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Due to the fragility of the ophiuroid (brittle star) skeleton, the bulk of the group’s fossil record consists of dissociated ossicles preserved as microfossils. In spite of their great potential as basis for taxonomic and phylogenetic studies, however, ophiuroid ossicles from the Paleozoic have received very little attention so far. Here, we provide an exhaustive taxonomic assessment of such fossils retrieved from sieving residues from the Silurian of Gotland, Sweden. This material was used in a previous study to describe two key taxa that allowed constraining the origin of the extant ophiuroid clade. The remaining taxa belonging to that same lineage are described in the present paper. The evidence at hand suggests that the stem of the extant ophiuroid clade was formed by two genera, Ophiopetagno and Ophiolofsson gen. nov., including six and five species, respectively, and spanning at least the upper Llandovery through upper Ludlow. We conclude that Ophiopetagno and Ophiolofsson represent sister genera that coexisted through most of the Silurian in the shallow tropical seas of Gotland. They underwent repeated body size reductions in correlation with environmental perturbations, with Ophiopetagno paicei eventually giving rise to Muldaster haakei; the first member of the living Ophiuroidea. Herein, we also introduce two new clades, Ankhurida clade nov. and Ophiovalida clade nov., and the following eight new species: Ophiolofsson joelmciveri gen. et sp. nov., O. obituary gen. et sp. nov., O. immolation gen. et sp. nov., O. archspire gen. et sp. nov., O. hendersonorum gen. et sp. nov., Ophiopetagno bonzo sp. nov., O. kansas sp. nov., O. doro sp. nov.; and two probably new species in open nomenclature: Ophiopetagno sp. 1, and Ophiopetagno sp. 2.
Holocarpic oomycetes infecting freshwater diatoms are obligate endobiotic parasites reported from a wide range of habitats. So far, the taxonomy of and phylogeny of most species remains unresolved, since most have not been reported throughout the past decades and sequence data are available for only the four species, Aphanomycopsis bacillariacearum, Diatomophthora gillii, Ectrogella bacillariacearum, and the recently-discovered species Miracula moenusica. In the current study, a new freshwater diatom parasite resembling Ectrogella bacillariacearum in the sense of Scherffel was discovered from pennate diatoms (Ulnaria acus, Ulnaria ulna) collected from the small stream Einbúalækur on Víkurskarð, North Iceland and investigated for its life cycle and phylogenetic placement. In contrast to the original description, Scherffel reports an achlya-like spore discharge for Ectrogella bacillariacearum. The phylogenetic reconstruction and morphological characterisation in this study revealed that Scherffel’s E. bacillariacearum is largely unrelated to the epitype of the species and is a member of the early-diverging genus Miracula. Consequently, the new species is described as M. einbuarlaekurica in the present study. This adds a second freshwater member to the genus, demonstrating the high ecological adaptability of the genus, which thrives in both freshwater and marine ecosystems.
Exploration of macrofungi in the Jammu and Kashmir regions, India, led to the discovery of two novel species, described herein as Lactarius indohirtipes sp. nov. and L. sharmai sp. nov. This paper presents detailed morphological descriptions and illustrations, as well as a phylogenetic analysis based on nrITS sequences. A comparative analysis with related species is also included.
A new species of Paranthrene Hübner (Lepidoptera: Sesiidae) from the northern midwest United States
(2024)
A clearwing moth species, Paranthrene sogaardi Taft and Smith, 2024, new species, is described from Michigan and Minnesota. The recognition of this new species is based on a phylogeny estimated from mitochondrial cytochrome oxidase I and wingless DNA sequences of 25 specimens representing all Paranthrene Hübner species including individuals from various locations. Paranthrene sogaardi new species was monophyletic and differed from Paranthrene tabaniformis Rottenburg, 1775 by a mean of 7.6% COI pairwise “p” distance, coloration, and genitalic morphology.
ZooBank registration. urn:lsid:zoobank.org:pub:8B9ED6CE-4DEB-4CB2-9235-3CA593F4D9DB
Taxonomic analysis of the genital plates and associated structures in Ophiuroidea (Echinodermata)
(2024)
Recently, new insights have been gained from the ophiuroid skeleton that were instrumental in the inference of a new phylogeny. The so far least studied ossicles are the adradial and abradial genital plates and the radial shields, which articulate with each other and support the genital slit and disc. In addition, the inner sides of the oral shields and madreporites have never been examined in detail. The present study utilized SEM, micro-CT and digital photography to document and examine these structures in 57 species from 28 of the currently accepted 34 families of Ophiuroidea. Early ontogeny and fossils were also considered. Previously, mainly the articular structures had been analysed, but the overall shape of the genital plates was here found to hold important phylogenetic signals. A long-neglected ossicle was re-discovered and studied in detail for the first time, here named the oral genital plate. It was recognized in all Ophintegrida, but was found to be absent in all Euryophiurida. The oral genital plate articulates with the oral shield and supports the proximal part of the genital slit wall. Abradial and oral genital plates were found to be absent in species that lack genital slits, but the adradial genital plate was always present. Numerous new morphological characters with potential phylogenetic signals were identified, described and figured in detail. A pre-existing character matrix was extended and revised with these new data, as well as with recently revised data on oral papillae, and a Bayesian phylogenetic analysis was performed. This phylogeny largely agrees with the current molecular hypothesis, but some branches were not supported.
During exploratory surveys of the fungal diversity in Margalla Hills National Park, Islamabad, we collected a new species of the genus Hymenopellis R.H.Petersen. This is the second report of any species of this genus from Pakistan. Hymenopellis areolata F.Razzaq & Khalid sp. nov. is characterized by an areolate pileus, small basidiospores, and transitional pileipellis (hymeniderm and epithelium) with small pileocystidia. Molecular phylogenetic analyses of the nucleotide sequences of nrITS and nrLSU regions, and morphological data support the description of this new species. A comparison with other closely related species confirmed that the newly described species is distinct from others.
Genomic analysis of Pyrginae Burmeister, 1878 (Lepidoptera: Hesperiidae Latreille, 1809) with an emphasis on the tribes Achlyodini Burmeister, 1878 and Carcharodini Verity, 1940 reveals many inconsistencies between the resulting phylogeny and the current classification. These problems are corrected by proposing new taxa, changing the ranks of others, or synonymizing them, and transferring species between genera. As a result, five subtribes, one genus, 20 subgenera, and one species are proposed as new: Cyclosemiina Grishin, new subtribe (type genus Cyclosemia Mabille, 1878), Ilianina Grishin, new subtribe (type genus Iliana E. Bell, 1937), Nisoniadina Grishin, new subtribe (type genus Nisoniades Hübner, [1819]), Burcina Grishin, new subtribe (type genus Burca E. Bell and W. Comstock, 1948), and Pholisorina Grishin, new subtribe (type genus Pholisora Scudder, 1872), all in Carcharodini; Lirra Grishin, new genus (type species Leucochitonea limaea Hewitson, 1868) in Pythonidina Grishin, 2019; Trifa Grishin, new subgenus (type species Tagiades jacobus Plötz, 1884), Tuberna Grishin, new subgenus (type species Pythonides contubernalis Mabille, 1883), Ebona Grishin, new subgenus (type species Quadrus eboneus E. Bell, 1947), Noctis Grishin, new subgenus (type species Achlyodes accedens Mabille, 1895), and Cyrna Grishin, new subgenus (type species Achlyodes cyrna Mabille, 1895) of Quadrus Lindsey, 1925; Liddia Grishin, new subgenus (type species Helias pallida R. Felder, 1869), Minna Grishin, new subgenus (type species Achlyodes minna Evans, 1953), and Thilla Grishin, new subgenus (type species Eurypterus later Mabille, 1891) of Eantis Boisduval, 1836; Torgus Grishin, new subgenus (type species Ouleus gorgus E. Bell, 1937) of Iliana E. Bell, 1937; Fenops Grishin, new subgenus (type species Cabares enops Godman and Salvin, 1894) of Polyctor Evans, 1953; Bezus Grishin, new subgenus (type species Pellicia bessus Möschler, 1877) and Macarius Grishin, new subgenus (type species Pellicia macarius Herrich-Schäffer, 1870) of Nisoniades Hübner, [1819]; Quadralis Grishin, new subgenus (type species Pterygospidea extensa Mabille, 1891) of Gorgopas Godman and Salvin, 1894; Menuda Grishin, new subgenus (type species Nisoniades menuda Weeks, 1902) and Narycus Grishin, new subgenus (type species Pythonides narycus Mabille, 1889) of Perus Grishin, 2019; Bovaria Grishin, new subgenus (type species Achlyodes cyclops Mabille, 1876), Sebia Grishin, new subgenus (type species Nisoniades eusebius Plötz, 1884), and Stolla Grishin, new subgenus (type species Pholisora balsa E. Bell, 1937) of Bolla Mabille, 1903; Vulga Grishin, new subgenus (type species Achlyodes vulgata Möschler, 1879) and Capilla Grishin, new subgenus (type species Helias aurocapilla Staudinger, 1876, currently a junior subjective synonym of Hesperia musculus Burmeister, 1875) of Staphylus Godman and Salvin, 1896; and Quadrus (Zera) vivax Grishin, new species (type locality in Brazil: Rio de Janeiro). The following 10 are subgenera, not genera or synonyms: Ouleus Lindsey, 1925 and Zera Evans, 1953 of Quadrus Lindsey, 1925; Atarnes Godman and Salvin, 1897 and Eburuncus Grishin, 2012 of Milanion Godman and Salvin, 1895; Pachyneuria Mabille, 1888 and Austinus O. Mielke and Casagrande, 2016 of Sophista Plötz, 1879; Hemipteris Mabille, 1889 and Mictris Evans, 1955 of Pellicia Herrich-Schäffer, 1870; and Hesperopsis Dyar, 1905 and Scantilla Godman and Salvin, 1896 of Staphylus Godman and Salvin, 1896. The following 7 are species, not subspecies: Quadrus (Ebona) cristatus (Steinhauser, 1989) (not Quadrus (Ebona) negrus (Nicolay, 1980)), Quadrus (Quadrus) ophia (A. Butler, 1870) (not Quadrus (Quadrus) lugubris (R. Felder, 1869)), Quadrus (Zera) gellius (Mabille, 1903) and Quadrus (Zera) servius (Plötz, 1884) (not Quadrus (Zera) hyacinthinus (Mabille, 1877)), Mimia pazana Evans,1953 (not Mimia phidyle (Godman and Salvin, 1894)), Polyctor (Polyctor) dagua Evans, 1953 (not Polyctor (Polyctor) polyctor (Prittwitz, 1868)), and Staphylus (Vulga) satrap Evans, 1953 (not Staphylus (Vulga) saxos Evans, 1953); and these 8 are species, not synonyms: Quadrus (Zera) menedemus (Godman and Salvin, 1894) (not Quadrus (Zera) tetrastigma (Sepp, [1847])), Pellicia (Pellicia) bilinea Mabille, 1889 (not Pellicia (Pellicia) dimidiata Herrich-Schäffer, 1870), Pellicia (Hemipteris) nema Williams and Bell, 1939 (not Pellicia (Pellicia) theon Plötz, 1882), Bolla (Bovaria) sodalis Schaus, 1913 (not Bolla (Bolla) imbras (Godman and Salvin, 1896)), Bolla (Bovaria) aplica (E. Bell, 1937) (not Bolla (Sebia) eusebius (Plötz, 1884)), Bolla (Sebia) chilpancingo (E. Bell, 1937) (not Bolla (Bolla) subapicatus (Schaus, 1902)), and Bolla (Stolla) madrea (R. Williams and E. Bell, 1940) and Bolla (Stolla) hazelae (Hayward, 1940) (not Bolla (Stolla) zorilla (Plötz, 1886)). The following 2 are junior subjective synonyms: Achlyodes erisichthon Plötz, 1884 of Quadrus (Zera) servius (Plötz, 1884) (not a subspecies of Quadrus (Zera) tetrastigma (Sepp, [1847]) and Staphylus subapicatus Schaus, 1902 of Bolla (Bolla) imbras (Godman and Salvin, 1896). Furthermore, we propose the following additional new genus-species combination: Gindanes homer (Evans, 1953), Gindanes nides (O. Mielke and Casagrande, 2002), Gindanes maraca (O. Mielke and Casagrande, 1992), Gindanes jenmorrisae (Shuey and Ramírez. 2022), Gindanes tullia (Evans, 1953), Gindanes herennius (Geyer, [1838]), Gindanes proxenus (Godman and Salvin, 1895), Gindanes parallelus (Mabille, 1898), Gindanes braga (Evans, 1953), Gindanes hampa (Evans, 1953), Gindanes rosa (Steinhauser, 1989), Gindanes neivai (Hayward, 1940), Gindanes mundo (H. Freeman, 1979), Gindanes eminus (E. Bell, 1934), Quadrus (Trifa) francesius Freeman, 1969, Quadrus (Trifa) ineptus (Draudt, 1922), Quadrus (Trifa) jacobus (Plötz, 1884), Quadrus (Tuberna) lancea (Hewitson, 1868), Quadrus (Ebona) pescada (E. Bell, 1956), Lirra pteras (Godman and Salvin, 1895), and Lirra limaea (Hewitson, 1868) (not Pythonides Hübner, 1819); Quadrus (Cyrna) zora (Evans, 1953) (not Bolla Mabille, 1903); Eantis later (Mabille, 1891) and Eantis haber (Mabille, 1891) (not Aethilla Hewitson, 1868); Iliana (Torgus) gorgus (E. Bell, 1937) and Iliana (Torgus) taurus (Evans, 1953) (not Eantis Boisduval, 1836); Bolla (Stolla) evemerus (Godman and Salvin, 1896), Bolla (Stolla) chlora (Evans, 1953), Bolla (Stolla) astra (R. Williams and E. Bell, 1940), Bolla (Stolla) balsa (E. Bell, 1937), Bolla (Stolla) tridentis (Steinhauser, 1989), Bolla (Stolla) esmeraldus (L. Miller, 1966), Bolla (Stolla) chlorocephala (Latreille, [1824]), and Bolla (Stolla) incanus (E. Bell, 1932) (not Staphylus Godman and Salvin, 1896). Finally, lectotypes are designated for Achlyodes servius Plötz, 1884 (type locality in Brazil: Rio de Janeiro), Pellicia theon Plötz, 1882 (type locality in South America), and Nisoniades eusebius Plötz, 1884 (type locality in Central America). Unless stated otherwise, all subgenera, species, subspecies, and synonyms of mentioned genera and species are transferred with their parent taxa, and others remain as previously classified.
ZooBank registration. http://zoobank.org/B9AFA1A9-8664-4F31-B4D9-ACF7780C7CC6
Few species of Japygidae (Diplura) have been discovered in cave ecosystems despite their importance as large predators. A small collection of rare specimens of this hexapod group has allowed to explore the taxonomy of japygids from caves in New Zealand, Morocco and South Africa, and to describe one new genus: Imazighenjapyx Sendra & Sánchez-García gen. nov., as well as four new species: Austrjapyx wynbergensis Sendra & Sánchez-García sp. nov., Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov., Opisthjapyx naledi Sendra & Sánchez-García sp. nov. and Teljapyx aotearoa Sendra & Sánchez-García sp. nov. For each of the new taxa we give a comprehensive description of their habitats. These new findings resulted in a revision of the distribution and allowed to re-evaluate the morphological traits of the fifteen cave-adapted japygids species already known worldwide. The functional morphology of the remarkable abdominal pincers of Japygidae and their adaptation to predation are discussed, as well as their potential role in mating behaviour.
The present work reviews the deep-water cone fauna of New Caledonia and its Economic Exclusive Zone. It is based on the material collected for more than 40 years by oceanographic expeditions in the deep waters surrounding New Caledonia, organized by the Muséum national d’histoire naturelle-Paris/ORSTOM, then Institut de Recherche pour le Développement, in the framework of the Tropical Deep-Sea Benthos programme. A total of 2377 lots containing 5113 specimens collected in depths between 100 and 1260 m have been examined. About 770 specimens were collected live, and allowed for radular and DNA studies. A phylogenetic analysis, based on the cox1 gene, of the deep-water cone snail fauna of New Caledonia is presented, along with a detailed, fully illustrated taxonomic account with data on geographic and bathymetric distribution and radular morphology. A total of 76 different species of cone snails were identified among the collected material. Of these, 22 corresponded to typical shallow water species, which were most likely translocated into deeper water, whereas 54 could be considered true components of the deep water (below 100 m) cone snail fauna. Species of the genus Profundiconus represent 22%, whereas those of the genera Conasprella and Conus represent 28% and 50%, respectively. Eleven deep water cone species can be considered as endemic to the New Caledonia EEZ, representing 20.3% of the total. The most abundant species found (more than 400 specimens each) were Conus (Afonsoconus) bruuni, Conasprella (Boucheticonus) alisi, Conasprella (Conasprella) boucheti, and Profundiconus vaubani. The new species Conus (Taranteconus) samadiae sp. nov. is hereby described.
Bumblebees (Bombus Latreille, 1802), because of their large body size, bright colours and activity at times and places that coincide with biologists, are an example of a group of insects that is particularly well represented in museum collections. This is important if taxonomic revisions are to achieve greater comparability among species. Bumblebees have also attracted particular attention because they are especially ecologically and economically valuable for pollination in north temperate regions, where they are now becoming increasingly threatened. I argue that the what, the where, and the how of effective conservation management may be informed by understanding the divergent characteristics that have affected their biogeographical past: by helping us to see ‘the woods’, not just ‘the trees’, of their habitat needs. Identifying suitable habitat should be part of reconstructing historical biogeography within taxonomic revisions. For bumblebees, for example, biogeographical analysis associates major taxonomic groups either with flower-rich lowland grasslands or with flower-rich montane grasslands, highlighting their contrasting requirements for: nest sites, flowers of different depths, pollen-plant families, and especially the differing importance of early spring and late summer flowers for breeding success. This broad view of species groups helps filter the less important idiosyncrasies from local case studies in order to focus conservation actions.
The New World species of the genus Ceratoculicoides Wirth & Ratanaworabhan are described, illustrated and keyed in both sexes. Ceratoculicoides borkenti sp. nov., C. confusus sp. nov., C. grogani sp. nov., C. pacificus sp. nov. and C. propinquus sp. nov. are described, with C. confusus being the first record of the genus from South America (Colombia). Ceratoculicoides blantoni is a junior synonym of C. virginianus and the identity of C. longipennis remains unclear as the male morphospecies associated in taxonomic literature with the female type series is not conspecific. A morphological phylogeny of all extant species in the genus is presented and characters discussed. The moravicus species group is recognized for a clade of species with the lateral margins of the aedeagus straight or concave.
Bujurquina is the most widely distributed and species-rich genus of cichlids in the western Amazon of South America. In this study we describe a new species from Peru from a hypothesized reverse flowing river system. Prior to the origin of the modern Amazon River at 4.5 Ma, this river system had its headwaters on the Iquitos arch, one of several main structural arches (swells) in the Amazon. Prior to the origin of the modern Amazon these arches formed topographic barriers of drainage basins in lowland Amazonia. For our analyses we use morphological and molecular data, analyzed through multivariate statistics and molecular phylogenies, respectivelly. For all valid species in the genus (except B. cordemadi and B. pardus) we additionally for the first time provide photographs of live specimens. Based on DNA phylogeny and coloration patterns we demonstrate that Bujurquina is divided into two main clades and based on this we provide a dichotomous key for all the species.
The freshwater snail genus Mercuria is widely distributed in lowland waters across Western Europe, Northern Africa and the Mediterranean islands. Approximately two-thirds of the currently recognised species are described based on their shell morphology, which may vary within species due to biotic and abiotic factors. Recent molecular phylogenies that included numerous previously documented populations recovered 14 species clades, nine of which correspond to nominal species and five, to undescribed taxa. Here, we formally describe the five undescribed taxa as new species and provide morphological descriptions of the shell and other anatomical structures for three of the other inferred clades and for the species M. maceana to elucidate their taxonomic status and assess the utility of morphological characters for species delimitation in Mercuria. Taken together, the morphological and molecular evidence suggest new identifications and synonymies, having implications on the known geographic range of the studied species, including the type species M. similis. Anatomical measurements and geometric morphometric analysis of shell shape revealed no clear differentiation among the species analysed, predicting the importance of molecular data in elucidating the species diversity of the genus.
A new species, Clitopilus cretoalbus A.Izhar, Zaman, M.Asif, H.Bashir, Niazi & Khalid sp. nov., is described herein based on several collections from Punjab, Pakistan. It is characterized by a clitocyboid to somewhat omphaloid stature combined with a white pileus, decurrent lamellae, an almost central to slightly eccentric whitish relatively long stipe, the occurrence of cheilocystidia, and basidiospores with 6 to 9 ridges in polar view. Molecular phylogenetic analyses of nrITS and nrLSU performed using the maximum likelihood method supported the novelty of this Pakistani species and its placement within the genus Clitopilus section Scyphoides. A comparison with other morpho-anatomically close species confirmed that the newly described species is distinct from others.
The Lasioglossum (Dialictus) gemmatum species complex, also known as the L. tegulare species group and the L. parvum species complex, is a very common, widespread, diverse, and recognisable lineage of sweat bees, containing 22 previously described species and several known undescribed species. The species were recently revised for the eastern Nearctic region and the Greater Antilles, but remain poorly known in the western Nearctic along with most other L. (Dialictus). These characteristics make it a prime candidate for revision in ongoing taxonomic work on the western Nearctic L. (Dialictus). Here we present the results of this revision, including 10 new species descriptions, one new synonymy, a preliminary phylogeny, and keys to known Nearctic species. Species of the eastern Nearctic and a few primarily Neotropical species which can occur in the Nearctic are also included. We report that the L. (D.) gemmatum species complex is likely a monophyletic group arising from the L. (D.) comulum group, but that the enlarged tegula has arisen independently in at least two other L. (Dialictus) lineages, and it contains multiple cases of allopatric speciation. The following species are described as new: Lasioglossum (Dialictus) angelicum sp. nov., L. (D.) deludens sp. nov., L. (D.) diabolicum sp. nov., L. (D.) eremum sp. nov., L. (D.) gloriosum sp. nov., L. (D.) indagator sp. nov., L. (D.) holzenthali sp. nov., L. (D.) magnitegula sp. nov., L. (D.) profundum sp. nov., and L. (D.) rufodeludens sp. nov. Previously undescribed males of L. (D.) perparvum (Ellis, 1914) and L. (D.) pseudotegulare (Cockerell, 1896) and the female of L. (D.) gaudiale (Sandhouse, 1924) are diagnosed and figured for the first time. Lasioglossum (Dialictus) hunteri (Crawford, 1932) is a new subjective junior synonym of L. (D.) ellisiae (Sandhouse, 1924). Pre-2022 specimen records of L. (D.) hunteri and L. (D.) tegulariforme (Crawford, 1907) are attributable to a heterogeneous mix of species, and records of L. (D.) perparvum are likely attributable to L. (D.) deludens.
Two novel species of Russula (Russulaceae, Russulales), namely Russula indosenecis A.Ghosh, D.Chakr., K.Das & Buyck sp. nov. and R. pseudosenecis A.Ghosh, D.Chakr., K.Das & Buyck sp. nov. belonging to sect. Ingratae subg. Heterophyllidiae are proposed herein based on their morphological features and nrITS-based phylogenetic inferences. Both species belong to the Asian ʻR. punctipes-senecisʼ complex of sect. Ingratae. The acrid R. indosenecis was collected from subalpine forests associated with Abies densa, whereas the mild R. pseudosenecis associates with tropical forests dominated by Shorea robusta. Both species are distinct from the other species of this species complex in nrITS sequence data and from all other known species in subg. Heterophyllidiae in the strong amyloidity of their suprahilar spot.
Two new opilionid species from suborder Cyphophthalmi, family Sironidae, Siro franzi Karaman & Raspotnig sp. nov. and Siro ozimeci Karaman sp. nov., from Austria and Croatia respectively, are described and illustrated. Both species show a close relation to two other relict sironid species from the southern and eastern parts of the Alps, Siro valleorum and Siro crassus. All four species are treated here as a monophyletic, alpine group of genus Siro, opposed to the remaining two European sironids, S. rubens and S. carpaticus (palaeoeuropean Siro group). The history of the alpine Siro group parallels the history of a part of the dynamic European archipelago in the Mediterranean Tethys area, which became a part of the Alpine orogeny. Diversification of the alpine Siro group is the result of the orogenic evolution of the Alps, linked to the Austroalpine and South Alpine tectonic units.
During mycological surveys of different areas of Punjab, Pakistan, we collected a new species from the genus Candolleomyces. This is the first report of any species of this genus from Pakistan. Candolleomyces asiaticus M.Asif, A.Izhar, Niazi & Khalid sp. nov. is characterized by ellipsoid to oblong-ellipsoid (8.2 × 5 µm) basidiospores with indistinct germ pores, lageniform to utriform cheilocystidia without adhering deposits at the apices, absence of pleuro-, pileo- and caulocystidia, pileipellis with spherical, subglobose to columnar cells, and presence of clamp connections. The morpho-anatomical and molecular phylogenetic analysis of nrITS revealed its distinct phylogenetic position, further supporting the recognition of a new species.
The present study examines whether the chaetotaxy of the costal vein in the calyptrate families Fanniidae and Muscidae deserves more attention in phylogenetic and taxonomic contexts. An overview of the macrotrichia and their arrangement on wing vein C is given. Special attention is given to the presence/absence of ventral and dorsal setulae on the costal sectors CS1‒3. This is described as one variable character (A) with nine states (A0‒A8). Specimens of both sexes (when possible) of each species belonging to 4 of a total of 5 fanniid genera and 115 of a total of 179 muscid genus-group taxa were examined and scored for character A. It was found that the presumed ancestral state of character A differs between the two families. It is further shown that the main transformational trend in character A in Muscidae has been bi-directional, leading either to the loss of ventral setulae or the gain of dorsal setulae. The utility of character A in the Fanniidae and Muscidae is many-sided and involves taxa ranging from species to family. It is concluded that character A and other aspects of costal chaetotaxy deserve more attention in morphology-based studies of calyptrate flies.
Two new species of subgenus Heterophyllidiae subsection Cyanoxanthinae, Russula fusiformata Y.Song sp. nov. and R. purpureorosea Y.Song sp. nov., collected from the Dinghushan Biosphere Reserve (DHSBR), are described based on both morphology and a phylogenetic analysis of the internal transcribed spacer (ITS), further increasing Heterophyllidiae species diversity in the area. Differences between the two new species and related taxa are analyzed. The other 17 reported species of Russula subgenus Heterophyllidiae that have been collected from DHSBR during mushroom explorations since 2014 are also summarized. The dominant species and the ecological distribution of all 19 species are briefly discussed, and most species are presented in macrofungal plates.
Acrostilicus Hubbard, 1896 and Pachystilicus Casey, 1905 are North American genera known from only one and two species, respectively, and have never been a subject of a modern revision. In fact, Acrostilicus was not even properly described as its author provided only a sketchy diagnosis of the genus and species. Here, we provide a redescription of the genus Acrostilicus and its species and illustrate the habitus and male genital features. For the first time, we also redescribe Pachystilicus and its two species, and provide their differential diagnoses. Additionally, we tested the phylogenetic position of both genera. They were scored into a morphological matrix supplemented with molecular data and the analyses were run using Bayesian inference and maximum likelihood methods. A total of 119 morphological characters and 4859 bp of nuclear (28S, TP, Wg, CADA, CADC, ArgK) and mitochondrial (COI) sequences were analysed for 46 taxa. The results confirmed that both Acrostilicus and Pachystilicus are members of the subtribe Stilicina, but at the same time challenged the monophyly of the subtribe in its current composition. Additionally, we provided further evidence for non-monophyly of the subtribe Medonina and discussed the biology of Acrostilicus and Pachystilicus.
Our expanded efforts in genomic sequencing to cover additional skipper butterfly (Lepidoptera: Hesperiidae) species and populations, including primary type specimens, call for taxonomic changes to restore monophyly and correct misidentifications by moving taxa between genera and proposing new names. Reconciliation between phenotypic characters and genomic trees suggests three new tribes, two new subtribes, 23 new genera, 17 new subgenera and 10 new species that are proposed here: Psolosini Grishin, new tribe (type genus Psolos Staudinger, 1889), Ismini Grishin, new tribe (type genus Isma Distant, 1886), Eetionini Grishin, new tribe (type genus Eetion de Nicéville, 1895), Orphina Grishin, new subtribe (type genus Orphe Godman, 1901), Carystoidina Grishin, new subtribe (type genus Carystoides Godman, 1901), Fulvatis Grishin, new genus (type species Telegonus fulvius Plötz, 1882), Adina Grishin, new genus (type species Nascus adrastor Mabille and Boullet, 1912), Ornilius Grishin, new genus (type species Ornilius rotundus Grishin, new species), Tolius Grishin, new genus (type species Antigonus tolimus Plötz, 1884), Lennia Grishin, new genus (type species Leona lena Evans, 1937), Trida Grishin, new genus (type species Cyclopides barberae Trimen, 1873), Noxys Grishin, new genus (type species Oxynthes viricuculla Hayward, 1951), Gracilata Grishin, new genus (type species Enosis quadrinotata Mabille, 1889), Hermio Grishin, new genus (type species Falga ? hermione Schaus, 1913), Eutus Grishin, new genus (type species Cobalus rastaca Schaus, 1902), Gufa Grishin, new genus (type species Phlebodes gulala Schaus, 1902), Godmia Grishin, new genus (type species Euroto chlorocephala Godman, 1900), Rhomba Grishin, new genus (type species Eutychide gertschi Bell, 1937), Rectava Grishin, new genus (type species Megistias ignarus Bell, 1932), Contrastia Grishin, new genus (type species Hesperia distigma Plötz, 1882), Mit Grishin, new genus (type species Mnasitheus badius Bell, 1930), Picova Grishin, new genus (type species Vorates steinbachi Bell, 1930), Lattus Grishin, new genus (type species Eutocus arabupuana Bell, 1932), Gubrus Grishin, new genus (type species Vehilius lugubris Lindsey, 1925), Koria Grishin, new genus (type species Hesperia kora Hewitson, 1877), Corta Grishin, new genus (type species Eutychide lycortas Godman, 1900), Calvetta Grishin, new genus (type species Hesperia calvina Hewitson, 1866), Oz Grishin, new genus (type species Astictopterus ozias Hewitson, 1878), Praxa Grishin, new subgenus (type species Nascus prax Evans, 1952), Bron Grishin, new subgenus (type species Papilio broteas Cramer, 1780), Turis Grishin, new subgenus (type species Pyrgus 1955, and Synale Mabille, 1904 of Carystus Hübner, [1819]. The following 20 genera are treated as junior subjective synonyms: Leucochitonea Wallengren, 1857 of Abantis Hopffer, 1855; Sapaea Plötz, 1879 and Netrobalane Mabille, 1903 of Caprona Wallengren, 1857; Parasovia Devyatkin, 1996 of Sebastonyma Watson, 1893; Pemara Eliot, 1978 of Oerane Elwes and Edwards, 1897; Ankola Evans, 1937 of Pardaleodes Butler, 1870; Arotis Mabille, 1904 of Mnaseas Godman, 1901; Chalcone Evans, 1955, Hansa Evans, 1955, and Propertius Evans, 1955 of Metrocles Godman, 1900; Jongiana O. Mielke and Casagrande, 2002 of Cobaloides Hayward, 1939; Pamba Evans, 1955 of Psoralis Mabille, 1904; Brownus Grishin, 2019 of Styriodes Schaus, 1913; Mnasilus Godman, 1900 of Papias Godman, 1900; Sucova Evans, 1955 of Mnasitheus Godman, 1900; Pyrrhocalles Mabille, 1904 and Asbolis Mabille, 1904 of Choranthus Scudder, 1872; Miltomiges Mabille, 1903 of Methionopsis Godman, 1901; Sacrator Evans, 1955 of Thracides Hübner, [1819]; and Lychnuchoides Godman, 1901 of Perichares Scudder, 1872. Arunena Swinhoe, 1919 is a junior subjective synonym of Stimula de Nicéville, 1898 (not of Koruthaialos Watson, 1893). The following 27 names are species-level taxa (some in new combinations) reinstated from synonymy: Salantoia gildo (Mabille, 1888) (not Salatis cebrenus (Cramer, 1777)), Bungalotis corentinus (Plötz, 1882) (not Bungalotis midas (Cramer, 1775)), Telegonus cretellus (Herrich-Schäffer, 1869) (not Telegonus cassander (Fabricius, 1793)), Santa palica (Mabille, 1888) (not Chiothion asychis (Stoll, 1780)), Camptopleura cincta Mabille and Boullet, 1917 (not Camptopleura auxo (Möschler, 1879)), Camptopleura orsus (Mabille, 1889) (not Nisoniades mimas (Cramer, 1775)), Metron voranus (Mabille, 1891) and Metron fasciata (Möschler, 1877) (not Metron zimra (Hewitson, 1877)), Limochores catahorma (Dyar, 1916) (not Limochores pupillus (Plötz, 1882)), Pares viridiceps (Mabille, 1889) (not Thoon modius (Mabille, 1889)), Tigasis wellingi (Freeman, 1969) (not Tigasis arita (Schaus, 1902)), Rectava sobrinus (Schaus, 1902) (not Papias phainis Godman, 1900), Nastra subsordida (Mabille, 1891) (not Adlerodea asema (Mabille, 1891), previously in Eutychide Godman, 1900), Lerema pattenii Scudder, 1872 (not Lerema accius (J. E. Smith, 1797)), Lerema (Morys) ancus (Möschler, 1879) (not Cymaenes tripunctus theogenis (Capronnier, 1874)), Cobalopsis zetus (Bell, 1942) (not Cobalopsis nero (Herrich-Schäffer, 1869)), Lerema (Geia) etelka (Schaus, 1902) (not Lerema (Geia) geisa (Möschler, 1879), previously in Morys Godman, 1900), Cymaenes isus (Godman, 1900) (not Cymaenes trebius (Mabille, 1891)), Vehilius labdacus (Godman, 1900) (not Vehilius inca (Scudder, 1872)), Papias amyrna (Mabille, 1891) (not Papias allubita (Butler, 1877), previously in Mnasilus Godman, 1900), Papias integra (Mabille, 1891) (not Papias subcostulata (Herrich-Schäffer, 1870)), Metiscus atheas Godman, 1900 (not Hesperia achelous Plötz, 1882), Dion agassus (Mabille, 1891) (not Dion uza (Hewitson, 1877), previously in Enosis Mabille, 1889), Picova incompta (Hayward, 1942) (not Lerema (Morys) micythus (Godman, 1900), previously in Morys Godman, 1900), Lucida melitaea (Draudt, 1923) (not Lucida lucia (Capronnier, 1874)), Methionopsis modestus Godman, 1901 (not Methionopsis ina (Plötz, 1882)), and Thargella (Volus) volasus (Godman, 1901) (not Eutocus facilis (Plötz, 1884)). The following 57 taxa are elevated from subspecies to species, new status (some in new combinations): Dyscophellus doriscus (Hewitson, 1867) (not Dyscophellus porcius (C. Felder and R. Felder, 1862), Phocides vida (A. Butler, 1872) (not Phocides urania (Westwood, 1852)), Tagiades (Daimio) ceylonica Evans, 1932 (not Tagiades litigiosa Möschler, 1878), Tagiades (Daimio) tubulus Fruhstorfer, 1910 (not Tagiades sambavana Elwes and Edwards, 1897), Tagiades (Daimio) kina Evans, 1934, Tagiades (Daimio) sheba Evans, 1934, Tagiades (Daimio) martinus Plötz, 1884, Tagiades (Daimio) sem Mabille, 1883, and Tagiades (Daimio) neira Plötz, 1885 (not Tagiades trebellius (Hopffer, 1874)), Tagiades (Daimio) korela Mabille, 1891 and Tagiades (Daimio) presbyter Butler, 1882 (not Tagiades nestus (C. Felder, 1860)), Tagiades obscurus Mabille, 1876, Tagiades ravi (Moore, [1866]), Tagiades atticus (Fabricius, 1793), Tagiades titus Plötz, 1884, Tagiades janetta Butler, 1870, Tagiades inconspicua Rothschild, 1915, and Tagiades hovia Swinhoe, 1904 (not Tagiades japetus (Stoll, [1781])), Tagiades silvia Evans, 1934 and Tagiades elegans Mabille, 1877 (not Tagiades gana (Moore, [1866])), Tapena bornea Evans, 1941 and Tapena minuscula Elwes and Edwards, 1897 (not Tapena thwaitesi Moore, [1881]), Darpa dealbata (Distant, 1886) (not Darpa pteria (Hewitson, 1868)), Perus manx (Evans, 1953) (not Perus minor (Schaus, 1902)), Canesia pallida (Röber, 1925) (not Carrhenes canescens (R. Felder, 1869)), Carrhenes conia Evans, 1953 (not Carrhenes fuscescens (Mabille, 1891)), Anisochoria extincta Hayward, 1933 and Anisochoria polysticta Mabille, 1876 (not Anisochoria pedaliodina (Butler, 1870)), Anisochoria verda Evans, 1953 (not Anisochoria minorella Mabille, 1898), Bralus alco (Evans, 1953) (not Bralus albida (Mabille, 1888)), Ephyriades jamaicensis (Möschler, 1879) (not Ephyriades brunnea (Herrich-Schäffer, 1865)), Koruthaialos (Stimula) frena Evans, 1949 (not Koruthaialos focula (Plötz, 1882)), Euphyes kiowah (Reakirt, 1866) (not Euphyes vestris (Boisduval, 1852)), Mnaseas inca Bell, 1930 (not Mnaseas bicolor (Mabille, 1889)), Metron hypochlora (Draudt, 1923) (not Metrocles schrottkyi (Giacomelli, 1911), previously in Metron Godman, 1900), Decinea huasteca (H. Freeman, 1969), Decinea denta Evans, 1955, and Decinea antus (Mabille, 1895) (not Decinea decinea (Hewitson, 1876)), Xeniades pteras Godman, 1900 (not Xeniades chalestra (Hewitson, 1866)), Xeniades difficilis Draudt, 1923 (not Xeniades orchamus (Cramer, 1777)), Xeniades hermoda (Hewitson, 1870) (not Tisias quadrata (HerrichSchäffer, 1869)), Hermio vina (Evans, 1955) (not Hermio hermione (Schaus, 1913), previously in Lento Evans, 1955), Cymaenes loxa Evans, 1955, (not Cymaenes laureolus (Schaus, 1913)), Niconiades peri (Evans, 1955) (not Rhinthon bajula (Schaus, 1902), previously in Neoxeniades Hayward, 1938), Gallio danius (Bell, 1941) (not Vehilius seriatus (Mabille, 1891)), Gallio massarus (E. Bell, 1940) (not Gallio garima (Schaus, 1902) previously in Tigasis Godman, 1900), Cymaenes edata (Plötz, 1882), Cymaenes miqua (Dyar, 1913) and Cymaenes aequatoria (Hayward, 1940) (not Cymaenes odilia (Burmeister, 1878)), Lychnuchus (Enosis) demon (Evans, 1955) (not Lychnuchus (Enosis) immaculata (Hewitson, 1868), previously in Enosis Mabille, 1889), Naevolus naevus Evans, 1955 (not Naevolus orius (Mabille, 1883)), Lucida scopas (Mabille, 1891), Lucida oebasus (Godman, 1900), and Lucida leopardus (Weeks, 1901) (not Lucida lucia (Capronnier, 1874)), Corticea schwarzi (E. Bell, 1941) and Corticea sylva (Hayward, 1942) (not Corticea mendica (Mabille, 1898)), and Choranthus orientis (Skinner, 1920) (not Choranthus antiqua (Herrich-Schäffer, 1863), previously in Pyrrhocalles Mabille, 1904). Borbo impar bipunctata (Elwes and J. Edwards, 1897) is a valid subspecies, not a synonym of Borbo impar tetragraphus (Mabille, 1891), here placed in synonymy with Lotongus calathus (Hewitson, 1876), new synonym. We confirm the species status of Telegonus cassius (Evans, 1952) and Lerema (Morys) valda Evans, 1955. Euphyes chamuli Freeman, 1969 is placed as a subspecies of Euphyes kiowah (Reakirt, 1866), new status. The following 41 taxa are junior subjective synonyms, either newly proposed or transferred from synonymy with other species or subspecies: Telegonus mutius Plötz, 1882 of Euriphellus phraxanor (Hewitson, 1876), Telegonus erythras Mabille, 1888 of Dyscophellus damias (Plötz, 1882), Aethilla jaira Butler, 1870 of Telegonus cretellus (Herrich-Schäffer, 1869), Paches era Evans, 1953 of Santa palica (Mabille, 1888), Antigonus alburnea Plötz, 1884 of Tolius tolimus robigus (Plötz, 1884) (not of Echelatus sempiternus simplicior (Möschler, 1877)), Echelatus depenicillus Strand, 1921 of E. sempiternus simplicior (not of T. tolimus robigus), Antigonus aura Plötz, 1884 of Theagenes dichrous (Mabille, 1878) (not of Helias phalaenoides palpalis (Latreille, [1824])), Achlyodes impressus Mabille, 1889 of Camptopleura orsus (Mabille, 1889), Augiades tania Schaus, 1902 of Metron voranus (Mabille, 1891), Pamphila verdanta Weeks, 1906 of Metron fasciata (Möschler, 1877), Niconiades viridis vista Evans, 1955 of Niconiades derisor (Mabille, 1891), Pamphila binaria Mabille, 1891 of Conga chydaea (A. Butler, 1877) (not of Cynea cynea (Hewitson, 1876)), Psoralis concolor Nicolay, 1980 of Ralis immaculatus (Hayward, 1940), Hesperia dido Plötz, 1882 of Cynea (Quinta) cannae (Herrich-Schäffer, 1869) (not of Lerema lochius (Plötz, 1882)), Proteides osembo Möschler, 1883 of Cynea (Cynea) diluta (Herrich-Schäffer, 1869) (not of Cynea (Quinta) cannae (Herrich-Schäffer, 1869)), Cobalopsis brema E. Bell, 1959 of Eutus rastaca (Schaus, 1902), Psoralis panamensis Anderson and Nakamura, 2019 of Rhomba gertschi (Bell, 1937), Cobalus asella Herrich-Schäffer, 1869 of Amblyscirtes alternata (Grote and Robinson, 1867) (not of Amblyscirtes vialis (W. H. Edwards, 1862)), Papias trimacula Nicolay, 1973 of Nastra subsordida (Mabille, 1891), Pamphila bipunctata Mabille, 1889 and Sarega staurus Mabille, 1904 of Lerema pattenii Scudder, 1872 (not of Cymaenes lumina (Herrich-Schäffer, 1869), previously in Lerema Scudder, 1872), Hesperia aethra Plötz, 1886 of Lerema lineosa (Herrich-Schäffer, 1865) (not of Lerema (Morys) compta Butler, 1877), Megistias miaba Schaus, 1902 of Cobalopsis valerius (Möschler, 1879), Phanis sylvia Kaye, 1914 of Lerema etelka (Schaus, 1902) (not of Lerema (Geia) geisa (Möschler, 1879), previously in Morys Godman, 1900), Carystus odilia Burmeister, 1878, Pamphila trebius Mabille, 1891 and Megistias corescene Schaus, 1902 of Cymaenes lumina (Herrich-Schäffer, 1869), Hesperia phocylides Plötz, 1882 of Cymaenes edata (Plötz, 1882) (not of Lerema accius (J. E. Smith, 1797)), Pamphila xenos Mabille, 1898 of Vehilius inca (Scudder, 1872), Mnasilus guianae Lindsey, 1925 of Papias amyrna (Mabille, 1891), Pamphila nubila Mabille, 1891 of Papias integra (Mabille, 1891) (not of Cynea corisana (Plötz, 1882)), Enosis matheri H. Freeman, 1969 of Metiscus atheas Godman, 1900 (previously in Enosis Mabille, 1889), Hesperia infuscata Plötz, 1882 of Mnaseas derasa derasa (Herrich-Schäffer, 1870) (previously Arotis Mabille, 1904), (not of Papias subcostulata (Herrich-Schäffer, 1870)), Pamphila astur Mabille, 1891 of Metiscus angularis (Möschler, 1877) (not of Cymaenes tripunctus theogenis (Capronnier, 1874)), Anthoptus macalpinei H. Freeman, 1969 of Anthoptus inculta (Dyar, 1918), Methionopsis typhon Godman, 1901 of Methionopsis ina (Plötz, 1882), Methionopsis dolor Evans, 1955 of Thargella volasus (Godman, 1901), Hesperia cinica Plötz, 1882 of Dubiella dubius (Stoll, 1781), Cobalus disjuncta Herrich-Schäffer, 1869 of Dubiella dubius (Stoll, 1781) (not of Vettius lafrenaye (Latreille, [1824])), and Saliana vixen Evans, 1955 of Neoxeniades parna (Evans, 1955). The following are new and revised genusspecies combinations: Euriphellus cebrenus (Cramer, 1777) (not Salatis Evans, 1952), Gorgopas extensa (Mabille, 1891) (not Polyctor Evans, 1953), Clytius shola (Evans, 1953) (not Staphylus Godman and Salvin, 1896), Perus narycus (Mabille, 1889) (not Ouleus Lindsey, 1925), Perus parvus (Steinhauser and Austin, 1993) (not Staphylus Godman and Salvin, 1896), Pholisora litus (Dyar, 1912) (not Bolla Mabille, 1903), Carrhenes decens (A. Butler, 1874) (not Antigonus Hübner, [1819]), Santa palica (Mabille, 1888) (not Chiothion Grishin, 2019), Bralus nadia (Nicolay, 1980) (not Anisochoria Mabille, 1876), Acerbas sarala (de Nicéville, 1889) (not Lotongus Distant, 1886), Caenides sophia (Evans, 1937) (not Hypoleucis Mabille, 1891), Hypoleucis dacena (Hewitson, 1876) (not Caenides Holland, 1896), Dotta tura (Evans, 1951) (not Astictopterus C. Felder and R. Felder, 1860), Nervia wallengrenii (Trimen, 1883) (not Kedestes Watson, 1893), Testia mammaea (Hewitson, 1876) (not Decinea Evans, 1955), Oxynthes trinka (Evans, 1955) (not Orthos Evans, 1955), Metrocles argentea (Weeks, 1901) (not Paratrytone Godman, 1900), Metrocles scitula (Hayward, 1951) (not Mucia Godman, 1900), Metrocles schrottkyi (Giacomelli, 1911) (not Metron Godman, 1900), Niconiades derisor (Mabille, 1891) (not Decinea Evans, 1955), Paratrytone samenta (Dyar, 1914) (not Ochlodes Scudder, 1872), Oligoria (Cobaloides) locutia (Hewitson, 1876) (not Quinta Evans, 1955), Psoralis (Saniba) laska (Evans, 1955) (not Vidius Evans, 1955), Psoralis (Saniba) arva (Evans, 1955) and Psoralis (Saniba) umbrata (Erschoff, 1876) (not Vettius Godman, 1901), Psoralis (Saniba) calcarea (Schaus, 1902) and Psoralis (Saniba) visendus (E. Bell, 1942) (not Molo Godman, 1900), Alychna gota (Evans, 1955) (not Psoralis Mabille, 1904), Adlerodea asema (Mabille, 1891) and Adlerodea subpunctata (Hayward, 1940) (not Eutychide Godman, 1900), Ralis immaculatus (Hayward, 1940) (not Mucia Godman, 1900), Rhinthon braesia (Hewitson, 1867) and Rhinthon bajula (Schaus, 1902) (not Neoxeniades Hayward, 1938), Cymaenes lochius Plötz, 1882 (not Lerema Scudder, 1872), Paracarystus ranka (Evans, 1955) (not Thoon Godman, 1900), Tricrista aethus (Hayward, 1951), Tricrista canta (Evans, 1955), Tricrista slopa (Evans, 1955), Tricrista circellata (Plötz, 1882), and Tricrista taxes (Godman, 1900) (not Thoon Godman, 1900), Gallio madius (E. Bell, 1941) and Gallio seriatus (Mabille, 1891) (not Vehilius Godman, 1900), Gallio garima (Schaus, 1902) (not Tigasis Godman, 1900), Tigasis corope (HerrichSchäffer, 1869) (not Cynea Evans, 1955), Tigasis perloides (Plötz, 1882) (not Cymaenes Scudder, 1872), Amblyscirtes (Flor) florus (Godman, 1900) (not Repens Evans, 1955), Vidius fraus (Godman, 1900) (not Cymaenes Scudder, 1872), Nastra celeus (Mabille, 1891) (not Vehilius Godman, 1900), Nastra nappa (Evans, 1955) (not Vidius Evans, 1955), Vehilius warreni (Weeks, 1901) and Vehilius limae (Lindsey, 1925) (not Cymaenes Scudder, 1872), Cymaenes lumina (Herrich-Schäffer, 1869) (not Lerema Scudder, 1872), Cobalopsis valerius (Möschler, 1879) (not Cobalopsis Godman, 1900), Cobalopsis dictys (Godman, 1900) (not Papias Godman, 1900), Lerema (Morys) venias (Bell, 1942) (not Cobalopsis Godman, 1900), Papias latonia (Schaus, 1913) (not Cobalopsis Godman, 1900), Dion iccius (Evans, 1955) and Dion uza (Hewitson, 1877) (not Enosis Mabille, 1889), Vistigma (Vistigma) opus (Steinhauser, 2008) (not Thoon Godman, 1900), Saturnus fartuga (Schaus, 1902) (not Parphorus Godman, 1900), Phlebodes fuldai (E. Bell, 1930) (not Vettius Godman, 1901), Mnasitheus padus (Evans, 1955) (not Moeris Godman, 1900), Naevolus brunnescens (Hayward, 1939) (not Psoralis Mabille, 1904), Lamponia ploetzii (Capronnier, 1874) (not Vettius Godman, 1901), Mnestheus silvaticus Hayward, 1940 (not Ludens Evans, 1955), Rigga spangla (Evans, 1955) (not Sodalia Evans, 1955), Corticea vicinus (Plötz, 1884) (not Lento Evans, 1955), Mnasalcas thymoetes (Hayward, 1942) (not Mnasicles Godman, 1901), Mnasalcas boyaca (Nicolay, 1973) (not Pamba Evans, 1955), Vertica brasta (Evans, 1955) (not Lychnuchus Hübner, [1831]), Carystina discors Plötz, 1882 (not Cobalus Hübner, [1819]), Zetka irena (Evans, 1955) (not Neoxeniades Hayward, 1938), and Neoxeniades parna (Evans, 1955) (not Niconiades Hübner, [1821]). The following are new or revised species-subspecies combinations: Tagiades neira moti Evans, 1934, Tagiades neira canonicus Fruhstorfer, 1910, Tagiades sheba vella Evans, 1934, Tagiades sheba lola Evans, 1945, Tagiades korela biakana Evans, 1934, Tagiades korela mefora Evans, 1934, Tagiades korela suffusus Rothschild, 1915, Tagiades korela brunta Evans, 1949, Tagiades ravi ravina Fruhstorfer, 1910, Tagiades atticus carnica Evans, 1934, Tagiades atticus nankowra Evans, 1934, Tagiades atticus helferi C. Felder, 1862, Tagiades atticus balana Fruhstorfer, 1910, Tagiades inconspicua mathias Evans, 1934, Tagiades hovia kazana Evans, 1934, Tagiades elegans fuscata de Jong and Treadaway, 2007, Tagiades elegans semperi Fruhstorfer, 1910, Metron hypochlora tomba Evans, 1955, Decinea denta pruda Evans, 1955, and Choranthus orientis eleutherae (Bates, 1934) (previously in Pyrrhocalles Mabille, 1904). In addition to the abovementioned changes, the following new combinations involve newly proposed genus group names: Fulvatis fulvius (Plötz, 1882) and Fulvatis scyrus (E. Bell, 1934) (not Salatis Evans, 1952); Adina adrastor (Mabille and Boullet, 1912) (not Bungalotis Watson, 1893); Nascus (Praxa) prax Evans, 1952, Nascus (Bron) broteas (Cramer, 1780), and Nascus (Bron) solon (Plötz, 1882) (not Pseudonascus Austin, 2008); Chirgus (Turis) veturius (Plötz, 1884); Paches (Tiges) liborius (Plötz, 1884), and Paches (Tiges) mutilatus (Hopffer, 1874) (not Antigonus Hübner, [1819]); Paches (Tiges) exosa (A. Butler, 1877); Tolius tolimus (Plötz, 1884) and Tolius luctuosus (Godman & Salvin, 1894) (not Echelatus Godman and Salvin, 1894); Ancistroides (Ocrypta) caerulea (Evans, 1928), Ancistroides (Ocrypta) renardi (Oberthür, 1878), Ancistroides (Ocrypta) waigensis (Plötz, 1882), Ancistroides (Ocrypta) aluensis (Swinhoe, 1907), Ancistroides (Ocrypta) flavipes (Janson, 1886), and Ancistroides (Ocrypta) maria (Evans, 1949) (not Notocrypta de Nicéville, 1889); Lennia lena (Evans, 1937), Lennia binoevatus (Mabille, 1891), Lennia maracanda (Hewitson, 1876), and Lennia lota (Evans, 1937) (not Leona Evans, 1937); Trida barberae (Trimen, 1873) and Trida sarahae (Henning and Henning, 1998) (not Kedestes Watson, 1893); Noxys viricuculla (Hayward, 1951) (not Oxynthes Godman, 1900); Xeniades (Tixe) quadrata (Herrich-Schäffer, 1869), Xeniades (Tixe) rinda (Evans, 1955), Xeniades (Tixe) putumayo (Constantino and Salazar, 2013) (not Tisias Godman, 1901); Gracilata quadrinotata (Mabille, 1889) (not Styriodes Schaus, 1913); Hermio hermione (Schaus, 1913) (not Lento Evans, 1955); Cynea (Nycea) hycsos (Mabille, 1891), Cynea (Nycea) corisana (Plötz, 1882), Cynea (Nycea) popla Evans, 1955, Cynea (Nycea) iquita (E. Bell, 1941), Cynea (Nycea) robba Evans, 1955, Cynea (Nycea) melius (Geyer, 1832), and Cynea (Nycea) irma (Möschler, 1879); Eutus rastaca (Schaus, 1902) (not Eutychide Godman, 1900); Eutus yesta (Evans, 1955) (not Thoon Godman, 1900); Eutus mubevensis (E. Bell, 1932) (not Tigasis Godman, 1900); Gufa gulala (Schaus, 1902) (not Mucia Godman, 1900); Gufa fusca (Hayward, 1940) (not Tigasis Godman, 1900); Godmia chlorocephala (Godman, 1900) (not Onophas Godman, 1900); Rhomba gertschi (E. Bell, 1937) (not Justinia Evans, 1955); Mnasicles (Nausia) nausiphanes (Schaus, 1913) (not Tigasis Godman, 1900); Amblyscirtes (Flor) florus (Godman, 1900) (not Repens Evans, 1955); Rectava ignarus (E. Bell, 1932) (not Papias Godman, 1900); Rectava vorgia (Schaus, 1902) (not Cobalopsis Godman, 1900); Rectava nostra (Evans, 1955) (not not Vidius Evans, 1955); Lerema (Geia) geisa (Möschler, 1879) and Lerema (Geia) lyde (Godman, 1900) (not Morys Godman, 1900); Contrastia distigma (Plötz, 1882) (not Cymaenes Scudder, 1872); Mit (Mit) badius (E. Bell, 1930) (not Styriodes Schaus, 1913); Mit (Mit) gemignanii (Hayward, 1940), (not Mnasitheus Godman, 1900); Mit (Rotundia) schausi (Mielke and Casagrande, 2002), (not Enosis Mabille, 1889); Picova steinbachi (E. Bell, 1930) (not Saturnus Evans, 1955); Lattus arabupuana (E. Bell, 1932) (not Eutocus Godman, 1901); Gubrus lugubris (Lindsey, 1925) (not Vehilius Godman, 1900); Thargella (Pseudopapias) tristissimus (Schaus, 1902) (not Papias Godman, 1900); Koria kora (Hewitson, 1877) (not Justinia Evans, 1955); Justinia (Septia) septa Evans, 1955; Corta lycortas (Godman, 1900) (not Orthos Evans, 1955); Vertica (Brasta) brasta (Evans, 1955) (not Lychnuchus Hübner, [1831]); Calvetta calvina (Hewitson, 1866) (not Cobalus Hübner, [1819]); Neoxeniades (Bina) gabina (Godman, 1900) (not Orthos Evans, 1955); Oz ozias (Hewitson, 1878) and Oz sebastiani Salazar and Constantino, 2013 (not Lychnuchoides Godman, 1901); and Carystoides (Balma) balza Evans, 1955 and Carystoides (Balma) maroma (Möschler, 1877). Finally, unless stated otherwise, all subgenera, species, subspecies and synonyms of mentioned genera and species are transferred together with their parent taxa, and taxa not mentioned in this work remain as previously classified.
Obligate endoparasitic oomycetes are known to ubiquitously occur in marine and freshwater diatoms, but their diversity is still largely unexplored. Many of these parasitoids are members of the early-diverging oomycete lineages (Miracula, Diatomophthora), others are within the Leptomitales of the Saprolegniomycetes (Ectrogella, Lagenisma) and some have been described in the Peronosporomycetes (Aphanomycopsis, Lagenidium). Even though some species have been recently described and two new genera were introduced (Miracula and Diatomophthora), the phylogeny and taxonomy of most of these organisms remain unresolved. This is contrasted by the high number of sequences from unclassified species, as recently revealed from environmental sequencing, suggesting the presence of several undiscovered species. In this study, a new species of Miracula is reported from a marine centric diatom (Minidiscus sp.) isolated from Skagaströnd harbor in Northwest Iceland. The morphology and life cycle traits of this novel oomycete parasite are described herein, and its taxonomic placement within the genus Miracula is confirmed by molecular phylogeny. As it cannot be assigned to any previously described species, it is introduced as Miracula islandica in this study. The genus Miracula thus contains three described holocarpic species (M. helgolandica, M. islandica, M. moenusica) to which likely additional species will need to be added in the future, considering the presence of several lineages known only from environmental sequencing that clustered within the Miracula clade.
The Afrotropical species of the sphecid genus Sphex are revised. Forty-six taxa are recognized, of which fifteen are newly described: Sphex abbotti nivarius subsp. nov., S. comorensis sp. nov., S. hades sp. nov., S. nefrens sp. nov., S. occidentalis sp. nov., S. pseudopraedator sp. nov., S. pseudosatanas sp. nov., S. pulawskii sp. nov., S. rufoclypeatus sp. nov., S. satanas memnon subsp. nov., S. schmideggeri sp. nov., S. schoutedeni malawicus subsp. nov., S. socotrensis sp. nov., S. stadelmanni rufus subsp. nov. and S. victoria sp. nov. A new subgenus, Menkeella subgen. nov., is proposed for S. paulinierii Guérin Méneville, 1843.
Four former synonyms are resurrected: Sphex pruinosus var. haemorrhoidalis Magretti, 1898, for which the replacement name of S. feijeni nom. nov. is proposed, S. camerunicus Strand, 1916, S. cinerascens Dahlbom, 1843 and S. abbotti W. Fox, 1891.
Nine of the previously valid names are shown to be junior synonyms: Sphex haemorrhoidalis basuto (Arnold, 1947) and S. haemorrhoidalis kobrowi (Arnold, 1928) = S. umtalicus Strand, 1916; S. incomptus anonymus Leclercq, 1955 = S. nigrohirtus Kohl, 1895; S. neavei (Arnold, 1928) = S. abbotti W. Fox, 1891; S. observabilis (R. Turner, 1918) = S. ahasverus Kohl, 1890; S. rufiscutis (R. Turner, 1918) and S. mochii Giordani Soika, 1942 = S. jansei Cameron, 1910; S. rufiscutis laevigatus Arnold, 1951 = S. gaullei Berland, 1927; and S. stadelmanni integer (Arnold, 1928) = S. stadelmanni Kohl, 1895. Three previously synonymized species are transferred as new synonyms to different species: Sphex conradti Berland, 1927 = S. camerunicus Strand, 1916; S. kilimandjaroensis Cameron, 1908 = S. abbotti W. Fox, 1891; and S. nigripes var. pachyderma Strand, 1916 = S. umtalicus Strand, 1916. For three species, the actual type localities are on different continents than their labels indicate: Sphex ahasverus Kohl, 1890, S. castaneipes Dahlbom, 1843 and S. optimus F. Smith, 1856, with the first occuring in Africa instead of Australia and the other two presumably occuring in South America instead of Africa.
The occurrence of the argentatus group in Sub-Saharan Africa is confirmed, and seven additional species groups are established, with all but one of them based on apomorphic morphological traits. A hypothesis on the phylogenetic relationships among the subgenera and species groups of Sphex is presented, and an identification key for both sexes as well as a placoid-based identification table for males are presented. Using Bayesian inference and maximum likelihood analysis on mitochondrial and nuclear sequence data generated from thirty-one of the recognized taxa, the monophyly of the proposed species groups is corroborated.
Pseudocetherinae (Hemiptera: Reduviidae) revisited: phylogeny and taxonomy of the lobe-headed bugs
(2022)
The concept of the previously monogeneric subfamily Pseudocetherinae (Hemiptera: Heteroptera: Reduviidae) is revised and expanded. We here transfer Gerbelius Distant, 1903, Kayanocoris Miller, 1954, Microvarus Jeannel, 1917, Paragerbelius Miller, 1958, and Voconia Stål, 1866 from Reduviinae to Pseudocetherinae and treat Kayanocoris, Microvarus, Paragerbelius, and Pseudocethera Villiers, 1963 as junior synonyms of Voconia, resulting in new combinations for Voconia conradti (Jeannel, 1917) comb. nov., V. ifana (Villiers, 1963) comb. nov., V. monodi (Villiers, 1963) comb. nov., V. motoensis (Schouteden, 1929) comb. nov., V. ornata (Distant, 1903) comb. nov., V. schoutedeni (Villiers, 1964) comb. nov., V. typica (Miller, 1958) comb. nov., and V. wegneri (Miller, 1954) comb. nov. We also describe 23 new species of Voconia: V. bakeri sp. nov., V. bracata sp. nov., V. brachycephala sp. nov., V. chrysoptera sp. nov., V. coronata sp. nov., V. decorata sp. nov., V. dolichocephala sp. nov., V. fasciata sp. nov., V. grandioculata sp. nov., V. hemera sp. nov., V. isosceles sp. nov., V. laosensis sp. nov., V. lasiosoma sp. nov., V. lirophleps sp. nov., V. loki sp. nov., V. mexicana sp. nov., V. minima sp. nov., V. nyx sp. nov., V. smithae sp. nov., V. tridens sp. nov., V. trinidadensis sp. nov., V. tuberculata sp. nov., and V. vittata sp. nov. Lectotypes are designated for Gerbelius confluens Distant, 1903, G. typicus Distant, 1903, V. conradti comb. nov., V. ornata comb. nov., and V. pallidipes Stål, 1866. A revised diagnosis and description of Pseudocetherinae are provided along with photographs of the species and of the male genitalia of 13 pseudocetherine and five closely related reduviine species. An identification key to the two genera of Pseudocetherinae as well as a key to species of Voconia are presented. A phylogenetic hypothesis is proposed for the relationships of Pseudocetherinae using parsimony analyses of 77 morphological characters.
Two species, namely Russula adwanitekae A.Ghosh, K.Das & Buyck sp. nov. and Russula purpureozonata K.Das, A.Ghosh & Buyck sp. nov. are proposed herein as new mushroom taxa from the Indian Himalaya based on their morphological features and ITS-based phylogenetic inferences. Both species belong to the crown clade of Russula subgenus Russula but with affinities to different subsections, viz. subsect. Laricinae Romagn. and subsect. Decolorantes Maire, respectively. Russula adwanitekae sp. nov. was collected in mixed temperate forests where it is most likely associated with conifers. It is distinct from several similarly looking, small, mild species with dark spore print and reddish lilac, orchid purple or greyish to deep magenta colored pileus in subsect. Laricinae by its sequence data (nrITS) or geographic distribution. Russula purpureozonata sp. nov. associates with Abies densa Griff., and possesses all typical features of Decolorantes, viz. the amyloid suprahilar spot on the spores, presence of pileogloeocystidia, the reddening then blackening context, equal lamellae and colored spore print. It reminds of North American R. californiensis Burl. and R. magna Beardslee under the microscope but has distinctly smaller spores and differs further in the unique coloration and concentrically zonated pileus margin. Macro- and micromorphological features are illustrated for both species. Their habitats, distributions and relationships with allied species are discussed.
Genomic sequencing and analysis of worldwide skipper butterfly (Lepidoptera: Hesperiidae) fauna points to imperfections in their current classification. Some tribes, subtribes and genera as they are circumscribed today are not monophyletic. Rationalizing genomic results from the perspective of phenotypic characters suggests two new tribes, two new subtribes and 50 new genera that are named here: Ceratrichiini Grishin, trib. n., Gretnini Grishin, trib. n., Falgina Grishin, subtr. n., Apaustina Grishin, subtr. n., Flattoides Grishin, gen. n., Aurivittia Grishin, gen. n., Viuria Grishin, gen. n., Clytius Grishin, gen. n., Incisus Grishin, gen. n., Perus Grishin, gen. n., Livida Grishin, gen. n., Festivia Grishin, gen. n., Hoodus Grishin, gen. n., Anaxas Grishin, gen. n., Chiothion Grishin, gen. n., Crenda Grishin, gen. n., Santa Grishin, gen. n., Canesia Grishin, gen. n., Bralus Grishin, gen. n., Ladda Grishin, gen. n., Willema Grishin, gen. n., Argemma Grishin, gen. n., Nervia Grishin, gen. n., Dotta Grishin, gen. n., Lissia Grishin, gen. n., Xanthonymus Grishin, gen. n., Cerba Grishin, gen. n., Avestia Grishin, gen. n., Zetka Grishin, gen. n., Turmosa Grishin, gen. n., Mielkeus Grishin, gen. n., Coolus Grishin, gen. n., Daron Grishin, gen. n., Barrolla Grishin, gen. n., Brownus Grishin, gen. n., Tava Grishin, gen. n., Rigga Grishin, gen. n., Haza Grishin, gen. n., Dubia Grishin, gen. n., Pares Grishin, gen. n., Chitta Grishin, gen. n., Artonia Grishin, gen. n., Lurida Grishin, gen. n., Corra Grishin, gen. n., Fidius Grishin, gen. n., Veadda Grishin, gen. n., Tricrista Grishin, gen. n., Viridina Grishin, gen. n., Alychna Grishin, gen. n., Ralis Grishin, gen. n., Testia Grishin, gen. n., Buzella Grishin, gen. n., Vernia Grishin, gen. n., and Lon Grishin, gen. n. In addition, the following taxonomic changes are suggested. Prada Evans is transferred from Hesperiinae to Trapezitinae. Echelatus Godman and Salvin, Systaspes Weeks, and Oenides Mabille are removed from synonymy and are treated as valid genera. The following genera are new junior subjective synonyms: Tosta Evans of Eantis Boisduval; Turmada Evans of Neoxeniades Hayward, Arita Evans of Tigasis Godman, and Alera Mabille of Perichares Scudder. Eantis pallida (R. Felder) (not Achlyodes Hübner), Gindanes kelso (Evans) (not Onenses Godman and Salvin), Isoteinon abjecta (Snellen) (not Astictopterus C. and R. Felder), Neoxeniades ethoda (Hewitson) (not Xeniades Godman), Moeris anna (Mabille) (not Vidius Evans), and Molo pelta Evans (not Lychnuchus Hübner) are new genus-species combinations. The following are species-level taxa: Livida assecla (Mabille) (not a subspecies of Livida grandis (Mabille), formerly Pythonides Hübner) and Alychna zenus (E. Bell) (not a junior subjective synonym of Alychna exclamationis (Mabille), formerly Psoralis Mabille); and Barrolla molla E. Bell (formerly Vacerra Godman) is a junior subjective synonym of Barrolla barroni Evans (formerly Paratrytone Godman). All these changes to taxonomic status of names are propagated to all names currently treated as subspecies (for species), subgenera (for genera) and synonyms of these taxa. Finally, taxa not mentioned in this work are considered to remain at the ranks and in taxonomic groups they have been previously assigned to.
The New World genus Axina Kirby (Coleoptera: Cleridae) is revised for the first time. Thirty-two new species are described: Axina acutipennis, A. adelosa, A. atmis, A. bahia, A. bella, A. brunnea, A. chiasta, A. furcula, A. heveli, A. ignota, A. klisis, A. latilinea, A. lobispinula, A. luzia, A. macilenta, A. megaspina, A.minas, A. ochra, A. oligocheia, A. ordinis, A. orcastomata, A. pallidioccabus, A. phallospina, A. piperata, A. pollex, A. polycaula, A. rio, A. schenklingi, A. spina, A. trinalis, A. villa, and A. vista. The 19 previously described species are: Axina analis Kirby, A. apicalis Pic, A. basalis Schenkling, A. bifasciata (Chevrolat), A. centrimaculata Schenkling, A. conspicua Schenkling, A. diversesignata Pic, A. equestris (Schenkling), A. fasciata Kirsch, A. fortipes Pic, A. lateralis Pic, A. longevittata Pic, A. munda Schenkling, A. nigrifrons Schenkling, A.parcepunctata Schenkling, A. picta Schenkling, A. plagiata Schenkling, A. proxima (Chevrolat), and A. sexmaculata Spinola. Two species Priocera equestris Schenkling and Priocera proxima Chevrolat, are transferred into the genus Axina becoming new combinations: Axina equestris (Schenkling) and Axina proxima (Chevrolat). Two names, Priocera podagrica Schenkling, 1900, and P. podagrica variety pygmaea Schenkling, 1902, are new synonymies of Priocera proxima Chevrolat, 1876. Lectotypes are here designated for nine species: Axina basalis Schenkling, 1900; Axina centrimaculata Schenkling, 1900; Axina conspicua Schenkling, 1900; Axina diversesignata Pic, 1946; Axina munda Schenkling, 1900; Axina nigrifrons Schenkling, 1906; Axina parcepunctata Schenkling, 1900; Axina picta Schenkling, 1907; and Axina plagiata Schenkling, 1900. It is proposed that Axina species are predators of lignicolous insects, particularly bark beetles. The species of Axina can be classified into eight species groups and a theory of their phylogenetic relationships is proposed via WINCLADA in conjunction with NONA. Of the 51 species that now comprise Axina, only one traversed the Panamanian portal before the Colombian Andes reached their modern altitudes. This work includes a generic-level morphological analysis, brief treatise of natural history, key to species, comments about Axinazoogeography, and hypotheses of species-group phylogeny.
A new dipluran species, Plusiocampa (Plusiocampa) imereti Sendra & Barjadze sp. nov., from the deep zone in three caves in the Imereti region, Georgia, is described. This new troglobitic Plusiocampa is an addition to four others known Diplura from around the Black Sea region, two Dydimocampa and two Plusiocampa s. str. The present study also provides the first CO1 sequences for the Plusiocampinae taxa and the first molecular data for cave-dwelling Plusiocampa species. Although bootstrap values were low, the maximum-likelihood phylogenetic tree grouped Plusiocampa (P.) imereti Sendra & Barjadze sp. nov. with two Plusiocampa s. str. species from Eastern Europe. Morphologically, P. (P.) imereti Sendra & Barjadze sp. nov. is closely related to two cave-dwelling species: Plusiocampa (Plusiocampa) glabra Condé, 1984 and Plusiocampa (P.) chiosensis Sendra & Gasparo, 2020. The new species can be distinguished by the presence of lateral anterior macrosetae on metanotum, more uneven claws, and the presence of 2+2 lateral anterior macrosetae on middle urotergites. The five species currently known for the Black Sea region inhabit caves located at low altitude but with no influence from former glacial or permafrost processes.
Numerous pseudoplasmodia containing myxospores belonging to the genus Cystodiscus were found in the gallbladder of Elachistocleis cesarii from Mato Grosso State, Brazil. Herein, we describe Cystodiscus elachistocleis sp. nov., using morphological and small subunit ribosomal DNA sequences. The mature myxospores were ellipsoid to ovoid, measuring in average 10.6 (9.8–11.2) μm in length and 6.2 (5.6–6.6) μm in width. Polar capsules were pyriform and equal in size measuring in average 3.6 (2.8–4.3) μm in length and 2.6 (2.2–3.1) μm in width. Polar filaments had 2–4 coils. The myxospores had 3–5 transverse ridges. The phylogenetic analysis showed Cystodiscus elachistocleis sp. nov. as a sister species of Cystodiscus cf. immersus 1, in a subclade formed by species that parasitize the amphibians gallbladder. This is the first species of Cystodiscus described parasitizing a species of Elachistocleis and the third species of Cystodiscus described in Brazil.
Five new species of the terrestrial snail genus Landouria Godwin-Austen, 1918 (Camaenidae) are described from northeastern Thailand, based on shell features, radular morphology, genital anatomy, and DNA sequence data: Landouria circinata sp. nov., L. tuberculata sp. nov., L. trochomorphoides sp. nov., L. chloritoides sp. nov., and L. elegans sp. nov. These species are phylogenetically well separated from each other by mtDNA phy-logeny and COI sequence divergences of 0.073–0.156. The record of Thaitropis Schileyko, 2004 (currently synonymized with Landouria) in Thailand is re-interpreted as referring to L. diplogramma (Möllendorff, 1902) comb. nov.
The genus Salvadora has not been subjected to a modern phylogenetic analysis. Described in 1853, its taxonomic history is complex and confusing. In this study, we evaluate the monophyly of the genus and present the first phylogenetic hypothesis based on an analysis of 66 qualitative and quantitative morphological characters of over 1000 specimens representing all described taxa across their entire distribution. Morphological characters were processed in Fast Morphology for subsequent phylogenetic analysis in PAUP under the maximum parsimony criterion. We obtained a single tree in which Salvadora appears as a monophyletic group with two clearly defined geographic species groups: a southern mexicana group and a northern grahamiae group. Based on our phylogenetic hypothesis, we evaluate the taxonomic status of all described taxa. Additionally, we include a diagnosis for all species, distribution maps, and an illustrated dichotomous taxonomic key of the genus.
Seven new species of the giant pill-millipede genus Zoosphaerium Pocock, 1895 are described from Madagascar: Z. nigrum sp. nov., Z. silens sp. nov., Z. ambatovaky sp. nov., Z. beanka sp. nov., Z. voahangy sp. nov., Z. masoala sp. nov. and Z. spinopiligerum sp. nov. All species are described based on drawings and scanning electron microscopy, while genetic barcoding of the COI gene was successful for six of the seven new species. Additional COI barcode information is provided for the first time for Z. album Wesener, 2009 and Z. libidinosum (de Saussure & Zehntner, 1897). Zoosphaerium nigrum sp. nov. and Z. silens sp. nov. belong to the Z. libidinosum species-group, Z. ambatovaky sp. nov. to the Z. coquerelianum species-group, Z. beanka sp. nov., Z. voahangy sp. nov. and Z. masoala sp. nov. to the Z. platylabum species-group and Z. spinopiligerum sp. nov. to the Z. piligerum species-group. Updated identification keys are provided for each species-group. Two western dry forest species, Z. silens sp. nov. and Z. voahangy sp. nov. are recorded from two localities, while the other five species are currently only known from their type localities. Of special conservation concern might be Z. ambatovaky sp. nov. from the lowland rainforest fragment of Ambatovaky, a nowadays isolated lowland rainforest, rapidly shrinking due to slash and burn agriculture. In addition to the new species, new locality data is provided for 11 species and numerous unidentifiable species of Zoosphaerium: Z. neptunus (Butler, 1872), Z. platylabum (de Saussure & Zehntner, 1902) and Z. piligerum (de Saussure & Zehntner, 1897) from the central eastern montane forests, as well as Z. ambrense Wesener, 2009, Z. aureum Wesener, 2009, Z. libidinosum, Z. corystoides Wesener, 2009, Z. discolor Wesener, 2009, Z. smaragdinum Wesener, 2009, Z. villosum Wesener & Sierwald, 2005 and Microsphaerotherium anjozorobe Wesener, 2009.
A new millipede species of the genus Sechelleptus Mauriès, 1980 is described and illustrated from Mayotte Island, Indian Ocean. This new species, S. arborivagus sp. nov., found on trees, looks particularly similar to the sympatric S. variabilis VandenSpiegel & Golovatch, 2007, but is much larger and has a very different ecological behavior. Phylogenetic analyses based on a concatenated dataset of the COI and 16S rRNA genes and including nine species of Spirostreptidae (including Sechelleptus, Doratogonus Attems, 1914, Bicoxidens Attems, 1928 and Spirostreptus Brandt, 1833), strongly support the monophyly of Sechelleptus. Despite the similarity of their genitalia, the molecular analyses also reveal a clear-cut genetic divergence between S. arborivagus sp. nov. and S. variabilis (22.55% for COI and 6.63% for 16SrRNA) and further suggest the presence of a higher diversity within the genus Sechelleptus on Mayotte.
New species, revision, and phylogeny of Ronzotherium Aymard, 1854 (Perissodactyla, Rhinocerotidae)
(2021)
Ronzotherium is one of the earliest Rhinocerotidae in Europe, which first appeared just after the Eocene/Oligocene transition (Grande Coupure), and became extinct at the end of the Oligocene. It is a large-sized rhinocerotid, with a special position in the phylogeny of this group, as being one of the earliest-branching true Rhinocerotidae. However, its intra-generic systematics has never been tested through computational phylogenetic methods and it is basically unknown. Its taxonomical history has gone through numerous complications, and thus we aim to provide here a complete revision of this genus, through phylogenetic methods. After a re-examination of all type specimens (five supposed species) as well as of most well-preserved specimens from all over Europe and ranging through the complete Oligocene epoch, we performed a parsimony analysis to test the position of some problematic specimens. According to our results, five species can be distinguished, Ronzotherium velaunum (type species), R. filholi, R. elongatum and R. romani as well as a new species: R. heissigi sp. nov. We also drastically re-interpret its anatomy and show that the ‘short-limbed’ “Diaceratherium” massiliae, described from Southern France, can be considered as a junior synonym of R. romani. Finally, we exclude the Asian species “Ronzotherium” orientale and “Ronzotherium” brevirostre from Ronzotherium and we consider R. kochi as a junior synonym of R. filholi.
Remipedia is a stygobitic group commonly associated with coastal anchialine caves. This class consists of 12 genera, ten of which are found within the Lucayan Archipelago. Herein, we describe a new species within the genus Godzillius from Conch Sound Blue Hole, North Andros Island, Bahamas. Godzillius louriei sp. nov. is the third known remipede observed from a subseafloor marine cave, and the first from the Godzilliidae. Remipedes dwell within notoriously difficult to access cave habitats and thus integrative and comprehensive systematic studies at family or genus level are often absent in the literature. In this study, all species of Godzillius are compared using morphological and molecular approaches. Specifically, the feeding appendages of G. louriei sp. nov., G. fuchsi Gonzalez, Singpiel & Schlagner, 2013 and G. robustus Schram, Yager & Emerson, 1986 were examined using scanning electron microscopy (SEM). Species of Godzillius are identified based on the spines of maxilla 1 segment 4 and by the denticles on the lacinia mobilis of the left mandible. A molecular phylogeny using the mitochondrial 16S rRNA and nuclear histone 3 genes recovered G. louriei sp. nov. within the Godzillius clade and 16S genetic distances revealed a 13–15% difference between species of Godzillius.
Most valvatiform genera of the gastropod family Hydrobiidae are narrow-range taxa. One exception is the genus Arganiella, which is comprised of three congeners: the type species A. pescei from the Apennine Peninsula, A. wolfi from the Iberian Peninsula and A. tabanensis from the Balkans. The genus assignment of the latter two species was based on morphological similarities with A. pescei in the shell, operculum, radula and genitalia. Given that the morphology of hydrobiids is sometimes susceptible to convergence, this study re-evaluates the taxonomic status of species of Arganiella by analysing mitochondrial (mtCOI) and nuclear (18S rRNA) sequences of topotypes or near topotypes to infer their phylogenetic position. Our phylogenetic analyses depicted Arganiella as a non-monophyletic group within Hydrobiidae, and sequence divergence among the three species ranged from 14.5 to 16.7% for mtCOI and 2.0 to 3.8% for 18S. We also re-examined the extent of morphological variation among species of Arganiella and found a few differences among them and other valvatiform genera. Consequently, we propose two new genera for A. wolfi and A. tabanensis. Our results conflict with the classification of valvatiform hydrobiid species solely based on traditional phenotypical methods and suggest further taxonomic evaluation within a molecular framework.
A new avian chewing louse genus Apomyrsidea gen. nov. is described based on species parasitizing birds in the family Formicariidae. Diagnostic characteristics and phylogenetic analyses were used to evaluate and confirm the generic status and merit its recognition as unique and different from Myrsidea Waterston, 1915. Three species previously belonging to the genus Myrsidea are placed in the new genus Apomyrsidea gen. nov. and are discussed: Apomyrsidea circumsternata (Valim & Weckstein, 2013) gen. et comb. nov., Apomyrsidea isacantha (Valim & Weckstein, 2013) gen. et comb. nov. and Apomyrsidea klimesi (Sychra in Sychra et al., 2006) gen. et comb. nov.
Commonly reported as a household pest throughout the northern hemisphere, Willowsia nigromaculata (Lubbock) is among the most abundant and widely distributed springtails. However, taxonomic uncertainty due to incomplete morphological descriptions based on specimens from different continents may lead to incorrect identifications and/or prevent the recognition of distinct lineages within this morphospecies. Here, we perform the first comprehensive morphological and genetic comparison between W. nigromaculata specimens collected from North America and Europe. Morphological and genetic evidence reveals that populations in the United States and France represent two distinct nigromaculata-like species, but a phylogenetic analysis indicates both species may also be present in Canada. Based on these results, we redescribe W. nigromaculata from France, provide a description for Willowsia neonigromaculata sp. nov. from the United States, and propose new diagnostic characters for their separation, including the number of inner appendages on the maxillary sublobal plate. We also highlight the need for morphological and molecular investigations of additional populations to better understand the diversity and distribution of W. nigromaculata and related species.
We report a new myxozoan, Myxobolus opsaridiumi sp. nov., infecting the ornamental fish Opsaridium ubangiensis (Pellegrin, 1901) collected from the Anga River near the city of Yaounde, Cameroon. Plasmodia were found in the skin, muscles and spleen. The overall prevalence of infection was 54.7% (288 parasitized fish out of 526 examined). The myxospores were ovoid to subspherical in frontal view and lenticular in lateral view. The valves were symmetrical and relatively thick, without edge markings. The myxospore measurements were 10.7 ± 0.14 (10–11.5) μm long, 9 ± 0.15 (8–10) μm wide and 6.2 ± 0.7 (5.6–7.2) μm thick. The two ovoid polar capsules were equal in size, converging and opening together at the anterior end, measuring 5 ± 0.07 (4.3–6.0) μm long and 2.7 ± 0.07 (2.2–3.0) μm wide. Polar filaments were coiled from 5 to 7 turns. Histopathological analysis revealed no inflammatory reaction associated with the infection. A BLAST search found that the newly obtained 18 rDNA sequence had a low sequence similarity with available sequences for Myxobolus on GenBank. A phylogenetical analysis based on ribosomal DNA partial sequences showed that M. opsaridiumi sp. nov. is closely associated with several species of Myxobolus infecting cyprinid fish.
A new monospecific genus belonging to the family Linyphiidae Blackwell, 1859, Nihonella gen. nov., is described using an integrative taxonomic approach based on the species N. chika gen. et sp. nov. The new genus is endemic to Western Honshu, Japan, and it shows distinctive genitalic and somatic characters of other genera of the subfamily Erigoninae Emerton, 1882. Nihonella gen. nov. is found only in the twilight and transition zones of caves in Okayama and Nara Prefectures. The phylogenetic position of Nihonella gen. nov. within the subfamily Erigoninae, and its relationship as a sister clade of the species of the group of Savignia Blackwell, 1833 (sensu Millidge 1977), is discussed on the basis of both, morphological and molecular evidence.
Four new species and one new subspecies of tateid freshwater gastropods are described from the north of the South Island of New Zealand, Catapyrgus jami sp. nov., Opacuincola lisannea sp. nov., O. gretathunbergae sp. nov., O. mete kahurangi ssp. nov. and Obtusopyrgus farri sp. nov. The species are integratively defined based on a combination of shell morphological, anatomical and mitochondrial DNA data. Morphological and anatomical data were generated by morphometrics, scanning electron microscopy, as well as micro-computed tomography. The genetic data were basis of phylogenetic analyses and incorporated into the diagnoses. The new taxa occur in springs or spring-like habitats, i.e., shallow, slow-flowing sections of small streams except for O. mete kahurangi subsp. nov., which was collected from rough rocks in a river, where the snails sat in small depressions. None of the species exceeded 2.75 mm in length. Opacuincola gretathunbergae sp. nov. and Obtusopyrgus farri sp. nov. are pigmented and true crenobionts, while C. jami sp. nov. and the sympatric Opacuincola lisannea sp. nov. have eyes of reduced size and lack epidermal pigment, hence, probably dwell in the transitional zone of epigean and groundwaters.
Members of the ATP‐binding cassette (ABC) transporter superfamily translocate a broad spectrum of chemically diverse substrates. While their eponymous ATP‐binding cassette in the nucleotide‐binding domains (NBDs) is highly conserved, their transmembrane domains (TMDs) forming the translocation pathway exhibit distinct folds and topologies, suggesting that during evolution the ancient motor domains were combined with different transmembrane mechanical systems to orchestrate a variety of cellular processes. In recent years, it has become increasingly evident that the distinct TMD folds are best suited to categorize the multitude of ABC transporters. We therefore propose a new ABC transporter classification that is based on structural homology in the TMDs:
The family Hahniidae is reported from Thailand for the first time. The genus Hexamatia gen. nov. and two new species, Hexamatia seekhaow gen. et sp. nov. and Hahnia ngai sp. nov., are described and illustrated. DNA sequences are provided for all the species reported here. The phylogenetic position of the novel genus Hexamatia gen. nov. and its relation to Hahnia are discussed. Based on these results, a new combination is proposed for Hexamatia senaria (Zhang, Li & Zheng, 2011) gen. et comb. nov. = Hahnia senaria. Known distribution of the species Hahnia saccata Zhang, Li & Zheng, 2011, originally described from China, is expanded. A brief review and notes on the taxonomy of the six-eyed hahniids are included.
This paper reports on the genus Cobbionema Filipjev, 1922 in Sweden with the description of four species and a revision of the genus. Cobbionema acrocerca Filipjev, 1922 is relatively small in size, with a tail that has a conical proximal and a digitate distal section. Cobbionema cylindrolaimoides Schuurmans Stekhoven, 1950 is similar to C. acrocerca in most characters except having a larger body size and heavily cuticularized mandibles. Cobbionema brevispicula sp. nov. is characterised by short spicules and a conoid tail. Cobbionema acuminata sp. nov. is characterised by a long two-part spicule, a conical tail and three (one mid dorsal and two ventrosublateral) sharply pointed tines in the anterior chamber of the stoma that are located more anterior than in all the other species. We also present a molecular phylogeny of the family based on the nearly full-length 18S and the D2-D3 expansion segment of the 28S rRNA genes. Maximum Likelihood and Bayesian trees inferred from both genes strongly support a clade that included Cobbionema, Demonema Cobb, 1894 and Halichoanolaimus de Man, 1888 and another clade with Gammanema Cobb, 1920 and Latronema Wieser, 1954 nested together. None of the trees supported the monophyly of the subfamilies Choniolaiminae and Selachinematinae.
In the last few years, a sharp increase in the number of descriptions of new species of West African cone snails, particularly from the Cabo Verde Archipelago, has taken place. In previous studies, we used mitogenome sequences for reconstructing robust phylogenies, which comprised in total 120 individuals representing the majority of species (69.7%) described from this biogeographical region (except Angolan endemics) and grouped into seven genera within the family Conidae. Here, we add another 12 individuals representing endemic species that were missing in the previous studies. We use the phylogenies to identify monophyletic groups and a genetic divergence threshold (0.2% uncorrected p distance) to determine the number of valid species. As a result, the number of valid West African cone species could be drastically reduced to at least 40%, indicating that some recent poor-quality descriptions loosely based on phenotypic characters prone to convergence such as the shape and color patterns of the shell have contributed substantially to taxonomic inflation. Several previously accepted species with a reduced geographical distribution now become phenotypic forms of the remaining valid species, which increase their distribution ranges. In contrast, several cryptic species are now uncovered and described. For instance, Africonus insulae sp. nov. and Kalloconus canariensis sp. nov. are hereby introduced as new species. A detailed systematic account with illustrations and relevant information is presented. Lectotypes are designated for Conus trochulus and Conus irregularis, and neotypes for Conus crotchii and Conus diminutus. According to our results, it is strongly recommended that any future introduction of new taxa names for cone snails from West Africa should be supported by molecular and/or anatomical rather than exclusively shell morphological data. The taxonomic decisions here taken have direct implications for conservation and will eventually require re-evaluation of the Red List risk status of an important number of species.
We describe Croton calcareus Riina & Mateo-Ram. sp. nov., a new species in Croton section Cyclostigma (dragon's blood trees) from the state of Chiapas (Mexico). This species is a small tree growing in dry forest on calcareous substrates. Both morphological and molecular data support C. calcareus sp. nov. as a new species closely related to C. redolens, another dry forest taxon from northern Venezuela. We provide illustrations, a distribution map and suggestions for species conservation status. The new species along with Croton draco are the only known representatives of C. section Cyclostigma occurring in Mexico.
The subfamily Sepiolinae (Mollusca: Cephalopoda: Sepiolidae), currently containing the genera Sepiola Leach, 1817, Euprymna Steenstrup, 1887, Inioteuthis Verrill, 1881, Rondeletiola Naef, 1921 and Sepietta Naef, 1912, is characterized by the hectocotylization of the left dorsal arm, i.e., its transformation into a copulatory organ thanks to modifications of sucker/pedicel elements. The hectocotylus morphology varies to a great extent across genera and species. In particular, one to several pedicels in its proximal third lose their sucker and become highly and diversely modified in shape to constitute a copulatory apparatus. An evolutionary gradient was observed in the copulatory apparatus morphology, from the simple modification into a papilla of just one pedicel from the third element of the ventral sucker row (some nominal species of Euprymna) to a quite complex structure involving several variously modified pedicels from both the ventral and dorsal sucker rows (Inioteuthis). In some species, elements in the distal portion of the hectocotylus may also be highly modified, such as the columnar suckers in Euprymna. The hectocotylian diversity allows to distinguish nine groups of species that do not match the current generic subdivision of Sepiolinae. Additional morphological characters (number of sucker rows on arms, female bursa copulatrix, occurrence and shape of visceral light organs, etc.) corroborate the subdivision of Sepiolinae into nine subtaxa, i.e., genera. Accordingly, a cladogram is drawn to describe the possible phylogenetic relationships among the nine clades. To comply with these results, all current genera are redefined and four new genera are described, namely Adinaefiola gen. nov., Boletzkyola gen. nov., Eumandya gen. nov. and Lusepiola gen. nov.
The aquatic biodiversity of springs and groundwater systems of North Africa remains largely unexplored. In an earlier field survey of Tunisian springs, a new gastropod genus, Bullaregia, was discovered as a phylogenetically independent lineage of uncertain position within the family Hydrobiidae. Here, we provide taxonomic and phylogenetic assignments for three newly collected populations of hydrobiids from springs in northern Tunisia based on morphological, anatomical and genetic (mtCOI and 18S) data. Among these and specimens of Bullaregia, major differences were observed in male and female genitalia as well as in mtCOI sequences (divergence 8.0–9.1%). Based on these findings, we describe two new genera and three new species: Belgrandiellopsis chorfensis gen. et sp. nov., Belgrandiellopsis secunda gen. et sp. nov. and Biserta putealis gen. et sp. nov. In all our phylogenetic analyses, these three new species were well resolved as a monophyletic group together with Bullaregia tunisiensis. Unexpectedly, this clade emerged as sister to the European valvatiform genera Corbellaria and Kerkia and not to the recently discovered clade of groundwater, conchologically similar, species living in Bulgaria (Balkan Peninsula). These Tunisian species are each locally endemic and form part of a newly discovered clade which in future systematic studies could eventually be identified as a distinct hydrobiid subfamily.
Detailed description and illustrations of immature Trictenotoma Gray, 1832 (Trictenotomidae Blanchard, 1845) are presented for the first time, based on larvae and pupae of T. formosana Kriesche, 1919. Characters exhibited by the mature larva are similar to those described by Gahan (1908) for T. childreni Gray, 1832, which was based on a single specimen. The phylogenetic position of Trictenotomidae has varied among Scarabaeoidea, Chrysomeloidea and Tenebrionoidea, though recent studies place the family clearly among the latter. Features of the immature stages described here corroborate this placement. Evidence supports placement within or near the "salpingid group" (Pythidae, Salpingidae, Boridae, Pyrochroidae). Distinguishing features of the mature trictenotomid larva include the absence of stemmata, antennal sensorium, urogomphal pit(s) and lip, the presence of paired series of longitudinal ridges on the meso- and metathorax and abdominal tergites 1–8 and sternites 2–8, a paired arcuate row of 12–15 asperities on the anterior margin of sternite 9 and relatively short, upturned urogomphi. The systematic position of trictenotomids within the Tenebrionoidea Latreille, 1802 is confirmed. The phylogenetic relationships among Trictenotomidae and other “salpingid group” members (e.g., Pythidae Solier, 1834 and Salpingidae Leach, 1815) are highlighted and discussed, solving an almost two centuries old puzzle in Coleoptera systematics.
Abiinae is the second-largest subfamily in Cimbicidae, a small family of true sawflies (Tenthredinoidea). The subfamily is adequately defined, but the generic classification has been unstable. Currently, only two genera are regarded as valid: Abia Leach, 1817 and Allabia Semenov & Gussakovskij, 1937. We evaluate the generic classification of Abiinae in a phylogenetic context. A total of 32 species (out of 57 described for the subfamily), including the type species of Allabia, Allabia infernalis (Semenov, 1896), are scored for 150 adult morphological characters. Results show some resolution, but only few clades can be circumscribed by consistent character combinations. Most of the characters that have previously been used to define genera are not congruent; consequently, most suggested genus definitions appear to be random character state combinations and few natural groups can be identified. For these reasons, we treat Allabia syn. nov. as a junior synonym of Abia and make the following additional taxonomic changes: Abia infernalis Semenov, 1896 comb. rev. and Abia malaisei (Semenov & Gussakovskij, 1937) syn. nov. For the purpose of long-term stability of the classification of Abiinae, we recommend recognizing only one genus, Abia, within the subfamily.
Holocarpic oomycetes are poorly known but widespread parasites in freshwater and marine ecosystems. Most of the holocarpic species seem to belong to clades that diverge before the two crown lineages of the oomycetes, the Saprolegniomycetes and the Peronosporomycetes. Recently, the genus Miracula was described to accommodate Miracula helgolandica, a holocarpic parasitoid of Pseudo-nitzschia diatoms, which received varying support for its placement as the earliest-diverging oomycete lineage. In the same phylogenetic reconstruction, Miracula helgolandica was grouped with some somewhat divergent sequences derived from environmental sequencing, indicating that Miracula would not remain monotypic. Here, a second species of Miracula is reported, which was found as a parasitoid in the limnic centric diatom Pleurosira leavis. Its life-cycle stages are described and depicted in this study and its phylogenetic placement in the genus Miracula revealed. As a consequence, the newly discovered species is introduced as Miracula moenusica.
Olpidiopsis is a genus of obligate holocarpic endobiotic oomycetes. Most of the species classified in the genus are known only from their morphology and life cycle, and a few have been examined for their ultrastructure or molecular phylogeny. However, the taxonomic placement of all sequenced species is provisional, as no sequence data are available for the type species, O. saprolegniae, to consolidate the taxonomy of species currently placed in the genus. Thus, efforts were undertaken to isolate O. saprolegniae from its type host, Saprolegnia parasitica and to infer its phylogenetic placement based on 18S rDNA sequences. As most species of Olpidiopsis for which sequence data are available are from rhodophyte hosts, we have also isolated the type species of the rhodophyte-parasitic genus Pontisma, P. lagenidioides and obtained partial 18S rDNA sequences. Phylogenetic reconstructions in the current study revealed that O. saprolegniae from Saprolegnia parasitica forms a monophyletic group with a morphologically similar isolate from S. ferax, and a morphologically and phylogenetically more divergent species from S. terrestris. However, they were widely separated from a monophyletic, yet unsupported clade containing P. lagenidioides and red algal parasites previously classified in Olpidiopsis. Consequently, all holocarpic parasites in red algae should be considered to be members of the genus Pontisma as previously suggested by some researchers. In addition, a new species of Olpidiopsis, O. parthenogenetica is introduced to accommodate the pathogen of S. terrestris.
Morphological and allozyme analyses suggested the occurrence of a pseudocryptic species in the Lasioglossum villosulum (Kirby, 1802) species complex (Hymenoptera: Halictidae). We analysed the morphology of more than 1500 specimens and the DNA barcode fragment of the cytochrome c oxidase subunit I (COI) of 102 specimens of this species complex from several Palaearctic countries. Our phylogenetic tree reconstructions, based on maximum likelihood and Bayesian inference revealed one clade corresponding to all specimens morphologically identified as Lasioglossum medinai (Vachal, 1895) and one divergent specimen morphologically identified as Lasioglossum berberum (Benoist, 1941). The other specimens, morphologically identified as L. villosulum, aggregated into at least three other lineages in our phylogenetic trees. The tree-based species delineations methods based on the Generalized Mixed Yule Coalescent (GMYC) model and the Bayesian Poisson Tree Process (bPTP) identified five to ten candidate species within the L. villosulum species complex, with L. medinai and L. berberum consistently recognized as separated from all other candidate species. Diagnostic morphological differences were found among L. medinai, L. berberum and the remaining specimens identified as L. villosulum. No diagnostic morphological differences were found to distinguish the different phylogenetic candidate species or lineages found within L. villosulum and L. medinai. Thus, both genetic and morphological approaches support the existence of L. medinai and L. berberum as distinct species from L. villosulum.
Revision of the land snail genus Landouria Godwin-Austen, 1918
(Gastropoda, Camaenidae) from Java
(2019)
A revision of the land snail genus Landouria Godwin-Austin, 1918 (Camaenidae) from Java reveals that this group represents the most diverse land snail radiation on that island. Only six species of Landouria were recognized from Java in the last revision of the genus based on shell characters. Our investigation, which also considers the genitalia as well as DNA sequences, shows that the diversity in Java is much higher. Based on newly collected specimens as well as museum material, twenty-eight species of Landouria from Java are described and figured. To stabilize the nomenclature, neotypes are designated for L. winteriana (Pfeiffer, 1842) and L. rotatoria (Pfeiffer, 1842). Sixteen species are described as new to science, i.e., L. naggsi sp. nov., L. parahyangensis sp. nov., L. nusakambangensis sp. nov., L. petrukensis sp. nov., L. tholiformis sp. nov., L. madurensis sp. nov., L. abdidalem sp. nov., L. sewuensis sp. nov., L. tonywhitteni sp. nov., L. sukoliloensis sp. nov., L. nodifera sp. nov., L. pacitanensis sp. nov., L. zonifera sp. nov., L. pakidulan sp. nov., L. dharmai sp. nov. and L. menorehensis sp. nov. Landouria conoidea (Leschke, 1914) comb. nov., L. intumescens (Martens, 1867) comb. nov., L. moussoniana (Martens, 1867) comb. nov., L. schepmani (Möllendorff, 1897) comb. nov. and L. leucochila (Gude, 1905) comb. nov. are considered valid species of the genus Landouria for the first time. Plectotropis kraepelini Leschke, 1914 syn. nov. is considered a probable synonym of L. winteriana (Pfeiffer, 1842), P. trichotrochium Möllendorff, 1897 syn. nov. is a synonym of L. epiplatia (Möllendorff, 1897) and the preoccupied name Helix squamulosa Martens, 1867 syn. nov. is a synonym of L. madurensis sp. nov. We estimate that there are actually more than fifty species of Landouria in Java because many shell samples could not be classified and because no material is available from several regions of the island. A molecular phylogeny reveals that the species from Java do not form a monophyletic group, but that at least one species from Timor is nested within Javanese clades. This means that the Oriental Landouria crossed Wallace's line, the supposed border between the Oriental and Australo-Papuan regions, at least twice and supports the conclusion that Wallace's line does not represent a more severe barrier for terrestrial organisms than other straits through the archipelago. Within the Javanese clades, species from western and eastern Java are mixed, indicating frequent dispersals also within Java.
In this paper we describe Macrobiotus canaricus sp. nov., a new tardigrade species of the Macrobiotus hufelandi group from the Canary Islands. Moreover, with the use of DNA sequencing, we confirm that Macrobiotus recens Cuénot, 1932 represents the hufelandi group, even though eggs laid by this species do not exhibit the typical hufelandi group morphology. Our study is based on both classical taxonomic methods that include morphological and morphometric analyses conducted with the use of light and scanning electron microscopy, and on the analysis of nucleotide sequences of four molecular markers (three nuclear: 18S rRNA, 28S rRNA, ITS-2, and one mitochondrial: COI). Our analyses revealed that M. canaricus sp. nov. is most similar to Macrobiotus almadai Fontoura et al., 2008 from the Archipelago of the Azores, from which it differs by the absence of granulation patches on the external and internal surfaces of legs I–III as well as by the absence of a cuticular pore in the centre of the external patch on legs I–III. Molecular sequences allowed us to pinpoint the phylogenetic positions of M. canaricus sp. nov. and M. recens within the M. hufelandi group.
Biosynthetic gene content of the "Perfume Lichens" Evernia prunastri and Pseudevernia furfuracea
(2019)
Lichen-forming fungi produce a vast number of unique natural products with a wide variety of biological activities and human uses. Although lichens have remarkable potential in natural product research and industry, the molecular mechanisms underlying the biosynthesis of lichen metabolites are poorly understood. Here we use genome mining and comparative genomics to assess biosynthetic gene clusters and their putative regulators in the genomes of two lichen-forming fungi, which have substantial commercial value in the perfume industry, Evernia prunastri and Pseudevernia furfuracea. We report a total of 80 biosynthetic gene clusters (polyketide synthases (PKS), non-ribosomal peptide synthetases and terpene synthases) in E. prunastri and 51 in P. furfuracea. We present an in-depth comparison of 11 clusters, which show high homology between the two species. A ketosynthase (KS) phylogeny shows that biosynthetic gene clusters from E. prunastri and P. furfuracea are widespread across the Fungi. The phylogeny includes 15 genomes of lichenized fungi and all fungal PKSs with known functions from the MIBiG database. Phylogenetically closely related KS domains predict not only similar PKS architecture but also similar cluster architecture. Our study highlights the untapped biosynthetic richness of lichen-forming fungi, provides new insights into lichen biosynthetic pathways and facilitates heterologous expression of lichen biosynthetic gene clusters.
Neuropogonoid species in the lichen-forming fungal genus Usnea exhibit great morphological variation that can be misleading for delimitation of species. We specifically focused on the species delimitation of two closely-related, predominantly Antarctic species differing in the reproductive mode and representing a so-called species pair: the asexual U. antarctica and the sexual U. aurantiacoatra. Previous studies have revealed contradicting results. While multi-locus studies based on DNA sequence data provided evidence that these two taxa might be conspecific, microsatellite data suggested they represent distinct lineages. By using RADseq, we generated thousands of homologous markers to build a robust phylogeny of the two species. Furthermore, we successfully implemented these data in fine-scale population genomic analyses such as DAPC and fineRADstructure. Both Usnea species are readily delimited in phylogenetic inferences and, therefore, the hypothesis that both species are conspecific was rejected. Population genomic analyses also strongly confirmed separated genomes and, additionally, showed different levels of co-ancestry and substructure within each species. Lower co-ancestry in the asexual U. antarctica than in the sexual U. aurantiacoatra may be derived from a wider distributional range of the former species. Our results demonstrate the utility of this RADseq method in tracing population dynamics of lichens in future analyses.
A new classification of Ophiuroidea, considering family rank and above, is presented. The new family and superfamily taxa in O’Hara et al. (2017) were proposed to ensure a better readability of the new phylogeny but are unavailable under the provisions of the ICZN. Here, the morphological diagnoses to all 33 families and five superfamilies are provided. Ten new families, Ophiosphalmidae fam. nov., Ophiomusaidae fam. nov., Ophiocamacidae fam. nov., Ophiopteridae fam. nov., Clarkcomidae fam. nov., Ophiopezidae fam. nov., Ophiernidae fam. nov., Amphilimnidae fam. nov., Ophiothamnidae fam. nov. and Ophiopholidae fam. nov., are described. The family Ophiobyrsidae Matsumoto, 1915, not yet discovered in the previous publication, is added, based on new molecular data. A new phylogenetic reconstruction is presented. Definitions of difficult-to-apply morphological characters are given.
Synchroidae Lacordaire, 1859 is a taxonomically and biologically poorly known group. In the present paper, diagnostic characters used to separate genera are analysed and the phylogenetic relationships within this family are preliminarily investigated. Results suggest that the characteristic Synchroa pangu Hsiao, Li, Liu & Pang, 2016 can be removed to establish a new genus, Thescelosynchroa gen. nov. The new combination, T. pangu (Hsiao, Li, Liu & Pang) gen. et comb. nov., is proposed. The definitions of Synchroa Newman, 1838 and Synchroina Fairmaire, 1898 are revised. Moreover, morphological analysis and character comparison also suggest that the familial placement of Mallodrya subaenea Horn, 1888 is questionable. Six species are re-examined and rediagnosed: Synchroa chinensis Nikitsky, 1999, S. elongatula Nikitsky, 1999, S. formosana Hsiao, 2015, S. melanotoides Lewis, 1895, S. punctata Newman, 1838 and Synchroina tenuipennis Fairmaire, 1898. The male of S. chinensis and the female of S. formosana are described for the first time. Synchroa elongatula and Synchroina tenuipennis are newly recorded from Laos and Indonesia, respectively. We also hypothesize that the Eastern Asian-North American disjunction of Synchroa could be connected to a Mid-Late Tertiary migration of plants via the Bering Land Bridge.
The Australian wolf spider genus Tetralycosa Roewer, 1960, with Lycosa meracula Simon, 1909 (junior synonym of Lycosa oraria L. Koch, 1877) as type species, is revised to include 13 species, eight of which are described as new here: Tetralycosa adarca sp. nov., T. alteripa (McKay, 1976), T. arabanae Framenau, Gotch & Austin, 2006, T. baudinettei sp. nov., T. caudex sp. nov., T. eyrei (Hickman, 1944), T. floundersi sp. nov., T. halophila sp. nov., T. oraria (L. Koch, 1876), T. orariola sp. nov., T. williamsi sp. nov., T. wundurra (McKay, 1979) comb. nov. and T. rebecca sp. nov. Members of Tetralycosa are halotolerant, exclusively inhabiting saline environments such as coastal beaches, and mound springs, clay pans and salt lakes in the Australian interior. A phylogenetic analysis of the genus identified a monophyletic clade of eight species that live permanently on the barren surface of salt lakes suggesting a single radiation into this extremely inhospitable habitat. Some of these Tetralycosa species are currently known from single salt lakes only and with increasing disturbances of these systems by mining, agriculture an
With 280 accepted species, the genus Riccardia S.F.Gray (Aneuraceae) is one of the most speciose genera of simple thalloid liverworts. The current classification of this genus is based on morphological and limited-sampling molecular studies. Very few molecular data are available and a comprehensive view of evolutionary relationships within the genus is still lacking. A phylogeny focusing on relationships within the large genus Riccardia has not been conducted. Here we propose the first worldwide molecular phylogeny of the genus Riccardia, based on Bayesian inference and parsimony ratchet analyses of sequences from three plastid regions (psbA-trnH, rps4, trnL-F). The results support the monophyly of Riccardia and a new monospecific genus, Afroriccardia Reeb & Gradst. gen. nov., is described based on molecular and morphological evidence. The results indicate that several currently recognized infrageneric divisions and a few species are not monophyletic, suggesting that further analyses are needed to arrive at a proper understanding of the phylogeny of the genus. Although evidence for an Andean clade was found, most of the species appear scattered in different clades without clear geographical segregation. Broader sampling and further analyses are necessary in order to improve our understanding of the phylogeny of this poorly known liverwort genus.
Acrodiscus Zanardini is a poorly known monotypic endemic Mediterranean genus based on A. vidovichii (Menegh.) Zanardini. Rarely reported, its reproductive structures have remained undocumented, leaving its exact taxonomic position uncertain. Solely on the basis of its vegetative structure, Zanardini provisionally placed it in the family Cryptonemiaceae of the order Cryptonemiales (currently the Halymeniaceae of the Halymeniales), although he was uncertain as to whether the new genus actually belonged to that family or should instead be included in the Gigartinaceae of the Gigartinales (where Meneghini had originally placed it). In the present study we have extensively sampled A. vidovichii and documented its vegetative and tetrasporangial features. As well, we provide molecularsequence data (COI-5P, rbcL, LSU) that indicate its phylogenetic affinities. We confirm Acrodiscus as a member of the Halymeniaceae and its status as an independent genus. Searches of several institutional herbaria have allowed us to locate and lectotypify Meneghini’s Chondrus? vidovichii by the discovery of his original material now held at the Herbarium Horti Pisani (Pisa, Italy).
The Thyropygus opinatus subgroup (Diplopoda: Harpagophoridae) of the T. allevatus group in Thailand is revised. Based on a phylogenetic analysis of mtDNA sequence data, it is merged with the T. bifurcus subgroup to form an extended T. opinatus subgroup. Nine new species are described: Thyropygus cimi sp. nov. and T. forceps sp. nov. from Nakhonsrithammarat Province, T. culter sp. nov., T. planispina sp. nov., T. undulatus sp. nov. and T. ursus sp. nov. from Krabi Province, T. mesocristatus sp. nov. from Songkhla Province, T. navychula sp. nov. from Phang-Nga Province and T. sutchariti sp. nov. from Phetchaburi Province.
This study deals with the biodiversity and distribution of cavernicolous Amphipoda in caves of the Arabika massif (Western Caucasus). The Sarma, Trojka and Orlinoe Gnezdo caves were explored during speleological expeditions over the years 2011–12. Two new species of Amphipoda were found: a sub-surface dweller Zenkevitchia sandroruffoi sp. nov. is reported from the Sarma, Trojka and Orlinoe Gnezdo caves at depths from -30 m to -350 m; the second one, a deep dweller Adaugammarus pilosus gen. et sp. nov. is reported from the Sarma Cave at depths of -1270 to -1700 m. Adaugammarus gen. nov. shares similarities with Typhlogammarus Schäferna, 1907 and Zenkevitchia Birstein, 1940. The species Anopogammarus birsteini Derzhavin, 1945 is also re-described herein based on new samples that suggest close affinity of this species with the family Gammaridae. The original taxonomic combination is resurrected for Zenkevitchia revazi Birstein & Ljovuschkin, 1970, comb. resurr. (from Anopogammarus Derzhavin, 1945). To accommodate morphologically different species in the genus Zenkevitchia, two new groups are proposed. These are the admirabilis-group (Z. admirabilis Birstein, 1940 and Z. yakovi Sidorov, 2015) and the sandroruffoi-group (Z. sandroruffoi sp. nov. and Z. revazi). An updated molecular (mt-cox1) phylogeny, an identification key to the genera and a distribution map for the typhlogammarid amphipod species of Transcaucasia are provided.
Phylogenetic reconstruction from transposable elements (TEs) offers an additional perspective to study evolutionary processes. However, detecting phylogenetically informative TE insertions requires tedious experimental work, limiting the power of phylogenetic inference. Here, we analyzed the genomes of seven bear species using high-throughput sequencing data to detect thousands of TE insertions. The newly developed pipeline for TE detection called TeddyPi (TE detection and discovery for Phylogenetic Inference) identified 150,513 high-quality TE insertions in the genomes of ursine and tremarctine bears. By integrating different TE insertion callers and using a stringent filtering approach, the TeddyPi pipeline produced highly reliable TE insertion calls, which were confirmed by extensive in vitro validation experiments. Analysis of single nucleotide substitutions in the flanking regions of the TEs shows that these substitutions correlate with the phylogenetic signal from the TE insertions. Our phylogenomic analyses show that TEs are a major driver of genomic variation in bears and enabled phylogenetic reconstruction of a well-resolved species tree, despite strong signals for incomplete lineage sorting and introgression. The analyses show that the Asiatic black, sun, and sloth bear form a monophyletic clade, in which phylogenetic incongruence originates from incomplete lineage sorting. TeddyPi is open source and can be adapted to various TE and structural variation callers. The pipeline makes it possible to confidently extract thousands of TE insertions even from low-coverage genomes (∼10×) of nonmodel organisms. This opens new possibilities for biologists to study phylogenies and evolutionary processes as well as rates and patterns of (retro-)transposition and structural variation.
The New World genus Chariessa Forster (Coleoptera: Cleroidea: Cleridae) is revised and includes C. catalina Opitz, new species, C. elegans Horn, C. dichroa (LeConte), C. floridana Schaeffer, C. pilosa (Forster), C. texana Wolcott, C. ramicornis Perty, C. vestita (Chevrolat), and C. duponti (Spinola). Enoplium pilosa var. marginata Say is synonymized with Chariessa pilosa Forster. Lectotypes are designated for C. pilosa (Forster), C. ramicornis Perty, and C. vestita (Chevrolat). Available information indicates that Chariessa adult and immature individuals are predatory on lignicolous insects with a particular affinity for cerambycids and buprestids that infest species of oak. It is postulated that Pleistocene speciation generated the North American components of Chariessa with more ancient southern species generated during the Middle Tertiary; after closures of the Middle American portals and orogeny of the South American Andes. Included in this treatise is a discussion of natural history, key to species, narratives of zoogeography and phylogeny, one diagram of a phylogenetic tree, 35 line drawings, eight SEM micrographs, twelve habitus photographs, nine photographs of male genitalia, and five distributional maps.
Background: The current taxonomy of the African giraffe (Giraffa camelopardalis) is primarily based on pelage pattern and geographic distribution, and nine subspecies are currently recognized. Although genetic studies have been conducted, their resolution is low, mainly due to limited sampling. Detailed knowledge about the genetic variation and phylogeography of the South African giraffe (G. c. giraffa) and the Angolan giraffe (G. c. angolensis) is lacking. We investigate genetic variation among giraffe matrilines by increased sampling, with a focus on giraffe key areas in southern Africa.
Results: The 1,562 nucleotides long mitochondrial DNA dataset (cytochrome b and partial control region) comprises 138 parsimony informative sites among 161 giraffe individuals from eight populations. We additionally included two okapis as an outgroup. The analyses of the maternally inherited sequences reveal a deep divergence between northern and southern giraffe populations in Africa, and a general pattern of distinct matrilineal clades corresponding to their geographic distribution. Divergence time estimates among giraffe populations place the deepest splits at several hundred thousand years ago.
Conclusions: Our increased sampling in southern Africa suggests that the distribution ranges of the Angolan and South African giraffe need to be redefined. Knowledge about the phylogeography and genetic variation of these two maternal lineages is crucial for the development of appropriate management strategies.
We compared Chatham Island endemic species Xanthocnemis tuanuii to its congenerics from the New Zealand South Island: X. zealandica (newly collected specimens)and X. sinclairi (type specimens plus newly collected material). Two independent tests were performed –geometric morphometrics and molecular. Both analyses were consistent in supporting the status of X. tuanuiias a good species. Species differed statistically in the following morphological traits: head (dorsal view), male appendages (dorsal, lateral, posterior and ventral views), thorax (dorsal view), and penis (dorsal and lateral view). In addition to the original diagnostic features (mainly shape of the male superior appendages), a new morphological character is suggested here which reliably distinguishes the species based on the shape of the inferior appendages. There was no statistical support for the species status of X. sinclairi. The only feature re-ported as diagnostic (lower lobe of male superior appendages) was found to be variable and insufficient to warrant the previously proposed taxonomic rank for X. sinclairi. Molecular analysis of specimens showing identical appendages to the X. sinclairi holotype grouped them with X. zealandica specimens. Therefore X. sinclairi is synonymised with X. zealandica.
Three fossil leafhopper inclusions from Eocene Baltic amber, representing three new extinct genera and species, are described and illustrated. Eomegophthalmus lithuaniensis gen. et sp. nov. is tentatively placed in Megophthalminae, although it may represent the stem group from which Megophthalminae, Ulopinae, and Membracidae arose. Xestocephalites balticus gen. et sp. nov. and Brevaphrodella nigra gen. et sp. nov. are placed in Aphrodinae: Xestocephalini based on the structure of the head, leg chaetotaxy, and male genital capsule. These new genera and species represent the oldest known representatives of their respective subfamilies and the latter is the oldest known brachypterous adult leafhopper.
The classification of the largest subfamily of leafhoppers, Deltocephalinae, including 38 tribes, 923 genera, and 6683 valid species, is reviewed and revised. An updated phylogeny of the subfamily based on molecular (28S, Histone H3) and morphological data and an expanded taxon sample (37 taxa not included in previous analyses) is presented. Based on the results of these analyses and on the morphological examination of many representatives of the subfamily, the classification of the tribes and subtribes of Deltocephalinae is revised. Complete morphological descriptions, illustrations, lists of the included genera, and notes on their distribution, ecology, and important vector species are provided for the 38 recognized tribes and 18 subtribes. A dichotomous key to the tribes is provided. All names in the taxonomic treatments are hyperlinked to online resources for individual taxa which are supported by a comprehensive database for Deltocephalinae compiled using the taxonomic database software package 3I. The online functionality includes an interactive key to tribes and subtribes and advanced database searching options. Each taxon (subspecies through subfamily) has a unique taxon webpage providing nomenclatural information, lists of included taxa, an automated description (if available), images (if available), distributional information, bibliographic references and links to outside resources. Some observations and trends regarding the history of taxonomic descriptions in Deltocephalinae are reported. Four new tribes are described: Bahitini tribe nov. (25 genera), Bonsapeiini tribe nov. (21 genera), Phlepsiini tribe nov. (4 genera), and Vartini tribe nov. (7 genera). The circumscription and morphological characterization of Scaphoideini Oman, 1943 (61 genera) is substantially revised. Eleven new species are described: Acostemma stilleri sp. nov., Arrugada linnavuorii sp. nov., Drabescus zhangi sp. nov., Parabolopona webbi sp. nov., Goniagnathus emeljanovi sp. nov., Hecalus hamiltoni sp. nov., Scaphoideus omani sp. nov., Dwightla delongi sp. nov., Abimwa knighti sp. nov., Gannia viraktamathi sp. nov., and Doratulina dmitrievi sp. nov. Some family-group level taxonomic changes are made: Platymetopiini Haupt, 1929, Anoterostemmini Haupt, 1929, and Allygidiina Dmitriev, 2006 are synonymized with Athysanini Van Duzee, 1892, syn. nov.; Procepitini Dmitriev, 2002 is synonymized with Cicadulini Van Duzee, 1892, syn. nov.; Listrophorini Boulard, 1971 is synonymized with Chiasmini Distant, 1908, syn. nov.; Adamini Linnavuori & Al-Ne’amy, 1983, Dwightlini McKamey, 2003, and Ianeirini Linnavuori, 1978 are synonymized with Selenocephalini Fieber, 1872 syn.nov., and all three are now recognized as valid subtribes in their parent tribe. New placements of many genera to tribe and subtribe are made, and these are described in individual taxon treatments.
Cteniogaster, a new genus of small ground spiders is described from Kenya and Tanzania. It encompasses seven new species, three of which are known from both sexes: C. toxarchus sp. nov., the type species, C. conviva sp. nov. and C. hexomma sp. nov. Three species are known from females only: C. lampropus sp. nov., C. sangarawe sp. nov. and C. taxorchis sp. nov. and one only from males: C. nana sp. nov. The new genus can be recognised by the presence of a posterior ventral abdominal f eld of strong setae and anterior lateral spinnerets with enlarged piriform gland spigots in males. A cladistic analysis attributes the genus to Liocranidae, Cybaeodinae. The results of the analysis performed do not produce an unequivocal autapomorphy for Liocranidae, but provide a combination of non-homoplasious character changes that offers significant potential for recognising genera as Liocranidae. Moreover, robust apomorphies are determined within Liocranidae for the subfamilies Liocraninae and Cybaeodinae. Based on these fi ndings Toxoniella Warui & Jocqué, 2002 is transferred from Gallieniellidae to Liocranidae, Cybaeodinae. Jacaena Thorell, 1897, Plynnon Deeleman-Reinhold, 2001 and Teutamus Thorell, 1890 are transferred to Corinnidae, Phrurolithinae and Montebello Hogg, 1914 to Gnaphosidae. Itatsina Kishida, 1930 is synonymised with Prochora Simon, 1886.
Members of the balloon vine genus, Cardiospermum, have been extensively moved around the globe as medicinal and horticultural species, two of which are now widespread invasive species; C. grandiflorum and C. halicacabum. A third species, C. corindum, may also have significant invasion potential. However, in some regions the native status of these species is not clear, hampering management. For example, in South Africa it is unknown whether C. halicacabum and C. corindum are native, and this is a major constraint to on-going biological control programmes against invasive C. grandiflorum. We review the geography, biology and ecology of selected members of the genus with an emphasis on the two most widespread invaders, C. halicacabum and C. grandiflorum. Specifically, we use molecular data to reconstruct a phylogeny of the group in order to shed light on the native ranges of C. halicacabum and C. corindum in southern Africa. Phylogenetic analyses indicate that southern African accessions of these species are closely related to South American taxa indicating human-mediated introduction and/or natural long distance dispersal. Then, on a global scale we use species distribution modelling to predict potential suitable climate regions where these species are currently absent. Native range data were used to test the accuracy with which bioclimatic modelling can identify the known invasive ranges of these species. Results show that Cardiospermum species have potential to spread further in already invaded or introduced regions in Australia, Africa and Asia, underlining the importance of resolving taxonomic uncertainties for future management efforts. Bioclimatic modelling predicts Australia to have highly favourable environmental conditions for C. corindum and therefore vigilance against this species should be high. Species distribution modelling showed that native range data over fit predicted suitable ranges, and that factors other than climate influence establishment potential. This review opens the door to better understand the global biogeography of the genus Cardiospermum, with direct implications for management, while also highlighting gaps in current research.
Camel spiders (Arachnida: Solifugae) are one of the arachnid groups characterised by a prosomal dorsal shield composed of three distinct elements: the pro-, meso- and metapeltidium. These are associated respectively with prosomal appendages one to four, five, and six. What is less well known, although noted in the historical literature, is that the coxae of the 4th and 5th prosomal segments (i.e. walking legs 2 and 3) of camel spiders are also separated ventrally by a distinct membranous region, which is absent between the coxae of the other legs. We suggest that this essentially ventral division of the prosoma specifically between coxae 2 and 3 is homologous with the so-called sejugal furrow (the sejugal interval sensu van der Hammen). This division constitutes a fundamental part of the body plan in acariform mites (Arachnida: Acariformes). If homologous, this sejugal furrow could represent a further potential synapomorphy for (Solifugae + Acariformes); a relationship with increasing morphological and molecular support. Alternatively, outgroup comparison with sea spiders (Pycnogonida) and certain early Palaeozoic fossils could imply that the sejugal furrow defines an older tagma, derived from a more basal grade of organisation. In this scenario the (still) divided prosoma of acariform mites and camel spiders would be plesiomorphic. This interpretation challenges the textbook arachnid character of a peltidium (or ‘carapace’) covering an undivided prosoma.
Birds are characterized by pneumatization of their skeletons by epithelial diverticula from larger, air-filled cavities. The diverticula-or 'air sacs'-that invade the postcranium result from outgrowths of the lungs; poslcranial pneumaticity has been very well studied. Much more poorly understood are the air sacs that pneumatize the skull. Study or craniofacial pneumaticity in modern birds (Neornithes) indicates the presence of two separate systems: nasal pneumaticity and tympanic pneumaticity, The lacrimal and maxillary bones arc pneumatized by diverticula of the main paranasal cavity, the antorbital sinus. There are five tympanic diverticula in neornithines that pneumatize the quadrate, articulare and the bones of the braincase. The pneumatic features of the following six genera of Mesozoic birds are examined: Archaeopteryx, Ellaliornis, Baplomis, Parahesperornis, Hesperornis and lchthyornis. Despite the 'archaic' aspect of most of these birds, many of the pneumatic features of neornithines are found in .Mesozoic birds and are considered primitive for Aves. The phylogenetic levels at which most of the avian pneumatic features arose within Archosauria are uncertain. Until the phylogenetic levels at which homologous pneumatic features arose are determined, it is unwise to use most pneumatic characters in the discussion of avian origins. Within avian phylogeny, Ornithurae and Neornithes are well-supported by pneumatic synapomorphies. There is a trend towards reduction of craniofacial pneumaticity within Hesperornithiformes. Witthin Neornithes, four derived pneumatic characters suggest that the Palaeognathae (ratites and tinamous) is monophyletic.
Lepidoptera phylogeny and systematics : the state of inventorying moth and butterfly diversity
(2007)
The currently recognized robust support for the monophyly of the Lepidoptera (and the superorder Amphiesmenoptera comprising Lepidoptera + Trichoptera) is outlined, and the phylogeny of the principal lineages within the order is reviewed succinctly. The state of the taxonomic inventory of Lepidoptera is discussed separately for ‘micro-moths’, ‘macro-moths’ and butterflies, three assemblages on which work has followed historically somewhat different paths. While currently there are about 160,000 described species of Lepidoptera, the total number of extant species is estimated to be around half a million. On average, just over one thousand new species of Lepidoptera have been described annually in recent years. Allowing for the new synonyms simultaneously established, the net increase in species numbers still exceeds 800/year. Most of the additions are foreseeable in the micro-moth grade, but even for butterflies ca 100 species are added annually. Examples of particularly interesting new high-rank taxa that have been described (or whose significance has become realized) since the middle of the 20th century include the non-glossatan lineages represented by Agathiphaga and Heterobathmia and the heteroneuran families Andesianidae, Palaephatidae, Hedylidae and Micronoctuidae. Some thoughts on how present and future systematic lepidopterology might be prioritised are presented.
The taxonomy, diversity, and distribution of the aquatic insect order Trichoptera, caddisflies, are reviewed. The order is among the most important and diverse of all aquatic taxa. Larvae are vital participants in aquatic food webs and their presence and relative abundance are used in the biological assessment and monitoring of water quality. The species described by Linnaeus are listed. The morphology of all life history stages (adults, larvae, and pupae) is diagnosed and major features of the anatomy are illustrated. Major components of life history and biology are summarized. A discussion of phylogenetic studies within the order is presented, including higher classification of the suborders and superfamilies, based on recent literature. Synopses of each of 45 families are presented, including the taxonomic history of the family, a list of all known genera in each family, their general distribution and relative species diversity, and a short overview of family-level biological features. The order contains 600 genera, and approximately 13,000 species.
A cladistic analysis is presented of the hawkmoths of the tribe Acherontiini, Morgan´s Sphinx (Xanthopan morganii (Walker», and related genera. The study aims to test the monophyly of tribe Acherontiini; the hypothesis that all taxa with extremely long probosces (some Acherontiini, Meganoton rubescens, Neococytius, Xanthopan) form a monophyletic group, or at least fall within a single reasonably compact clade; and, within this group, to determine whether Xanthopan is more closely related to Acherontiini or to COCytillS and Neococytius. The data set comprises 109 characters derived from adult and immature stage morphology, biology and behaviour. These data were analysed using equal weighting, successive approximations character weighting (SACW) and implied weighting. All weighting schemes agreed on the monophyly of Acherontiini and of a group of genera comprising Amphimoea, Cocytius and Neococytius (the Cocytius group). Several other generic and suprageneric clades were also consistently recovered. However, those hawkmoths with extremely long probosces were never recovered as a monophyletic group. The relationships of Xanthopan were also ambiguous. Equal weighting and SACW placedXanthopan + Meganoton rztbescens (Butler) as sister to the COCytills group, while implied weighting placed Xanthopan as sister to Acherontiini. This latter relationship is based primarily on shared possession of a pilifer/palp hearing organ. Further analyses suggested the two components of this organ were not biologically independent. Downweighting this feature accordingly resulted in all weighting schemes converging on the topology found by equal weighting. Exclusion of the incomplete subset of immature stage data had no effect under implied weighting but equal weighting and SACW now recovered a Neotropical clade comprising Manduca. and the Cocytius group, while Xanthopan was placed with M. rubescens and Panogena. Downweighting the pilifer/palp hearing organ under implied weighting again caused convergence with the equal weighting/SACW results. Thus, the relationships of Xanthopan remain equivocal and further data, particularly from the immature stages, will be required to elucidate its phylogenetic position further.
Before the turn of the millenium the investigation of phylogenetic relationships was revolutionized by two major inputs, the use of molecular sequence data for phylogenetic reconstruction, paralleled by the sophistication of computer aided reconstruction methods. The ever growing number of data however did not only result in clarifications of open questions, but brought forth a number of new conflicting phylogenetic hypotheses. Sometimes they are wrongly referred to as conflicts between morphological and molecular approaches, which sporadically even culminated in the rejection of the usefulness of one of the two approaches (e.g. Scotland et al 2003). These scientists overlook the great advantage of having two a priori largely independent data sets (Wägele 2001) which in a synthetic way enable the greatest progress in phylogenetic research. However, solely putting data together will not suffice to choose among conflicting hypotheses. The increasing number of conflicts necessitates approaches that go beyond mere data congruence, but searching for the possible reasons of conflicts. In the present paper, problems in the reconstruction of the phylogenetic origin of Hexapoda, as well as of the early branchings within the Hexapoda, will exemplify approaches of critical re-evaluation and testing of data used in morphological data matrices for phylogenetic analyses. The early cladogenetic events of hexapods are especially suited for such a discussion for several reasons. The hexapods, as the most species-rich group of organisms, look back at a long and multi-faceted history of taxonomic and phylogenetic studies, culminating in a number of conflicting hypotheses. Triggered by incongruences with morphological analyses the reconstruction of the hexapodan roots likewise became a hot-spot of molecular research activities during^the last two decades. Furthermore the phylogenetic positions of the oldest lineages branching off within the hexapodan clade, the Diplura, Protura and Collembola, are in particular very difficult to reconstruct. While at least the latter two are well defined by morphological autapomorphies their phylogenetic position could not be reconstructed unambiguously, since their morphology seems highly derived with respect to the hexapodan ground pattern.