subfamily Xeromelissinae based on 248 morphological characters, many of which are novel and illustrated. A total of 47 ingroup species was included in the analysis, representing at least two divergent members from each described genus or subgenus and seven taxa which did not readily fall within previously described subgenera. Three most parsimonious trees were found (length, 1508; consistency index, 40; retention index, 70). The result shows that Xeromelissa renders Chilimelissa paraphyletic, and the 20 known Chilimelissa species are reassigned to Xeromelissa, giving the following new combinations: X. luisa (Toro & Moldenke) (this is the type species of Chilimelissa), X. mucar (Toro & Moldenke), X. xanthorhina (Toro), X. brevimalaris (Toro), X. rosie (Toro and Packer), X. laureli (Toro and Packer), X. chusmiza (Toro), X. longipalpa (Toro), X. pedroi (Toro & Moldenke), X. australis (Toro & Moldenke), X. chillan (Toro & Moldenke), X. farellones (Toro & Moldenke), X. machi (Toro), X. minuta (Toro & Moldenke), X. nortina (Toro & Moldenke), X. sielfeldi (Toro & Moldenke), X. obscura (Toro & Moldenke), X. irwini (Toro & Moldenke), X. nolanai (Toro & Moldenke) and X. rozeni (Toro & Moldenke). Group support was estimated using symmetric resampling and group supported/contradicted (GC) ratios, which compare the frequency of each most parsimoniously resolved clade with the alternative arrangement that was most commonly found in resampling. Relationships among subgenera for Chilicola are weakly supported. By contrast, when three previously synonymized subgenera (Stenoediscelis, Heteroediscelis and Oediscelisca) are resurrected, there is good support for all subgenera (GC ≥ 99), except two: a paraphyletic Oediscelis and a polyphyletic Anoediscelis. Both of these subgenera became monophyletic following successive approximations character weighting. Four distinctive new subgenera are described: Unicarinicola Packer subgen.n. , Obesicola Packer subgen.n. , Capitatiscopa Packer subgen.n. and Toroediscelis Packer subgen.n. , and a revised key to the subgenera of Chilicola is provided.     

Two New Species of Austrophlebioides Campbell and Suter (Ephemeroptera: Leptophlebiidae) from Australia, with Notes on the Genus

Two new species of Austrophlebioides Campbell and Suter, A. marchanti sp.nov. from Victoria, and A. booloumbi sp.nov. from southeastern Queensland, are described, necessitating a slight modification to the generic definition. A. decipiens (Harker) comb.n. is transferred to Austrophlebioides from Deleatidium.     

A new species of Thraulus from Nanjing (eastern China) with single first gill (Insecta: Ephemeroptera: Leptophlebiidae)

Thraulus femoratus sp. nov., a new species found in eastern China, is described from imagos, subimagos, nymphs, and eggs. The nymphs are unique in the genus as well as in the Thraulus group because of their single lanceolate gills 1. The imagos can be differentiated from those of related species by the following combination of characters: slightly robust hindwing with a bluntly rounded costal projection that is situated over half the distance from wing base to apex, darkly pigmented basal portion of hindwing, and subapical dark band on hind femur. Subimagos resemble the imagos in color pattern. Eggs have eight pairs of long threads on the polar portion and slight ridges on the surface. The reduced gills 1 in combination with entirely fringed gills 2-7 represent a new evolutionary lineage and ecological type in the Thraulus group, but provide limited information regarding phylogenetic relationships.     

Systematics, cladistics and biogeography of the American genus Farrodes (Ephemeroptera: Leptophlebiidae: Atalophlebiinae)

Cladistic and biogeographic analyses of the genus Farrodes are presented. Two species groups are delineated within Farrodes: F. caribbianus and F. bimaculatus. Three species formerly placed in other genera– Thraulus caribbianus Traver, Thraulus roundsi Traver and Homothraulus maculatus (Needham & Murphy)–are transferred to Farrodes. The species of the F. caribbianus species group are revised. Three new species are described: F. savagei from Venezuela, F. maya and F. mexicanus from Mexico. Keys to separate the two species groups of Farrodes and the species of the F caribbianus species group are provided. Successive cladistic analyses were carried out on both adult and nymphal characters using Hennig86 and CLADOS. The matrix was composed of all available data (nymphal characters were missing for some species), from nymphal and adult stages separately and on taxa represented by both adult and nymphal characters. Species of the genera Simothraulopsis and Homothraulus (components of the Farrodes lineage) were included in the analyses, and Ecuaphkbia was used as the outgroup. Results of the four analyses are compatible. The historical biogeography of Farrodes , with a distribution from nordiern Argentina to southern Texas, is analysed using the program COMPONENT. Areas of endemism are established, and some of their relationships compared with those of other groups available in the literature.     

Intersexuality in Farrodes Peters, 1971 (Ephemeroptera: Leptophlebiidae: Atalophlebiinae) from Brazil

Gynandromorphism or intersexuality is a rare condition where the individual has male and female features. In this report we described two imagoes of Farrodes xingu Domínguez et al. (1996) from Brazil presenting intersex traits. We considered the arranged sexual characters as antero-posterior and bilateral. One individual has a male head, abdomen full of eggs and the subgenital plate with a malformed outset of penes (antero-posterior), while the other individual has half of the head with a male-like turbinate eye and the other half with a regular female-like one, and with malformed male genitalia (bilateral).     

Phylogeny of Callibaetis (Ephemeroptera: Baetidae) based on morphology of nymphs and adults

Callibaetis is considered to be one of the most problematic genera among mayflies by a series of taxonomic inaccuracies that have accumulated over the last two centuries. Despite these taxonomic problems, two independent hypotheses of species groups have been proposed. In the first hypothesis, three species groups for North America were proposed, and in the second, three species groups were proposed for South American. In these hypotheses, the generic delimitation and monophyly of Callibaetis have not been evaluated under a cladistic framework. Taking this into account, the objectives of this study were to verify the monophyly of Callibaetis and whether the groups of species proposed for the genus are corroborated as natural. The matrix included 128 morphological characters and one habitat character, 119 discrete characters were compiled (101 of nymphs and 18 of adults), and six imaginal characters are related to pigmentation pattern. Continuous characters were ratios and were represented by 10 characters (nine for nymphs and one for adults). The data set was analysed under implied weights. Group support was estimated with relative Bremer support and frequency differences. The results corroborated the monophyletic nature of Callibaetis and the generic status of Callibaetoides; however, the groups proposed for North and South American Callibaetis species were not corroborated. Our study indicated four groups of species for the genus, which we proposed as subgenera: Callibaetis, Abaetetuba subgen. n. Aiso subgen. n. and Cunhaporanga subgen. n.