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The ‘acantherpestes’ group of dragon millipedes, formerly placed in the genus Desmoxytes Chamberlin, 1923, is revised and assigned to the new genus Nagaxytes Srisonchai, Enghoff & Panha gen. nov. Desmoxytes acantherpestes Golovatch & Enghoff, 1994 is the type species of the new genus and is redescribed as N. acantherpestes (Golovatch & Enghoff, 1994) gen. et comb. nov. Three new species are described from Thailand: N. erecta Srisonchai, Enghoff & Panha gen. et sp. nov. and N. gracilis Srisonchai, Enghoff & Panha gen. et sp. nov. from Kanchanaburi Province, and N. spatula Srisonchai, Enghoff & Panha gen. et sp. nov. from Tak Province. All new species are endemic to western Thailand and all are restricted to limestone habitats. Complete illustrations of external morphological characters, an identification key, and a distribution map are provided.
The bathyal kinorhynch fauna along the Northwest American continental rise is explored, with emphasis on species of Echinoderidae Zelinka, 1894. Seven species of Echinoderes Claparède, 1863 are described as new to science: E. anniae sp. nov., E. dubiosus sp. nov., E. hamiltonorum sp. nov., E. hviidarum sp. nov., E. juliae sp. nov., E. lupherorum sp. nov. and E. yamasakii sp. nov. Three known species, Echinoderes hakaiensis Herranz, Yangel & Leander, 2017, E. cf. unispinosus Yamasaki, Neuhaus & George, 2018 and Fissuroderes higginsi Neuhaus & Blasche, 2006, are reported. The numerous new species indicate that the deep-sea still holds a great, undiscovered diversity of kinorhynchs, and that Echinoderes, as is also the case in more shallow, coastal waters, represents an important component of the deep-sea kinorhynch fauna. The presence of E. hakaiensis in the deepsea sediments demonstrates that the species may occur at a great depth range, and suggests that depth may play a less important role for the distribution of some kinorhynch species. The finding of the Northeast Atlantic species E. cf. unispinosus and the Southwest Pacific species Fissuroderes higginsi could indicate that kinorhynch species in the deep-sea may cover considerably larger distributional ranges than is assumed for coastal species of Echinoderidae.
The family Plectopylidae is divided into two subfamilies: Sinicolinae subfam. nov. (included extant genera: Gudeodiscus Páll-Gergely, 2013, Endothyrella Zilch, 1959, Halongella Páll-Gergely, 2015, Sicradiscus Páll-Gergely, 2013, Sinicola Gude, 1899) and Plectopylinae Möllendorff, 1898 (included genera: Chersaecia Gude, 1899, Hunyadiscus Páll-Gergely, 2016, Naggsia Páll-Gergely & Muratov, 2016, Plectopylis Benson, 1860). The Eocene fossil Plectopyloides Yen, 1969 is classified into the Sinicolinae. The Plectopylinae are revised mainly based on historical type and non-type material, and the material of the Florida Museum of Natural History, collected in Thailand in the 1980s. The following species-group taxa are described as new: Chersaecia auffenbergi sp. nov., Chersaecia densegyrata sp. nov., Chersaecia mogokensis sp. nov., Chersaecia reversalis sp. nov., Chersaecia scabra sp. nov., Chersaecia shiroiensis subnagaensis subsp. nov., Hunyadiscus tigrina sp. nov., Naggsia oligogyra sp. nov., Plectopylis crassilabris sp. nov., Plectopylis malayana sp. nov. and Plectopylis thompsoni sp. nov. The genus Endoplon Gude, 1899 is treated as a synonym of Chersaecia. Consequently, the two species classified in Endoplon are members of Chersaecia: Chersaecia brachyplecta (Benson, 1863) comb. nov. and Chersaecia smithiana (Gude, 1897) comb. nov. The genus Plectopylis is redefined, and includes only species with fused anterior and posterior lamellae. Thus, the following species are moved from Plectopylis to Chersaecia: Chersaecia feddeni (Blanford, 1865) comb. nov., Chersaecia goniobathmos (Ehrmann, 1922) comb. nov., Chersaecia leucochila (Gude, 1897) comb. nov., Chersaecia magna (Gude, 1897) comb. nov. and Chersaecia woodthorpei (Gude, 1899) comb. nov. Altogether thirteen species and varieties are moved to the synonymy of valid species: Helix (Plectopylis) brachydiscus Godwin-Austen, 1879 syn. nov., Helix (Plectopylis) ponsonbyi Godwin-Austen, 1888 syn. nov., Plectopylis (Chersaecia) kengtungensis Gude, 1914 syn. nov., Plectopylis (Chersaecia) degerbolae Solem, 1966 syn. nov., Plectopylis lissochlamys Gude, 1897 syn. nov., Helix repercussa Gould, 1856 syn. nov., Plectopylis achatina var. obesa Gude, 1898 syn. nov., Plectopylis achatina var. infrafasciata Gude, 1898 syn. nov. Plectopylis achatina var. venusta Gude, 1898 syn. nov., Plectopylis achatina var. castanea Gude, 1898 syn. nov., Plectopylis achatina var. breviplica Gude, 1898 syn. nov., Plectopylis achatina var. repercussoides Gude, 1899 syn. nov., Plectopylis linterae var. fusca Gude, 1898 syn. nov. Plectopylis (Chersaecia) simplex Solem, 1966 is a subspecies of Chersaecia perarcta (Blanford, 1865), whereas Plectopylis muspratti Gude, 1897 is a subspecies of Chersaecia nagaensis (Godwin-Austen, 1875).
Ants of the Tetramorium solidum group occur in Africa, with the vast majority of species endemic to the arid regions of southern Africa. The first revision of the genus was published more than 30 years ago and ant surveys have since considerably expanded the number of specimens available for study. The revision of this group reveals five new species, expanding the total number to 19. Almost all the species in this group occur in the southern parts of the Afrotropical region, with the exception of T. setuliferum Emery, 1895 and T. rothschildi (Forel, 1907). These two species have broad distributions within African grasslands and savannas, with T. setuliferum occurring in southern Africa and T. rothschildi in East Africa and the Sahel. Five new species are described in this revision: T. aisha sp. nov., T. brigitteae sp. nov., T. duncani sp. nov., T. lerouxi sp. nov. and T. margueriteae sp. nov. An illustrated key is presented and descriptions of new species are provided, supported by montage images and distribution maps.
We describe two new frog species of the genus Guibemantis Dubois, 1992 (Mantellidae) from northern Madagascar. Both species are placed in the subgenus Pandanusicola Glaw & Vences, 1994 and, like most of their relatives, appear to only inhabit the leaf axils of Pandanus plants. Guibemantis albomaculatus sp. nov. is distinguished from other closely related species by light colored dorsolateral stripes and abundant small white spots found on its flanks and limbs. Guibemantis woosteri sp. nov. also has light-colored dorsolateral stripes and small white spots, but differs by having strongly banded forelimbs and a reddish coloration around the eye orbits. Mitochondrial DNA sequences corroborate the identity of these two new species, which appear to be regional endemics in northern and north-eastern Madagascar, respectively. These descriptions bring the species count in Pandanusicola to thirteen.
Four new species of Lobolibethra Hennemann & Conle, 2007 (Diapheromerinae: Diapheromerini: “Clonistria-group”) are described and illustrated. Lobolibethra carbonelli sp. nov. and L. pozuzoae sp. nov. from eastern Peru are described from both sexes and the eggs, L. tricarinata sp. nov. from southeast Ecuador is described from the female and male, and L. verruculosa sp. nov. from eastern Peru is described from the male. The previously unknown males of the Peruvian L. mainerii (Giglio-Tos, 1910) and L. mutica Hennemann & Conle, 2007 are described and illustrated. Dyme iconnicoffi Caudell, 1918 is shown to represent the previously unknown male of L. mainerii (Giglio-Tos, 1910) and is synonymised (syn. nov.). An updated key is presented for the eleven known species of Lobolibethra.
Flower flies (Diptera, Syrphidae) of French Polynesia, with the description of two new species
(2018)
The flower flies (Diptera, Syrphidae) of French Polynesia are revised. A total of nine syrphid species were recorded from the five archipelagos of French Polynesia. Among them are two species new to science, Allograpta jacqi Mengual & Ramage sp. nov. and Melanostoma polynesiotes Mengual & Ramage sp. nov., and a new record for this country, Syritta aenigmatopatria Hardy, 1964. We provide DNA barcodes for all flower fly species of French Polynesia, making the syrphid fauna of this country the first one in the world to be entirely barcoded. New data on biology, flowers visited and some taxonomic notes are provided. An identification key for the species of Syrphidae in French Polynesia is given, as well as an identification key for the species of Melanostoma Schiner, 1860 in the Australasian and Oceanian Regions.
Based on a study of 2150 specimens of the genus Trechus Clairville, 1806 from the Bale Mts, Mount Chillalo, and Mount Enkuolo, southern Ethiopia, the current knowledge regarding systematics and distribution of the genus in the area is summarized. Twenty-six new species are described and figured in detail: Trechus abalkhasimi sp. nov. (southeastern slope of Bale Mts, below Mt Abalkhasim); T. adaba sp. nov. (western Bale Mts, above Adaba); T. angavoensis sp. nov. (western Bale Mts, above Dodola); T. balesilvestris sp. nov. (western Bale Mts, above Adaba); T. bombi sp. nov. (southern escarpment of Bale Mts, Harenna Forest); T. colobus sp. nov. (western Bale Mts); T. depressipennis sp. nov. (northeastern margin of Sanetti Plateau, Bale Mts); T. dodola sp. nov. (western Bale Mts, above Dodola); T. fisehai sp. nov. (northern slope of Bale Mts, above Goba); T. grandipennis sp. nov. and T. hagenia sp. nov. (southern escarpment of Bale Mts, Harenna Forest); T. haggei sp. nov. (northeastern margin of Sanetti Plateau); T. harenna sp. nov. and T. harryi sp. nov. (southern escarpment of Bale Mts, Harenna Forest); T. iridescens sp. nov. (southeastern slope of Bale Mts, below Mt Abalk-hasim); T. mattisi sp. nov. (southern escarpment of Bale Mts, Harenna Forest); T. mekbibi sp. nov. (southern and western part of Bale Mts); T. minitrechus sp. nov. (northeastern slope of Mt Enkuolo); T. nanulus sp. nov. (southern escarpment of Bale Mts, Harenna Forest); T. nigrifemoralis sp. nov. and T. oppositus sp. nov. (western Bale Mts, above Dodola); T. rira sp. nov. (southern escarpment of Bale Mts, upper Rira Valley); T. sanettii sp. nov. (southeastern slope of Sanetti Plateau, Bale Mts); T. tragelaphus sp. nov. (western Bale Mts, above Dodola); T. transversicollis sp. nov. (southern escarpment of Bale Mts, Harenna Forest); T. wiersbowskyi sp. nov. (southern escarpment of Bale Mts, upper Rira Valley). New distributional data are provided for T. baleensis (Basilewsky, 1974), T. bastianinii Magrini & Sciaky, 2006, T. batuensis Magrini & Sciaky, 2006, T. chillalicus Jeannel, 1936, T. clarkeianus (Basilewsky, 1974), T. culminicola Jeannel, 1936, T. ericalis Magrini, Quéinnec & Vigna Taglianti, 2013, T. gallorites Jeannel, 1936, T. gypaeti Vigna Taglianti & Magrini, 2010, T. oromiensis Magrini, Quéinnec & Vigna Taglianti, 2012, T. relictus Magrini, Quéinnec & Vigna Taglianti, 2012 and T. rotundicollis (Basilewsky, 1974). Cothresia robini Basilewsky, 1974 is considered a junior synonym of T. chillalicus Jeannel, 1936. A key to all species known to occur in the Bale Mts and adjacent volcanos is presented.
Twenty-two new species of the genus Eviulisoma Silvestri, 1910, from the Eastern Arc Mountains, Tanzania, are described: E. acaciae sp. nov., E. aequilobatum sp. nov., E. akkariae sp. nov., E. angulatum sp. nov., E. articulatum sp. nov., E. biquintum sp. nov., E. breviscutum sp. nov., E. cetafi sp. nov., E. chitense sp. nov., E. commelina sp. nov., E. coxale sp. nov., E. ejti sp. nov., E. grumslingslak sp. nov., E. kalimbasiense sp. nov., E. navuncus sp. nov., E. nessiteras sp. nov., E. ottokrausi sp. nov., E. paradisiacum sp. nov., E. sternale sp. nov. and E. zebra sp. nov. from the Udzungwa Mts, E. culter sp. nov. from the Rubeho Mts and E. kangense sp. nov. from the Kanga Mts. Eviulisoma kwabuniense Kraus, 1958, and E. dabagaense Kraus, 1958, both from the Udzungwa Mts, are redesribed based on new material. Notes are provided on E. iuloideum (Verhoeff, 1941) based on type material. Eoseviulisoma Brolemann, 1920, is synonymized under Eviulisoma, based on newly collected material of E. julinum (Attems, 1909), type species of Eoseviulisoma. New material of Suohelisoma ulugurense Hoffman, 1964, type species of Suohelisoma Hoffman, 1964, has revealed that the gonopod structure is more similar to that of Eviulisoma than originally thought, but Suohelisoma is retained as a valid genus. Four species groups are recognized among Eviulisoma species from the Udzungwa Mts, but the need for a revision of the entire genus is emphasized. Two types of epizootic fungi are recorded from Eviulisoma spp., and an enigmatic amorphous mass, which may be a kind of plugging substance, is recorded from the gonopod tips and excavated sixth sternum of several species.
Representatives of the subgenus Helochares (s. str.) Mulsant, 1844 of China are revised. One new species, H. (s. str.) songi sp. nov., is described from Guangxi, China. All species known from China are redescribed. A diagnosis and a differential diagnosis are provided for each species. Helochares
fuliginosus d’Orchymont, 1932 is recorded for the first time from China and Cambodia. Additional distribution records of H. atropiceus Régimbart, 1903 and H. pallens (MacLeay, 1825) are provided from China. The habitus and aedeagus of all species are illustrated, and a key for the identification of Chinese species of the subgenus is provided.
Description of Hemicordulia tuiwawai sp. nov. from Kadavu Island, Fiji (Odonata: Corduliidae)
(2019)
Hemicordulia tuiwawai sp. nov. (Odonata: Corduliidae) is described and diagnosed based on material collected from Kadavu Island, Fiji; holotype: Wainitayuki River about 750 m above Baidamudamu village, -19.0916, 178.1038; 37 m a.s.l., 06 June 2016, M. Marinov leg. This species is distinguished from its congeners in the field by the contrasting colouration – dark green metallic body with bright yellow spots on the synthorax and base of the abdomen. This pattern is comparable to H. pacifica Selys, 1871. However, Hemicordulia tuiwawai sp. nov. can be recognised by the larger size and unique shape of the caudal appendages and genital hamule (in males) and vulvar scale (in females).
The South American genus Phantasca Redtenbacher, 1906 (Phasmatodea: Diapheromeridae: Diapheromaerinae) is re-diagnosed and revised at the species level. The precedingly unknown eggs are described for the first time. The genus Pterolibethra Günther, 1940 (type species: P. heteronemia Günther, 1940) is re-synonymised, with Phantasca (syn. nov.) and consequently the two species originally contained, P. heteronemia Günther, 1940 and P. poeciloptera Günther, 1940, are transferred to Phantasca (comb. rev.). P. laeta Conle, Hennemann & Gutierréz, 2011 is not congeneric and is transferred to the genus Jeremiodes Hennemann & Conle, 2007 (Cladomorphinae: Cladomorphini; comb. nov.). Two species are removed from Bacteria Berthold, 1827 and transferred to Phantasca; these are B. quadrilobata Chopard, 1911 and B. montana Redtenbacher, 1906 (comb. nov.). Six new species are described: P. adiposa sp. nov., P. amabile sp. nov., P. femorata sp. nov., P. guianensis sp. nov., P. nigrolineata sp. nov. and P. ruboligata sp. nov. The male and egg of P. quadrilobata (Chopard, 1911) are described and illustrated for the first time. The genus now contains 13 species that are distributed throughout the northern half of South America. A key as well as detailed descriptions and illustrations are presented for all known species.
Palaeosynthemis opaca sp. nov., a new dragonfly from Papua New Guinea (Anisoptera: Synthemistidae)
(2016)
A new species of the synthemistid genus Palaeosynthemis is described from the northern slopes of Papua New Guinea’s central cordillera. It is distinguished from all congeners by having females with large, nearly black patches at the bases of both wings. The male is most similar to P. cervula and P. feronia, but it differs from those species in having superior anal appendages less than three times as long as S10, basally enlarged and otherwise unarmed vs basally not enlarged (feronia) and almost four times as long as S10 and armed (cervula). Characters of the adults (both sexes) are illustrated and the affinities of the species are discussed.
Two new Cordulegaster species are described and illustrated by drawings, photographs and SEM images. The first one, C. plagionyx sp. nov., was discovered in NW Azerbaijan in a low-mountain forest landscape; the second, C. nachitschevanica sp. nov., occurs in the subalpine zone of Nakhichevan AR, south of the main territory of Azerbaijan. Both new taxa look generally similar to C. insignis Schneider, 1845; however, each of them reveals unique features and distinctive combinations of characters that set them apart from other species of the genus. Both new species exhibits new types of sex dimorphism previously unknown in Cordulegaster. Some traits related to the structure of male appendages and important details of colouration prevent both C. plagionyx sp. nov. and C. nachitschevanica sp. nov., from being reliably classified under any of two widely accepted groups of species within the genus Cordulegaster: the boltonii-group and the bidentata-group.
Drepanosticta kosterini sp. nov. (holotype ♂, from Gunung Penrissen, Kuching Division, Sarawak, Malaysian Borneo, deposited in RMNH) is described from both sexes. It is the sister species of D. actaeon Laidlaw, 1934; a fresh description of the male of D. actaeon and the first description of the female are given, along with discussion of variation in this species. Both D. actaeon and D. kosterini are considered to belong to a species group also including D. rufostigma (Selys, 1886) and a preliminary discussion of variation in this species is given, along with illustrations of both sexes. A neighbour joining COI gene tree for D. actaeon and D. kosterini is presented. The relationships of D. actaeon, D. kosterini and D. rufostigma to other members of the Platystictidae are briefly discussed.
A new species of the genus Teinobasis Kirby is described from the Muller Range in Western Province, Papua New Guinea. Its male is distinguished from all other Teinobasis species by having a pale labrum, an extensively bright orange thorax, and ventrally bowed superior anal appendages that are markedly shorter than the plump, apically rounded inferiors. Characters of the male are illustrated, and the affinities of the new species are discussed.
Two new species of damselflies are described from central Halmahera in North Maluku Province, Indonesia. They are Drepanosticta pararudicula sp. nov. (Holotype MZB. ODON. 19257) and Nososticta halmahera sp. nov. (Holotype MZB. ODON. 19265). The two species are most similar to the Moluccan taxa D. rudicula and N. moluccensis respectively and their descriptions bring the total number of Drepanosticta species known from Halmahera to five and of Nososticta to two.
Specimens of Neurothemis disparilis Kirby, 1889, N. fluctuans (Fabricius, 1793), N. fulvia (Drury, 1773), N. ramburii (Brauer, 1866), N. stigmatizans (Fabricius, 1775) and N. terminata Ris, 1911, including their subspecies, were studied with the main focus on the morphology of the vesica spermalis, wing maculation, wing venation, abdominal markings and vulvar scales. The results were compared with species descriptions and directly with type specimens where possible. The vesica spermalis, especially the medial process, is useful at least in separating species groups and supports the traditional differentiation methods using wing maculation and venation. The use of other characters in accessing specific status, coupled with known distribution patterns, is discussed. The following taxonomic changes are proposed: Neurothemis manadensis (Boisduval, 1835) stat. nov., Neurothemis papuensis (Lieftinck, 1942) stat. nov. and Neurothemis taiwanensis sp. nov. is described (27.5.1998, Kenting, Pingtung County/Taiwan, L. M. Juang leg.; holotype is deposited at Taiwan Forestry Research Institute, Taipeh, Taiwan). The type of Polyneura palliata Rambur, 1842 was rediscovered at MNHN and designated as lectotype; a lectotype for Neurothemis nicobarica Brauer, 1867 housed at NHMW is designated. The holotype of Neurothemis incerta Brauer, 1867 was rediscovered and synonymized with N. ramburii.
The genus Pseudolatirus Bellardi, 1884, with the Miocene type species Fusus bilineatus Hörnes, 1853, has been used for 13 Miocene to Early Pleistocene fossil species and eight Recent species and has traditionally been placed in the fasciolariid subfamily Peristerniinae Tryon, 1880. Although the fossil species are apparently peristerniines, the Recent species were in their majority suspected to be most closely related to Granulifusus Kuroda & Habe, 1954 in the subfamily Fusininae Wrigley, 1927. Their close affinity was confirmed by the molecular phylogenetic analysis of Couto et al. (2016). In the molecular phylogenetic section we present a more detailed analysis of the relationships of 10 Recent Pseudolatirus-like species, erect two new fusinine genera, Okutanius gen. nov. (type species Fusolatirus kuroseanus Okutani, 1975) and Vermeijius gen. nov. (type species Pseudolatirus pallidus Kuroda & Habe, 1961). Five species are described as new for science, three of them are based on sequenced specimens (Granulifusus annae sp. nov., G. norfolkensis sp. nov., Okutanius ellenae gen. et sp. nov.) and two (G. tatianae sp. nov., G. guidoi sp. nov.) are attributed to Granulifusus on the basis of conchological similarities to sequenced species. New data on radular morphology is presented for examined species.
This paper presents an improved diagnosis and definition of the genus Stenaelurillus Simon, 1886, with new taxonomic and faunistic data for 23 species. The genera Microheros Wesołowska & Cumming, 1999 and Mashonarus Wesołowska & Cumming, 2002 are synonymized with Stenaelurillus. Six new species are described: Stenaelurillus bandama sp. nov. (♂♀, from Côte d’Ivoire), S. belihuloya sp. nov. (♂, from Sri Lanka), S. jocquei sp. nov. (♂♀, from Cameroon), S. pseudoguttatus sp. nov. (♂, from Namibia), S. senegalensis sp. nov. (♂♀, from Senegal), and Stenaelurillus siyamae sp. nov. (♀, from Sudan). Lectotypes are designated for two species: S. albopunctatus Caporiacco, 1949 (♂) from Kenya and S. werneri Simon, 1906 (♀) from South Sudan. Six new combinations are proposed: Aelurillus ambiguus (Denis, 1966), comb. nov. (ex Stenaelurillus); Evarcha werneri (Simon, 1906), comb. nov. (ex Stenaelurillus); Phlegra davidi (Caleb, Mungkung & Mathai, 2015), comb. nov. (ex Mashonarus); Stenaelurillus brandbergensis (Wesołowska, 2006), comb. nov. (ex Mashonarus); Stenaelurillus guttatus (Wesołowska & Cumming, 2002), comb. nov. (ex Mashonarus); and S. termitophagus (Wesołowska & Cumming, 1999), comb. nov. (ex Microheros). Two species names are synonymized: Evarcha elegans Wesołowska & Russell-Smith, 2000 with E. werneri comb. nov.; and Aelurillus sahariensis Berland & Millot, 1941 with Stenaelurillus nigricaudus Simon
The Siriella brevicaudata species group from the West Indo-Pacific, defined and designated by Murano & Fukuoka (2008), previously contained five nominal species. In this study we describe five new species in the brevicaudata group: S. bassi sp. nov. from the Bass Strait, southern Australia, S. occulta sp. nov. from the Arabian Gulf, S. muranoi sp. nov. from the coast of Northern Territory, Australia, S. tabaniocula sp. nov. from Ningaloo Reef of Western Australia and Lodestone Reef off Queensland, and S. talbotae sp. nov. from Lizard Island, Queensland, Australia. Furthermore, Siriella hanseni W.M. Tattersall, 1922 from India and S. vincenti W.M. Tattersall, 1927 from South Australia are redescribed based on re-examination of their type material. A re-examination of specimens subsequently attributed to these two species from other geographical regions showed that these were misidentifications, partly representing three of the new species described herein. Siriella gibbosa (Ledoyer, 1970), which was previously synonymized with S. brevicaudata Paulson, 1875 by Bačescu, is revalidated and included within the brevicaudata group. Siriella lacertilis Talbot, 2009, from Lizard Island, is placed within the brevicaudata group. Diagnostic features for all the members of the group and the group itself are updated. As a result of the present study, the brevicaudata group now comprises 12 valid species.
Homollea Arènes (Rubiaceae, subfamily Ixoroideae, tribe Pavetteae) is a genus of shrubs and small trees endemic to western and northern Madagascar. The genus comprises five species occurring in dry deciduous forest, often in limestone areas. The five species are narrow endemics and their conservation status is either Endangered (4 species) or Critically Endangered (1 species). Homollea is characterized by few-flowered, pseudo-axillary, pedunculate inflorescences, well-developed calyces with the lobes much longer than the tube, laterally flattened seeds with a shallow, elongated to linear hilum and entire endosperm, ovules arising from the upper margin of the placenta, and, pollen grains with supratectal elements in the shape of microgemmae. Until now, three species were known and their descriptions are amended. Two further species, H. furtiva De Block sp. nov. and H. septentrionalis De Block sp. nov., are described as new for science. The five species are dealt with in detail: descriptions, distribution maps, conservation assessments, illustrations, lists of exsiccatae and an identification key are given.
A new monotypic genus of flatid planthoppers (Hemiptera: Fulgoromorpha: Flatidae), Medleria gen. nov., is described for Medleria caudata gen. et sp. nov. (type species) from the island of Socotra (Yemen). Habitus, male and female external and internal genital structures of the new species are illustrated and compared with similar taxa. Medleria caudata gen. et sp. nov. is probably endemic to Socotra where it is known to date from a small area of the Dixam mountain plateau only.
The concept of the jumping spider genus Pochytoides Berland & Millot, 1941 is reviewed, based on the examination of described and undescribed species. Pochytoides is elevated from the subgeneric to the generic rank and a short diagnosis and description of the genus are presented. Redescriptions or descriptions of all species are provided together with a key to the species. Two new combinations are proposed: Pochytoides perezi (Berland & Millot, 1941) comb. nov. and P. poissoni (Berland & Millot, 1941) comb. nov. (both from Pochyta). Pochyta remyi Berland & Millot, 1941 originally placed in the subgenus Pochytoides is excluded; new combination Thiratoscirtus remyi (Berland & Millot, 1941) comb. nov. is proposed for it (but its generic status is uncertain). Six new species are described: Pochytoides monticola sp. nov., P. obstipa sp. nov., P. lamottei sp. nov., P. patellaris sp. nov., P. securis sp. nov. and P. spiniger sp. nov. The genus has a West African distribution.
The Palaearctic species of Ismaridae Thomson, 1858 are reviewed. Thirteen species of Ismaridae are recognized from the Palaearctic. Five species are described as new: Ismarus brevis Kim & Lee sp. nov. from the Russian Far East and South Korea; I. distinctus Kim, Notton & Ødegaard sp. nov. from Norway and the United Kingdom; I. excavatus Kim & Lee sp. nov. from China, Japan and South Korea; I. similis Kim, Notton & Lee sp. nov. from the United Kingdom and I. tripotini Kim & Lee sp. nov. from South Korea. Ismarus apicalis Kolyada & Chemyreva, 2016 is newly recorded from China, France, Japan and South Korea; I. dorsiger (Haliday, 1831) from France, Montenegro, Norway, South Korea and Switzerland; I. flavicornis (Thomson, 1858) from Bulgaria and Norway; I. grandis Alekseev, 1978, I. halidayi Förster, 1850 and I. multiporus Kolyada & Chemyreva from Japan and South Korea; I. rugulosus Förster, 1850 from Austria and I. spinalis Kolyada & Chemyreva, 2016 from China, Japan and South Korea. An identification key to all species found in the Palaearctic region is presented.
Two new species of the family Selachinematidae Cobb, 1915 from an intertidal sandy flat in the East China Sea are described. Both species are distinct rapacious nematodes, preying on other nematodes. Gammanema magnum sp. nov. is characterized by its leaf-like labial setae, large loop-shaped amphideal fovea in the male and small spiral ones in the female, an anterior buccal cavity reinforced by 12 rhabdia, each rhabdion with about six denticles, and two unequally long spicules. Gammanema magnum sp. nov. is a striking species, exhibiting extreme sexual dimorphism in the amphideal fovea, but lacking a gubernaculum or precloacal supplements, all features by which it can be differentiated from congeneric species. Synonchium caudatubatum sp. nov. is characterized by a posterior buccal cavity, with three mandibles each with a large tooth flanked by three smaller teeth on both sides, a transverse oval-shaped multispiral amphideal fovea, and a bluntly rounded tail with a protuberant caudal duct. Synonchium caudatubatum sp. nov. is unique in the genus in having a protuberant caudal duct in the tail end. Furthermore, it differs from its congeners by its mandibles, which have seven teeth each. Synonchium caudatubatum sp. nov. lacks a gubernaculum and a precloacal supplement, both features present in congeners except S. depressum Gerlach, 1954.
A new species complex, the eparmata complex, is established within the subgenus Phortica s. str., based on eight known and five new species, all of which are endemic to the Oriental Region: P. bipartita (Toda & Peng, 1992), P. eparmata (Okada, 1977), P. lanuginosa Chen & Toda, 2007, P. latipenis Chen & Gao, 2005, P. pangi Chen & Wen, 2005, P. setitabula Chen & Gao, 2005, P. unipetala Chen & Wen, 2005 and P. zeta Chen & Toda, 2007; P. jadete sp. nov., P. kava sp. nov., P. mengda sp. nov., P. wongding sp. nov. and P. yena sp. nov. A key to all species of this complex is provided. Barcoding sequences (mitochondrial COI gene) were obtained for 22 specimens of five known and the five abovementioned new species. The intra- and inter-specific pairwise K-2P (Kimura’s two-parameter) distances of COI were determined. Phylogenetic analysis was performed using Bayesian inference based on COI sequences, confirming the monophyletic status of the eparmata complex, which is distinct from the species complexes of magna, omega, variegata and another two ungrouped species.
The Quedius mutilatus group, a very poorly known presumably monophyletic complex of wingless, possibly hypogean species confined to the Tien-Shan Mountains, is characterized as such for the first time. Newly available material clarified the identity of Q. mutilatus Eppelsheim, 1888 and Q. kalabi Smetana, 1995, each hitherto known from a handful of non-conspecific and vaguely georeferenced specimens only. Additional material is reported for Q. equus Smetana, 2014 and bionomics for all these four species of the group are summarized.
Bees of the genus Lasioglossum (Hymenoptera: Halictidae) from Greater Puerto Rico, West Indies
(2018)
The species of Lasioglossum from Greater Puerto Rico are reviewed. Nine species are recognized, including five new species described herein: asioglossum (Dialictus) genaroi sp. nov., L. (D.) dispersum sp. nov., L. (D.) enatum sp. nov., L. (D.) monense sp. nov. and L. (D.) amona sp. nov. The latter two are known only from Mona Island. Keys and images are provided to assist in identification. Details of nesting biology, floral hosts and distribution are provided where available. Three species, L. (D.) parvum (Cresson, 1865), L. (D.) busckiellum (Cockerell, 1915), and L. (D.) mestrei (Baker, 1906) are removed from the list of species for Puerto Rico. Details on their revised distribution are provided. Three new records for Haiti, L. (D.) gundlachii (Baker, 1906), L. (D.) ferrerii (Baker, 1906) and L. (D.) busckiellum are documented. Notes on other species in the Greater Antilles are provided, including the synonymy of Lasioglossum bruesi (Cockerell, 1912) and L. jamaicae (Ellis, 1914) under L. gemmatum (Smith, 1853).
The genus Raveniola Zonstein, 1987 is found to be represented in Western Asia by 16 species: ♂♀ R. adjarica sp. nov. (Georgia), ♂ R. anadolu sp. nov. (Turkey), ♂ R. arthuri Kunt & Yağmur, 2010 (Turkey), ♂ R. birecikensis sp. nov. (Turkey), ♂♀ R. dunini sp. nov. (Armenia, Azerbaijan, Iran), ♂♀ R. hyrcanica Dunin, 1988 (Azerbaijan), ♂ R. marusiki sp. nov. (Iran), ♂ R. mazandaranica Marusik, Zamani & Mirshamsi, 2014 (Iran), ♂♀ R. micropa (Ausserer, 1871) (Turkey), ♀ R. nana sp. nov. (Turkey), ♂♀ R. niedermeyeri (Brignoli, 1972) (Iran), ♂♀ R. pontica (Spassky, 1937) (Russia, Georgia), ♀ R. sinani sp. nov. (Turkey), ♂♀ R. turcica sp. nov. (Turkey), ♂♀ R. vonwicki Zonstein, 2000 (Iran) and ♂♀ R. zaitzevi (Charitonov, 1948) (Azerbaijan, Georgia) = ♀ Brachythele recki Mcheidze, 1983, syn. nov. Eight species are described as new; others are redescribed from types and/or conspecific material. Males of R. micropa and R. zaitzevi, hitherto unknown, are described for the first time. Data on the variability, relationships, distribution and ecology of all considered species are also provided.
The species of the subgenus Conocetus Desbrochers des Loges, 1875 are reviewed and Polydrusus (Conocetus) transjordanus sp. nov. is described. Upon examination of the holotype of Polydrusus bardus Gyllenhal, 1834, it was observed that the species hitherto determined sensu auctorum as P. bardus was a misidentification. The specimen in question was therefore unnamed and is thus newly described as Polydrusus (Conocetus) crinipes sp. nov. Polydrusus femoratus (Stierlin, 1888) is a junior synonym of P. angustus (Lucas, 1854). Polydrusus gracilicornis Kiesenwetter, 1864, P. cylindrithorax (Desbrochers des Loges, 1900) and P. quadraticollis (Desbrochers des Loges, 1902) are proposed as junior synonyms of P. bardus. Polydrusus zurcheri (Schilsky, 1912) is proposed as a junior synonym of P. grandiceps (Desbrochers des Loges, 1875). Polydrusus kahri Kirsch, 1865 is transferred from subgenus Conocetus to Denticonocetus subgen. nov., with P. siculus Desbrochers des Loges, 1872 and P. vodozi Desbrochers des Loges, 1903 both recognized as new junior synonyms of P. kahri. The lectotypes of P. gracilicornis, P. zurcheri, P. marcidus Kiesenwetter, 1864, P. gracilis (Stierlin, 1888), P. rhodiacus (Schilsky, 1912) and P. grandiceps are designated. A key, figures, label data and distribution maps are provided for all species, except for P. longus (Stierlin, 1884), for which no specimens were available for examination, and whose placement in the subgenus Conocetus remains uncertain (thus categorized as incertae sedis). Polydrusus angustus is recorded for the first time for Italy, P. rhodiacus for mainland Turkey and P. festae (Solari, 1925) for Greece.
Five species of the terrestrial diatom genus Luticola D.G.Mann were found during a taxonomic survey of two small volcanic islands, Ile Amsterdam and Ile Saint-Paul (Southern Indian Ocean). Apart from the two already known Luticola species L. beyensii Van de Vijver et al. and L. subcrozetensis Van de Vijver et al., two new species are described: L. ivetana Chattová & Van de Vijver sp. nov. and L. vancampiana Chattová & Van de Vijver sp. nov. Finally, one, up to now unknown, Luticola species is briefly discussed and illustrated. Detailed morphological descriptions of these taxa are provided based on both light and scanning electron microscopy observations. Morphological features of the new species are compared to morphologically similar taxa, and notes on their ecology and biogeography are added.
The genus Synaldis Foerster, 1863 is recorded in the Neotropical region for the first time. Five new Neotropical species, S. brasiliense Peris-Felipo, sp. nov., S. fritzi Peris-Felipo, sp. nov., S. longiflagellaris Peris-Felipo, sp. nov., S. magnioculis Peris-Felipo, sp. nov., and S. novateutoniae Peris-Felipo, sp. nov., are described and illustrated. The original combination for Synaldis ulmicola Ashmead, 188
Siamopsis gen. nov., described here, belongs to a group of genera with the right valve overlapping the left valve in the subfamily Cypridopsinae Kaufmann, 1900 of the family Cyprididae Baird, 1845. The distinguishing characters of the new genus are in the morphology of its valves and soft parts. The postero-dorsal margin of the internal left valve is plate-like protruded. The morphology of this plate varies in different species, e.g., some species bear a tooth-like tubercle on the plate. The posterior margin of the right valve is recurved inwardly at ca mid-height, resulting in the occurrence of a lobe-like expansion that can clearly be seen in the dorsal and caudal views of the carapace. In addition, the other diagnostic soft part features of the new genus are the cylindrical caudal ramus, the presence of two t-setae on the female A2 penultimate segment, the very elongated terminal segment of the Mx1 palp, the morphology of the two large bristles (tooth bristles) of the Mx1 third endite (one smooth, one serrated) and the absence of d-seta on T1. In the present paper, five new species are described under this new genus: Siamopsis renateae gen. et sp. nov., S. suttajiti gen. et sp. nov., S. conspecta gen. et sp. nov., S. khoratensis gen. et sp. nov. and Siamopsis planitia gen. et sp. nov. A key to the species of Siamopsis gen. nov. is also provided.
Two species of the nematode family Diplopeltidae are described from Skagerrak. The new genus Belgopeltula gen. nov. is proposed for Diplopeltula belgica Vincx & Gourbault, 1992 and is characterised by: amphidial fovea circular in female and double-loop-shaped in male; excretory pore located at the level of cephalic setae bases; oral opening on the dorsal side of the body; pharynx subdivided into strongly muscularised fusiform corpus and weakly muscularised narrow and long postcorpus; female didelphic with antidromously reflexed ovaries; supplements absent. Mudwigglus micramphidium sp. nov. is characterised by: a body of 0.6 mm long; cephalic sensilla 1.5 μm long; amphidial fovea loop-shaped, 8 μm long and 3.5 μm wide; gymnostom without cuticularised ring; tail elongate conoid, with subcylindrical distal part; terminal setae absent; spicules 15 μm long; gubernaculum present; two midventral precloacal setae. It is distinguished from M. macramphidium Leduc, 2013 in having shorter amphidial fovea, shorter spicules and presence of two precloacal setae. Redescription of Diplopeltis cylindricauda Allgén, 1932 is provided based on type material. Diplopeltula minuta Vitiello, 1972 is transferred to the genus Mudwigglus Leduc, 2013. Diplopeltis cylindricauda Allgén, 1932, Diplopeltula laminata Vitiello, 1972 and Diplopeltula cassidaignensis Vitiello, 1972 are transferred to the genus Pseudaraeolaimus Chitwood, 1951.
We review the three species currently placed in the genus Xylopertha Guérin-Méneville, 1845, and describe a new species, Xylopertha elegans sp. nov., from Turkey. We propose the following new synonymy: Xylopertha Guérin-Méneville, 1845 (= Paraxylogenes Damoiseau, 1968); Xylopertha reflexicauda (Lesne, 1937) (= Paraxylogenes pistaciae Damoiseau, 1968). We give details of the sexual dimorphism, and summarise information on the distribution and biology of all species. A key to the species of Xylopertha is provided.
Two Mastogloia Thwaites ex W.Sm. taxa were found during a survey of the diatom flora of Lac de Guiers, Senegal. Based on all currently available literature, one taxon could be identified as M. belaensis M.Voigt, a species formerly described from Pakistan. The second species showed some resemblance to M. braunii Grunow. Analysis of the type of M. braunii revealed, however, important morphologic differences, leading to the description of a new species from the Senegal population: M. senegalensis Van de Vijver, Fofana, Sow & Ector sp. nov. The present paper describes this new species and discusses and illustrates the morphology of M. belaensis and the type of M. braunii. All taxa are discussed with morphologically similar taxa.
Three new scale insect species, Coccidohystrix daedalea Gavrilov-Zimin sp. nov., Mirococcopsis ptilura Gavrilov-Zimin sp. nov. (both from the family Pseudococcidae) and Cryptinglisia millari Gavrilov-Zimin sp. nov. (family Coccidae), are described and illustrated from the Western Cape Province of South Africa.
Recent expeditions (NANHAI 2014, DONGSHA 2014 and ZHONGSHA 2015) conducted in deep waters of the South China Sea obtained interesting material of various spider crabs (Majoidea) including several new records for the area, and two new species of epialtids of the genera Oxypleurodon Miers, 1885 and Stegopleurodon Richer de Forges & Ng, 2009. Two poorly known species, previously only known from their types, Rochinia strangeri Serène & Lohavanijaya, 1973 and R. kagoshimensis (Rathbun, 1932) comb. nov., are redescribed, refigured, and their taxonomy discussed.
Revision of the genus Spilopteron Townes, 1965 (Hymenoptera: Ichneumonidae: Acaenitinae) from Japan
(2017)
Ten Japanese species of the genus Spilopteron Townes, 1965 are recognized. Five new species, S. albiventre sp. nov., S. brachyurum sp. nov., S. nigrum sp. nov., S. oblongulum sp. nov. and S. pseudonigrum sp. nov., are described from Japan. Morphological discrimination between most Japanese species is confirmed by sequence analysis of the mitochondrial COI gene, which indicates the following relationships: S. oblongulum sp. nov. + S. apicale (Matsumura, 1912), S. brachyurum sp. nov. + S. nigrum sp. nov. + S. pseudonigrum sp. nov., and S. tosaense (Uchida, 1934) + S. luteum (Uchida, 1930). A key to the Japanese species of Spilopteron is provided. This genus seems to have its center of diversity in the mid-latitude area of East-Asia.
Haplosclerid sponges possessing a unique asymmetric flagelliform type of sigmoid microsclere have been reported from all global oceans. This peculiar spicule, characterized by a circular or elliptical shape, with a longer and sharper curved ending at one side and a shorter and more gradually curved ending at the opposing side, is proposed to be termed ‘flagellosigma’. These sponges invariably also possess smaller normal sigmas while their skeletal structure of oxea megascleres is markedly confused. They are assigned to the large genus Haliclona Grant, 1841 (family Chalinidae) in a new subgenus, Haliclona (Flagellia) subgen. nov. The species belonging to the new subgenus are reviewed and four species new to science are described, Haliclona (Flagellia) indonesiae subgen. et sp. nov., H. (F.) amirantensis subgen. et sp. nov., H. (F.) hiberniae subgen. et sp. nov. and H. (F.) hajdui subgen. et sp. nov. One species, H. (F.) hentscheli nom. nov., is given a new name on account of secondary homonymy caused by its transfer to the genus Haliclona. One species remains unnamed because of paucity of material. Already known species, reassigned to the new subgenus are H. (F.) hamata subgen. et comb. nov., H. (F.) flagellifera subgen. et comb. nov., H. (F.) porosa subgen. et comb. nov., H. (F.) edaphus subgen. et comb. nov. and H. (F.) anataria subgen. et comb. nov. Additional species are likely hiding among many erroneous records of ‘Gellius flagellifer’ from wide ranging parts of the global oceans.
Big and beautiful: the Megaxyela species (Hymenoptera, Xyelidae) of East Asia and North America
(2017)
Megaxyela Ashmead, 1898 comprises 13 species, four of which are described as new and one is removed from synonymy: Megaxyela euchroma Blank, Shinohara & Wei sp. nov. from China (Zheijang), M. fulvago Blank, Shinohara & Wei sp. nov. from China (Hunan, Jiangsu, Zhejiang), M. inversa Blank & D.R. Smith sp. nov. from the USA (West Virginia), M. langstoni Ross, 1936 sp. rev. from the eastern USA, and M. pulchra Blank, Shinohara & Sundukov sp. nov. from China (Hubei, Jilin, Liaoning, Shaanxi, Tibet), South Korea (Kangwon-do) and Russia (Primorskiy Kray). The male of M. parki Shinohara, 1992 is described for the first time. A lectotype is designated for M. gigantea Mocsáry, 1909. A cladogram, based on COI sequences of seven species, is presented and interpreted in view of selected morphological characters. Records of M. fulvago sp. nov. from Hunan and of M. pulchra sp. nov. from Tibet extend the known distribution of Megaxyela in the Old World 600 kilometers farther south and 2500 kilometers farther west than previous records.
The genus Hybovalgus Kolbe, 1904 is represented by eight species on the Chinese mainland, many of which also inhabit northern Vietnam and Laos. Species of Hybovalgus are endemic to this area, and to the island of Taiwan. Until now, there is a lot of confusion in our knowledge of Hybovalgus on mainland China, due to erroneous descriptions of new species by European entomologists and incorrect identifications of specimens by local entomologists. Study of more material and many types has clarified this situation by better defining the species, synonymizing some of them, describing one new species, Hybovalgus calvus sp. nov. and recognizing the fact that females of two species were included in the new genus Excisivalgus Endrödi, 1952, which is here synonymized with Hybovalgus.
Japanese species of the genus Intybia are revised taxonomically, with the examination of the endophallic structure. Eight species, including one new species Intybia donan sp. nov. from Yonagunijima, are recognized. All species are described or redescribed with a key and figures. The endophallic structure contains one primary sclerite (gonoporal piece), three secondary sclerites (ligula, semigonoporal piece, and spinous plate) in some species, and a membranous basal area densely covered with many spines (spinous area). Based on the structures of the endophallus, the Japanese members of the genus are divided into two species groups (the histrio and pelegrini groups). The pelegrini species group is furthermore subdivided into three subgroups (subgroups 1–3). New distributional records are as follows: I. histrio from Hachijô-jima and Tanega-shima; I. niponica from Sakhalin and I. takaraensis from Tokuno-shima and Amami-Ôshima.
From a moss sample collected in the Manabí Province in Ecuador, we extracted 96 specimens of a new species of eutardigrade. No eggs were found. In order to obtain eggs, an in vitro culture was prepared. In total, 136 specimens (including ten simplex), one exuvia and 44 eggs (including two with embryos) of the new species were obtained. In addition to the traditional taxonomic description with morphometrics, light and scanning microscopy imaging, we also provide nucleotide sequences of three nuclear (18S rRNA, 28S rRNA, ITS-2) and one mitochondrial (COI) DNA fragments of the new species. Macrobiotus polypiformis sp. nov. belongs to the hufelandi group and is most similar to Ma. paulinae Stec, Smolak, Kaczmarek & Michalczyk, 2015, but differs from it mainly by the lack of dorso-lateral patches of granulation on the cuticle, egg processes with longer and more numerous filaments and in some morphometric characters of both eggs and adults. Moreover, we provide a short discussion on the modifications/abnormalities of the claws in eutardigrades and the possible consequences on the taxonomic status of Mesobiotus armatus (Pilato & Binda, 1996), suggesting its consideration as species inquirenda (with uncertain taxonomic status).
The genus Koiulus gen. nov. and its type-species, Koiulus interruptus gen. et sp. nov., are described from the Russian Far East. The new genus is compared with other genera of Mongoliulidae, in particular with Ussuriiulus Golovatch, 1980, also from the Russian Far East, with which it shares the absence of ozopores from individual body rings distributed along the body, a condition so far otherwise unknown in the superorder Juliformia. A synoptic table of genera and a list of species of Mongoliulidae are presented.
We describe a new minute species of the genus Pristimantis, P. boucephalus sp. nov., from the Yanachaga-Chemillén National Park, Región Pasco, Peru. The description is based on a freshly collected male specimen found at 2950 m a.s.l. in a cloud forest and four previously unidentified museum specimens consisting of two adult males, one subadult female and a juvenile from the Yanachaga-Chemillén National Park. The new species is mainly characterized by a snout–vent length of 13.4–14.5 mm in adult males (n = 3), and 12.5 mm in the only known subadult female, and is compared morphologically and genetically with other taxonomically and biogeographically relevant species of Pristimantis. The new species is characterized by its small size, disproportionally large head with short snout, absence of a tympanic annulus and membrane, and reddish-copper iris. Phylogenetically it belongs to a speciose clade, an as yet unnamed species group, comprising both montane (Andes, Guiana Shield) and lowland (Amazon) taxa from the northern part of South America. The new species is genetically close to the sympatric P. cruciocularis. Species of Pristimantis occurring in the Cordillera Yanachaga region in the Andes of central Peru are members of six divergent phylogenetic lineages.
Examination of leucothoid amphipods of the Red Sea has revealed seven species not previously reported from this location. Leucothoe minoculis sp. nov., Leucothoe pansa sp. nov., Leucothoe reimeri sp. nov., and Paranamixis sommelieri sp. nov. are described and the range of Leucothoe predenticulata Ledoyer, 1978, L. acutilobata Ledoyer, 1978 and L. squalidens Ledoyer, 1978 is expanded to include the Red Sea. Clarification of reports of L. acanthopus Schellenberg, 1928 and L. bannwarthi (Schellenberg, 1928) is provided and Leucothoe alani sp. nov. is described from outside the Red Sea.
The Chimarra lehibemavo species-group, new and endemic to Madagascar (Trichoptera, Philopotamidae)
(2017)
The Chimarra lehibemavo group is described to include thirteen new species: Chimarra lehibemavo sp. nov., C. cebegepi sp. nov., C. fenoevo sp. nov., C. forcellinii sp. nov., C. fotobohitra sp. nov., C. gattolliati sp. nov., C. gensonae sp. nov., C. jejyorum sp. nov., C. hamatra sp. nov., C. makiorum sp. nov., C. moramanga sp. nov., C. saha sp. nov. and C. tamara sp. nov. The adults are easily recognizable by their large size, yellow colour and the structure of the male genitalia. The membranous tergum IX and the absence of the mesal lobe of tergum X are observed in other lineages, but the strong asymmetrical deformation of the phallotheca is apomorphic. The group is monophyletic with unknown affinities, but a preliminary phylogenetic placement is suggested following genetic analysis of two specimens. With one exception, the species have restricted geographical distributions in Madagascar and inhabit rivers in eastern pristine rainforests.