A new species of Askola Peters, 1969 (Ephemeroptera: Leptophlebiidae) from Southeastern Brazil.

Askola fortuita sp. nov. is described based on male imagos from Southeastern Brazil, near Caparaó National Park. The new species is somewhat similar to A. froehlichi Peters, 1969 in regards to development of the male compound eyes and relative length of penis, but can be readily distinguished from this and other species of the genus by the following characteristics: dorsal portion of compound eyes widely meeting on meson of head; forewings hyaline; femora white, with a submedial mark and an apical band, both orangish brown; abdominal coloration mostly translucent white; genitalia with styliger plate washed with brown and red laterally, forceps segment I reddish-brown on basal » , remainder translucent white, penes yellowish brown; penes not extending beyond posterior margin of styliger plate.

Brevitentoria Weaver 1984 (Trichoptera: Integripalpia) of Esprito Santo State, Brazil: New records and new species.

In Brazil, about 900 species of Trichoptera have been recorded, with some species in all Brazilian states. Nevertheless, the collection effort is unequal, with several under-sampled regions. Despite being located entirely within the Atlantic Forest ecoregion, a hotspot of biodiversity, Esprito Santo State has a low known caddisfly richness when compared to nearby areas in the same ecoregion, especially for the infraorder Brevitentoria. This suggests the existence of a Trichoptera biodiversity knowledge gap. Aiming to overcome these taxonomic and distributional shortfalls, we performed a comprehensive inventory of the Brevitentoria species in the state. The sampled sites were distributed from North to South of the state covering 22 locations. In total, 3,420 adult specimens of Brevitentoria were analyzed, leading to a total of 40 species. Two families and 27 species are recorded for the first time from the state. Additionally, we describe three new species of the genera Phylloicus, Helicopsyche (Feropsyche), and Marilia. As a result of this survey, we increase by 100% the number of species of Brevitentoria known from the state, and by 30% for the number of known Trichoptera species. Based on incidence data from this inventory and from the literature, the Brevitentoria species richness was estimated to be about 72 species in Esprito Santo State.

A new species of Campsurus Eaton, 1868 (Ephemeroptera: Polymitarcyidae) from the Doce River Basin, Brazil.

The male imago of Campsurus mirim sp. nov., an integrant of the C. major species group, is described based on material from a lake at the Rio Doce basin, Southeastern Brazil. It can be distinguished from other species of the group by the following combination of characteristics: small to medium size (body 7.3 mm, forewings 6.1 mm); blackish pigments on dorsum of head widely and strongly marked in fresh material; abdominal color pattern with pale transverse dashes on terga IIIVII; posterior margin of sternum IX triangular at middle; pedestals flat with inner margin broadly rounded and with both posterolateral corners (inner and outer) approximately of the same length, both acute; penes relatively wide and large, ventrally curved on apical and with apex of main lobe slightly twisted.

Hagenulopsis Ulmer (Ephemeroptera: Leptophlebiidae): re-description, morphological notes and a new species from South America, Brazil.

Hagenulopsis diptera Ulmer, type species of the genus Hagenulopsis, was originally described based on imagos from Santa Catarina State, Southern Brazil. Misconceptions of H. diptera circumscription led to erroneous attribution of material from Minas Gerais and Esprito Santo, Southeastern Brazil, to H. diptera. Despite the increase in the number of species attributed to Hagenulopsis, little attention has been given to the type species. After comparative examination of photographs of the holotype and fresh material of H. diptera from Southeastern Brazil, we conclude that many specimens previously assigned to H. diptera represent a new species. Thus, we redescribe H. diptera and describe a new species Hagenulopsis perere sp. nov. based on eggs, nymphs and imagos. Diagnostic features of Hagenulopsis perere sp. nov. include cross veins between C and RP1 strongly clouded with brown and outer surface of mid femur with a brown spot at midlength. Finally, comments and new records are presented for Hagenulopsis minuta Spieth.

A new species of Campsurus Eaton, 1868 (Ephemeroptera: Polymitarcyidae) from Brazil.

Campsurus is the most species-rich genus in Polymitarcyidae, and it is divided into four groups. A new species is described and illustrated here based on male imagos. Campsurus molinerii sp. nov. seems to be similar to C. nessimiani, C. dasilvai, and C. cuyuniensis because of the shape of penes and pedestal. The new species differs from them by the main penis lobe being expanded laterally and slightly curved ventrally, with the apex acute and curved ventrally outwards, the apical portion almost cylindrical with apex in ventral view pointed, secondary membranous lobe cylindrical and relatively long, and the outer projection of pedestal with apical condyle process-like projection.

Mayfly taxonomy (Arthropoda: Hexapoda: Ephemeroptera) during the first two decades of the twenty-first century and the concentration of taxonomic publishing.

The twentieth anniversary of the first issue of Zootaxa (De Moraes Freire, 2001) provides an appropriate opportunity to reflect on some trends in global Ephemeroptera taxonomy publishing over the last two decades, with a focus on the description of new species and the outsized role of the journals Zootaxa and ZooKeys, in particular. Detailed reviews of world Ephemeroptera knowledge up to about 2000 were collected in a series of nine papers from a symposium on the subject, published together in the proceedings of the ninth International Conference on Ephemeroptera (Domínguez 2001). Domínguez Dos Santos (2014) provided updates and analysis for South America up to the year 2012. More recent detailed accounts of regional and taxonomic diversity, and other aspects of mayfly biology and ecology, were reviewed by Jacobus et al. (2019), while Ogden et al. (2019) discussed current issues involving higher classification.

Unmasking a gap: A new oligoneuriid fossil (Ephemeroptera: Insecta) from the Crato Formation (upper Aptian), Araripe Basin, NE Brazil, with comments on Colocrus McCafferty.

The Crato Formation (Araripe Basin) preserves one of the most diverse entomofaunas of the Cretaceous. Among the groups of insects, mayflies stand out in abundance, but among them oligoneuriids are especially rare. A newly discovered adult oligoneuriid from this unit is here described as Incogemina nubila gen. et sp. nov. and new subfamily Incogemininae. A phylogenetic analysis recovered the new taxon as the sister group to the species-rich and cosmopolitan Oligoneuriinae. The paratype of Colocrus indivicum, described as an "oligoneuriid" from the same unit, is here reviewed and considered as belonging to the family Hexagenitidae. The biogeographical and taxonomic implications of this discovery and the phylogenetic position of Incogemina nubila are discussed. Incogemina bridge an important morphological gap between the Oligoneuriinae and the extant Chromarcys. Also, it demonstrates that the divergence between Oligoneuriinae and Incogemininae probably occurred in South America.

Description of two related new species of Miroculis Edmunds, 1963 (Ephemeroptera: Leptophlebiidae) from Brazil and Colombia.

Two new species of Miroculis Edmunds, 1963 (Ephemeroptera: Leptophlebiidae) are described, diagnosed and illustrated based on adults and/or nymphs collected in Brazil and Colombia. Both species share an uncommon genitalia morphology, which includes a long penis that is curved at base and a long styliger plate. In the case of the species from Brazil, which is also described based on nymphs, gills are devoid of the characteristic lateral lobes found in the other species of the genus.

Revision of the Afrotropical genus Susua Lugo-Ortiz McCafferty 1998 (Ephemeroptera: Baetidae: Protopatellata).

The species originally described as Cheleocloeon sigiensis Gillies 2001 is moved to the genus Susua Lugo-Ortiz McCafferty 1998. Additional descriptions of Susua sigiensis (Gillies 2001) are given based on male and female imagoes reared from larvae from several localities in Tanzania (including the type locality in the Upper Sigi Basin). Susua niandanensis (Wuillot in Wuillot Gillies 1993) is redescribed based on larvae and male imagoes from Guinea, Mali, Côte d'Ivoire and Zambia. The genus Susua belongs to the plesiomorphon Protopatellata Kluge Novikova 2011 and has an Afrotropical distribution; Susua is closely related to Crassabwa Lugo-Ortiz McCafferty 1996. General characteristics of Susua are given based on the two known species.

Redescription of the Afrotropical genus Crassabwa Lugo-Ortiz & McCafferty 1996 (Ephemeroptera: Baetidae: Protopatellata).

Larvae, subimagoes, imagoes of both sexes and eggs of Crassabwa flava (Crass 1947) are described based on reared material from South Africa, Zambia and Tanzania. Larvae, subimagoes, imagoes of both sexes and eggs of a new species Crassabwa ludmilae sp. n. are described based on reared material from Zambia, Guinea and Mali. Larvae and details of a male subimago extracted from a mature larva of a new species, Crassabwa ameliae sp. n., are described from Zambia, Guinea and Mali. Diagnosis of the genus Crassabwa Lugo-Ortiz & McCafferty 1996 is corrected. Among species formerly attributed to Crassabwa, the species originally described as Centroptilum loweae Kimmins 1949 is moved to Cheleocloeon Wuillot & Gillies 1993 and synonymized with Ch. excisum (Barnard 1932): Cheleocloeon excisum (Barnard 1932) = Cheleocloeon loweae (Kimmins 1949) comb. n., syn. n. The species formerly attributed to Crassabwa and originally described as Centroptilum badium Kopelke 1979 belongs to a new genus, which will be described in a separate paper by N. Kluge. Crassabwa is closely related to Susua Lugo-Ortiz & McCafferty 1998; the species originally described as Cheleocloeon sigiense Gillies 2001 is treated here as Susua sigiense comb. n.; redescription of Susua and argumentation of this new combination will be given in a separate paper.

The imagos of some enigmatic members of the Hermanella complex (Ephemeroptera, Leptophlebiidae).

The imago stages of three species of the Hermanella complex are described mostly based on material from Roraima, northern Brazil: Hydrosmilodon gilliesae, Hydromastodon sallesi and Leentvaaria palpalis. Male imagos of Hydrosmilodon gilliesae and Leentvaaria palpalis both have a pair of large, broad projections at the posterior margin of the styliger plate, nearly covering the penis lobes; in Leentvaaria palpalis, however, these projections are fused. The male imago of Hydromastodon sallesi resembles Hydrosmilodon plagatus in that both species have a styliger plate with a robust projection that is curved towards the penis lobes. DNA barcoding is likely to be a powerful investigative tool for identifying and understanding species limits among these Ephemeroptera taxa, especially if it is used within an integrative taxonomic context. An updated identification key to the genera of the Hermanella complex is proposed.

Two new species of Caenis Stephens, 1835 (Ephemeroptera: Caenidae) from South America.

Caenis Stephens, 1835 is a relatively diverse South American genus of Caenidae (Ephemeroptera), with 22 recorded species, but much of its diversity remains poorly explored in this region. In the present study, two new species of Caenis with apically pointed forceps are described: Caenis amacayacu sp. nov., based on male adults from Colombia; and C. elidioi sp. nov, based on all life stages from several localities in Brazil.

Two new species of Bungona Harker, 1957 (Ephemeroptera: Baetidae) from Borneo, Indonesia.

Two new species of Bungona, belonging to the subgenera Chopralla Waltz & McCafferty, 1987 and Centroptella Braasch & Soldán, 1980, are described based on larvae from Kalimantan (Borneo, Indonesia). Bungona (Centroptella) papilionodes n. sp. is the third species described for the subgenus. It can be distinguished from Bungona (Centroptella) longisetosa (Braasch & Soldán, 1980) and Bungona (Centroptella) soldani (Müller-Liebenau, 1983) by the length of the maxillary palp, the presence or absence of an additional small denticle on the lateral margin of the distal incisor, and the spination of the paraproct. This new report of the subgenus greatly increases its geographic range of distribution, as it was known only from Sri-Lanka and China. Bungona (Chopralla) bintang n. sp. is the seventh species described for the subgenus Chopralla and the second described from Borneo. It differs from others species of the subgenus and especially from Bungona (Chopralla) pusilla (Müller-Liebenau, 1984) (Borneo) by the combination of lacking hindwing pads, the particular spination of distal margins of tergites, and the shape of the maxillary palp. The two new species fit into the recently revised concepts of Chopralla and Centroptella and confirm the characters used to support these taxa as valid subgenera.

FOREWORD: EDITORS' INTRODUCTION.

Since its establishment ZOOTAXA has become not only a rapid journal for zoological systematics but also a respected forum for discussions of all taxonomic matters, and it has gradually attained a distinguished position among other zoological journals by its special issues. These collections of papers treat varied themes such as the Carl Linnaeus legacy (Zhang & Shear 2007, Minelli et al. 2008), cataloguing metazoan life (Zhang 2011, 2013), and promoting and discussing the future of taxonomic sciences, for example modification of the International Code of Zoological Nomenclature (ICZN 2008). For these reasons we offer this special issue to celebrate the fruitful career of the eminent Brazilian researcher Dr. Angelo Barbosa Monteiro Machado ("Professor Angelo" to his friends and colleagues).

Phylogeny and biogeography of Asthenopodinae with a revision of Asthenopus, reinstatement of Asthenopodes, and the description of the new genera Hubbardipes and Priasthenopus (Ephemeroptera, Polymitarcyidae).

The Neotropical species of Asthenopodinae are revised in a formal phylogenetic context. The five known species of Asthenopus Eaton, 1871, together with other five new species were included in a cladistic analysis using morphological characters (continuous and discretes). Representatives of the Afro-Oriental group of the subfamily (Povilla Navás, 1912 and Languidipes Hubbard, 1984) were also included to test the monophyletic hypothesis traditionally accepted for the group. Additional taxa representing the other subfamilies of Polymitarcyidae were incorparated: Ephoron Williamson, 1802 (Polymitarcyinae) and Campsurus Eaton, 1868, Tortopus Needham & Murphy, 1924 and Tortopsis Molineri, 2010 (Campsurinae). A matrix of 17 taxa and 72 characters was analyzed under parsimony resulting in a single tree supporting the monophyly of the subfamily Asthenopodinae. Other results include the monophyly of the Afro-Oriental taxa (Povilla and Languidipes), the paraphyletic nature of Neotropical Asthenopodinae, and the recognition of four South American genera: Asthenopus (including Asthenopuscurtus (Hagen), 1861, Asthenopusangelae de Souza & Molineri, 2012, Asthenopusmagnus sp. n., Asthenopushubbardi sp. n., Asthenopusguarani sp. n.), Asthenopodes Ulmer, 1924, stat. n. (including Asthenopuspicteti Hubbard, 1975, stat. n., Asthenopodestraverae sp. n., Asthenopodeschumuco sp. n.), Priasthenopus gen. n. (including Priasthenopusgilliesi (Domínguez), 1988, comb. n.), and Hubbardipes gen. n. (including Hubbardipescrenulatus (Molineri et al.), 2011, comb. n.). Descriptions, diagnoses, illustrations and keys are presented for all Neotropical taxa of Asthenopodinae (adults of both sexes, eggs and nymphs). Additionally a key to the subfamilies and genera of Polymitarcyidae is included. A quantitative biogeographic analysis of vicariance is presented and discussed through the study of the "taxon history" of the group.

ResumenSe revisan las especies neotropicales de Asthenopodinae en un contexto filogenético. Las cinco especies de Asthenopus Eaton, 1871, junto con otras cinco nuevas especies, son analizadas cladísticamente a partir de caracteres morfológicos externos (continuos y discretos). También se incluyen representantes del grupo Afro-Oriental de la subfamilia (Povilla Navás, 1912 y Languidipes Hubbard, 1984) para comprobar la hipótesis de monofilia tradicionalmente aceptada para el grupo. Se incorporaron taxones adicionales representando las otras subfamilias de Polymitarcyidae: Ephoron Williamson, 1802 (Polymitarcyinae) y Campsurus Eaton, 1868, Tortopus Needham & Murphy, 1924 y Tortopsis Molineri, 2010 (Campsurinae). Se analizó bajo parsimonia una matriz de 17 taxa y 72 caracteres resultando un sólo árbol que apoya la monofilia de Asthenopodinae. Otros resultados incluyen la monofilia del grupo Afro-Oriental (Povilla y Languidipes), la naturaleza parafilética de los Asthenopodinae neotropicales, y el reconocimiento de cuatro géneros sudamericanos: Asthenopus (incluyendo a Asthenopuscurtus (Hagen), 1861, Asthenopusangelae de Souza & Molineri, 2012, Asthenopusmagnussp. n., Asthenopushubbardisp. n., Asthenopusguaranisp. n.), Asthenopodes Ulmer, 1924, stat. n. (Asthenopuspicteti Hubbard, 1975, stat. n., Asthenopodestraveraesp. n., Asthenopodeschumucosp. n.), Priasthenopusgen. n. (Priasthenopusgilliesi (Domínguez), 1988, comb. n.), y Hubbardipesgen. n. (Hubbardipescrenulatus (Molineri et al.), 2011, comb. n.). Se presentan descripciones, diagnosis, ilustraciones y claves para todos los taxones neotropicales de Asthenopodinae (adultos de ambos sexos, huevos y ninfas). Adicionalmente se incluye una clave para subfamilias y géneros de Polymitarcyidae. Se presenta un análisis biogeográfico cuantitativo de vicarianza y se discute la historia particular del grupo.

Redescription of Leptohyphes cornutus Allen, 1967 (Ephemeroptera: Leptohyphidae) and description of three related new species.

Leptohyphes cornutus Allen from Brazil (Santa Catarina) and Misiones, Argentina is re-studied from new material and newly diagnosed as follows: tubercles on head (two pairs), pronotum (two pairs, anterior pair very small), and mesonotum (two pairs), abdominal terga without paired tubercles, nor remnants of them; fore femur length/maximum width, 1.4-1.8; tarsal claws denticulation 4-11+1; hind wing pads present in females; gill formula 3/10/8/8/5. Three new species are described from the nymphal stage: 1) Leptohyphes airuoca sp. nov. from Minas Gerais and Rio de Janeiro (Brazil) characterized by: i) two paired tubercles on head, two pairs on pronotum, and one pair on mesonotum, abdominal terga VI-IX with single medial tubercle projecting from hind margin; ii) fore femur length/maximum width, 1.7-2.0; iii) tarsal claws denticulation 4+0; iv) hind wing pads absent in female; v) gill formula 3/9/9/7/6. 2) Leptohyphes cornutillus sp. nov. from Espirito Santo (Brazil), with: i) paired tubercles on head (two pairs), pronotum (two pairs, anterior pair very small), and mesonotum (three pairs), abdominal terga with remnants of paired tubercles on hind margin (blunt and short undulations in dorsal view); ii) fore femur length/maximum width, 2.1-2.3; iii) tarsal claws denticulation 6+1; iv) hind wing pads present in females; v) gill formula 3/11/9/9/6. And 3) Leptohyphes nebulosus sp. nov. from Espirito Santo (Brazil), with: i) two paired tubercles on head, one pair on pronotum, and one pair on mesonotum, abdominal terga VI-VII with paired submedian tubercles on hind margin; ii) fore femur length/maximum width, 1.6; iii) tarsal claws denticulation 5+1; iv) hind wing pads present in females; v) gill formula 3/9/9/9/6. 

Discovery of an alien species of mayfly in South America (Ephemeroptera).

Despite its wide, almost worldwide distribution, the mayfly genus Cloeon Leach, 1815 (Ephemeroptera: Baetidae) is restricted in the Western hemisphere to North America, where a single species is reported. In the Neotropics, except for some species wrongly attributed to the genus in the past, there are no records of Cloeon. Recently, however, specimens of true Cloeon were collected along the coast of Espírito Santo, Southeastern Brazil. In order to verify the hypothesis that this species was recently introduced to Brazil, our aim was to identify the species based on morphological and molecular characters and to confirm the presence of true representatives of the genus in the Neotropics. Our results revealed that the specimens found in Brazil belong to the Afrotropical species C. smaeleni Lestage, 1924. The identity of the species, its distribution, along with its previous absence in regularly sampled sites, is a clear sign that the specimens of C. smaeleni found in Espírito Santo are introduced, well established, and that the colonization took place very recently.

A new species of Miroculis Edmunds, 1963 (Ephemeroptera: Leptophlebiidae) from northeastern Brazil.

Miroculis stenopterus, sp. nov., is described based on material from State of Pernambuco, Northeastern Brazil. The imago of the new species can be distinguished, among other characteristics, by an uncommon forewing shape, long and narrow, similar to species of genus Microphlebia. The tentatively associated nymph can be distinguished from other species of the genus mainly by the abdominal color pattern and the ratio of labial palp segment 2 to segment 1 (1.01-1.04 x).