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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).