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The “trachystreptoform” species of Spirostreptidae, i.e., species which would formerly have been ascribed to the tribe Trachystreptini, from the Udzungwa Mountains are (re)described, including one new genus and five new species: Attemsostreptus reflexus Akkari & Enghoff, 2019, A. cataractae Enghoff sp. nov., A. leptoptilos Enghoff sp. nov., A. julostriatus Enghoff sp. nov., Lophostreptus tersus (Cook, 1896) (= L. ptilostreptoides Carl, 1909 syn. nov.), L. magombera Enghoff sp. nov., and Udzungwastreptus marianae Enghoff gen. et sp. nov. The type material of Lophostreptus regularis Attems, 1909 (= L. tersus) is discussed. The discussion includes paragraphs on the classification and the Udzungwa fauna of Spirostreptidae, on grouping of the Udzungwa trachystreptoform species in relation to altitude, and on the possibly recent immigration of A. reflexus and L. tersus into the Udzungwa Mts.
Three new species of the family Pyrgodesmidae Silvestri, 1896 from the Udzungwa Mountains are described: Cryptocorypha geminiramus sp. nov., Cryptocorypha cactifer sp. nov., and Cryptocorypha exovo sp. nov. Several additional, unidentifiable pyrgodesmids from the Udzungwa Mountains are recorded. Elythesmus enghoffi Hoffman, 1978 (Cryptodesmidae Karsch, 1880), is recorded from the East Usambara (type locality), West Usambara, Uluguru, Nguru (Kanga) and Udzungwa Mountains, and its variation in size and body ring numbers is analyzed. The surface sculpture of pro- and metazonites is discussed.
The fauna of the millipede family Odontopygidae in the Eastern Arc Mountains of Tanzania is reviewed. Species from the North Pare, South Pare, West Usambara, East Usambara, Nguru, Rubeho, Uluguru and Rungwe Mts are treated. The odontopygids of the Udzungwa Mts have been subject of a series of previous papers and are only treated marginally. Six new genera and 25 new species are described: Antipustia gen. nov., Aptyctosmilax gen. nov., Multipronopea gen. nov., Notogallanus gen. nov., Praludivera gen. nov., Uncodrama gen. nov., Antipustia hoteldolichoiuli gen. et sp. nov., Aptyctosmilax helenae gen. et sp. nov., Calyptomastix ingemanni sp. nov., Calyptomastix vuasu sp. nov., Calyptomastix xystopygoides sp. nov., Calyptomastix zoltani sp. nov., Chaleponcus jolantae sp. nov., Chaleponcus nesrineae sp. nov., Chaleponcus schioetzae sp. nov., Chaleponcus sergeii sp. nov., Chaleponcus soerensenae sp. nov., Geotypodon cristinae sp. nov., Lamelloramus frederiksenae sp. nov., Multipronopea agneteae gen. et sp. nov., Notogallanus mastacembalus gen. et sp. nov., Praludivera paralellamella gen. et sp. nov., Raduliverpa donatellae sp. nov., Spinotarsus axeli sp. nov., Syndesmogenus estelleae sp. nov., Uncodrama coronata gen. et sp. nov., Xystopyge bentemarieae sp. nov., Xystopyge doggartae sp. nov., Xystopyge hippocampus sp. nov., Xystopyge minnae sp. nov., and Xystopyge voluntariorum sp. nov. The discussion focuses on diversity and distribution patterns, the justification for monotypic (monospecific) genera, and the following morphological character types: the ozopore series, the limbus, the number of setae on the anal valves, the first pair of male legs, the gonopod sternum, and the sternum of the rudimentary 9th leg-pair.
For each of the 534 species of the millipede order Chordeumatida known from Europe, available information on taxonomy, distribution and habitat is summarized, and the distribution in 50 × 50 km UTM/MGRS squares is shown on a map. Comparisons between Chordeumatida and the equally-sized order Julida are made with respect to distribution patterns and history of exploration.
Two new species of giant pill-millipedes, Zephronia viridisoma Rosenmejer & Wesener sp. nov. and Sphaerobelum aesculus Rosenmejer & Wesener sp. nov., are described based on museum samples from southern Thailand. Zephronia viridisoma sp. nov. comes from Khao Lak, while the type locality of S. aesculus sp. nov. is on Phuket Island. Both species are described integratively, combining light microscopy, scanning electron microscopy, multi-layer photography, micro-CT scans and genetic barcoding. Genetic barcoding was successfully conducted for holotypes of both new species, which could be added to a dataset of all published sequences of the family Zephroniidae, including all described species from Thailand, Laos and Cambodia up to 2020. Genetic barcoding of the COI gene revealed another female of S. aesculus sp. nov., 160 km east of the type locality. Both new species are genetically distant from all other Zephroniidae from Thailand and surrounding countries, showing uncorrected p-distances of 16.8–23.1%. A virtual cybertype of a paratype of Z. viridisoma sp. nov. was created and made publically accessible.
A new giant species of the subfamily Rhynchoproctinae with strikingly bi-coloured, red-yellow legs contrasting with a black body is described: Alienostreptus bicoloripes sp. nov. from Vietnam. The new species is assigned to the genus Alienostreptus Pimvichai, Enghoff & Panha, 2010 hitherto comprising one species, A. alienus (Attems, 1936), and differing from other rhynchoproctine genera by having the femoral spine duplicated. Species of this genus share three synapomorphic characters of the subfamily, viz (1) anterior coxal fold forms deep concavity in posterior view, (2) posterior coxal fold very low, and (3) posterior coxal fold with mesal flap. A superficially very similar colourful species from Borneo, also with bi-coloured legs but clearly not belonging to Alienostreptus due to the position of ventral soft pads on male legs, is documented based on photographs.
A new genus and six new species of the family Gomphodesmidae from the Udzungwa Mts are described, including Pogoro alopias Rosenmejer & Enghoff sp. nov., Pogoro siren Rosenmejer & Enghoff sp. nov., Pogoropsis prolixopes Rosenmejer & Enghoff gen. et sp. nov., Emphysemastix frampt Olsen & Enghoff sp. nov., Agrophogonus hamulus Olsen & Enghoff sp. nov., and Agrophogonus pusillokiellandi Olsen & Enghoff sp. nov. Emphysemastix dracarys Olsen & Enghoff sp. nov. from Iringa city is described. Descriptive notes are given for Pogoro scharffi Hoffman, 2005, and Agrophogonus mwanihana Hoffmann, 2005. General gomphodesmid gonopod morphology is described and illustrated. A key to Udzungwa gomphodesmids is presented, as well as revised keys to all species of Emphysemastix and Agrophogonus. All gomphodesmid species from the Udzungwa Mts are mapped.
A large, newly collected material of the genus Aquattuor from the Udzungwa Mts, Tanzania, has been studied. Two new species are described, viz., A. mollilobus sp. nov. from the Udzungwa Mts and A. nguruensis sp. nov. from the Nguru Mts, Tanzania. Aquattuor claudiahempae Enghoff & Frederiksen, 2015, is recorded as new for the Udzungwa fauna. Characters from the first pair of male legs are illustrated for all Aquattuor species and are shown to be useful for species discrimination. A new gonopodal difference between A. submajor Enghoff, 2015 and A. udzungwensis Enghoff, 2015 is described and illustrated. Intrageneric relationships and distribution patterns are discussed. A distribution map and a key to all Aquattuor species are presented.
We revise the genus Attemsostreptus Verhoeff, 1941 based on type material of the type species, A. costatus Verhoeff, 1941, synonymise A. orobius (Kraus 1958) with A. costatus and describe a second species of the genus, A. reflexus sp. nov., collected from Kimboza Forest Reserve in Tanzania, and discuss the dubious tribe Trachystreptini.
Jeekelosoma Mauriès, 1985, is upgraded from subgenus status under Eviulisoma Silvestri, 1910 to full genus status. The type species, Jeekelosoma abadi (Mauriès, 1985) is redescribed based on topotypical material from a cave in Morocco. Jeekelosoma heptarachne sp. nov. and J. viginti sp. nov. are described from two further Moroccan caves.
This is the second part of a study of a large collection of Siphonophoridae Cook, 1895 from Brazil. Here, those without an abrupt division between the head and rostrum are considered. Two species have been distinguished and are described as Siphonophora setaepromissa sp. nov. and S. tuberculata sp. nov. Siphonophora hebetunguis (Attems, 1951) from the same region has many similarities with the two new species and is redescribed from type material. Comparisons have also been made with S. fuhrmanni Carl, 1914, and comments are made on the genera Cordillerium Verhoeff, 1941, Pterozonium Attems, 1951, Rhinosiphora Verhoeff, 1924, and Siphonophora Brandt, 1837. Finally, some suggestions for characters to look at in future collections of siphonophorids are made based on experience of this collection.
A large sample of Siphonophoridae from Brazil was studied; two morphological groups could be distinguished. Here species considered to be from the genus Columbianum Verhoeff, 1941 are examined in detail. The genus is known from Central and South America (Guatemala, Panama, Honduras, Guiana, Colombia, Peru and Brazil) and is characterised by a clear demarcation between head and rostrum in combination with long antennae, clearly surpassing the tip of the rostrum. A list of previously described species considered to belong to the genus is given; three new species are described: C. major sp. nov. has a large body size and a small head, C. nahvalr sp. nov. has a particularly pronounced domed head and a more castellated appearance to the body, C. adisi sp. nov. has a small body size and a very characteristic hind margin to the pleurites. Variation in the state of preservation of specimens hinders a diagnosis, but the examination of the accessory claw and details of the metazonital limbus and pleurite edges are helpful. Unusually for Diplopoda, the male gonopods are not very useful for identification. Ecological comments are given for each new species, one of which, C. adisi sp. nov., is from the seasonally flooded forest and appears to avoid inundation by climbing trees.
The ‘gigas’ group of dragon millipedes, formerly placed in the genus Desmoxytes Chamberlin, 1923, is revised and assigned to the new genus Gigaxytes gen. nov. Desmoxytes gigas Golovatch & Enghoff, 1994 is the type species of the new genus and is redescribed as G. gigas (Golovatch & Enghoff, 1994) gen. et comb nov. Three new species are described: G. fusca gen et sp. nov. from Thailand and Myanmar; G. parvoterga gen et sp. nov. and G. suratensis gen et sp. nov. from Thailand. All Gigaxytes species are endemic to small distribution areas in limestone habitats in South Thailand and South Myanmar. Illustrations of external morphological characters and an identification key to all known species are provided as well as a distribution map.
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.
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.
We describe five new species of fungi of the order Laboulbeniales Lindau growing on millipedes and belonging to the genera Diplopodomyces W.Rossi & Balazuc and Troglomyces S.Colla. Three new species of Diplopodomyces, viz. Diplopodomyces coronatus Santam., Enghoff & Reboleira sp. nov. living on Serboiulus spelaeophilus Gulicka, 1967 from Bulgarian caves, Diplopodomyces liguliphorus Santam., Enghoff & Reboleira sp. nov. on an unidentified species of Spirobolida from Sri Lanka, and Diplopodomyces ramosus Santam., Enghoff & Reboleira sp. nov. on Pachyiulus spp. from Turkey, Macedonia and Serbia; and two new species of Troglomyces, viz. Troglomyces dioicus Santam., Enghoff & Reboleira sp. nov. on Nepalmatoiulus sp. from Myanmar, and Troglomyces tetralabiatus Santam., Enghoff & Reboleira sp. nov. on Caucaseuma Strasser, 1970 and Heterocaucaseuma Antić & Makarov, 2016 from caves in Western Caucasus. Troglomyces dioicus sp. nov. is the first dioecious species described in the genus Troglomyces. Keys for all hitherto known species of Diplopodomyces and Troglomyces are presented, as is a discussion of the status of both genera. Additional records for Diplopodomyces lusitanipodos Santam., Enghoff & Reboleira and Troglomyces manfrediae S.Colla are also included.
Damacornu gen. nov. (type species: D. transversum gen. et sp. nov.), Geotypodon papei sp. nov. and Spinotarsus fortehamatus sp. nov. are described, and Helicochetus dimidiatus (Peters, 1855), H. mutaba Kraus, 1960 and Hoffmanides dissutus (Hoffman, 1963) are recorded from the Udzungwa Mts, Tanzania. A complete overview of the 39 odontopygid species now known from the Udzungwa Mts is given, including notes on endemism, biogeographical relationships and altitudinal distribution patterns.
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.
In this work we present a revision of the genus Ommatoiulus Latzel, 1884 in Portugal. Based on recently collected material and older museum samples, including type specimens, we describe six new species to science, viz. Ommatoiulus alacygni sp. nov., O. camurus sp. nov., O. denticulatus sp. nov., O. litoralis sp. nov., O. staglae sp. nov. and O. stellaris sp. nov. The species O. alacygni sp. nov., O. denticulatus sp. nov. and O. staglae sp. nov. described from the Algarve are outstanding by their extremely reduced mesomerital process. The species O. porathi (Verhoeff, 1893) and O. andalusius (Attems, 1927) are recorded and redescribed for the first time after their original description. The finding of O. andalusius – originally described from Andalusia in Spain – constitutes a new record for Portugal together with two species, viz. O. fuentei (Brolemann, 1920) and O. martensi Mauriès, 1969. The taxonomic status of several species is revised. Thus Archiulus (Schistocoxitus) cingulatus Attems, 1927 is here considered as a junior synonym of Ommatoiulus lusitanus (Verhoeff, 1895) while Schizophyllum cervinum Verhoeff, 1910 is synonymized with Ommatoiulus moreleti (Lucas, 1860). An identification key to all hitherto known Portuguese species of Ommatoiulus is presented as well as a distribution map illustrating the various species occurrences in the country.
Three new genera of Odontopygidae are described, all based on new species from the Udzungwa mountains, Tanzania, and all monotypic: Casuariverpa gen. nov. (type species: C. scarpa gen. et sp. nov.), Yia gen. nov. (type species: Y. geminispina gen. et sp. nov.), and Utiliverpa gen. nov. (type species: U. decapsulatrix gen. et sp. nov.). Similarities and differences between the new genera and other genera are discussed.
Two species of the genus Prionopetalum Attems, 1909, are recorded from the Udzungwa Mountains: P. asperginis sp. nov. and P. kraepelini (Attems, 1896). Prionopetalum stuhlmanni Attems, 1914, is synonymized under P. kraepelini. Odontopyge fasciata Attems, 1896, is transferred from Prionopetalum to Aquattuor Frederiksen, 2013, and new illustrations are given. A new illustrated key to species of Prionopetalum is provided.
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.
The new genus Geotypodon gen. nov. is described. It includes two species from the Udzungwa Mountains: G. millemanus gen. et sp. nov. (type species) and G. submontanus gen. et sp. nov., one species from nearby Iringa: G. iringensis gen. et sp. nov., and 18 previously described species hitherto incorrectly assigned to Odontopyge Brandt, 1841.
The genus Aquattuor Frederiksen, 2013 is revised. A. denticulatus Frederiksen, 2013 (type species) from the East Usambara Mts, Tanzania, is redescribed, and six new species are described: A. claudiahempae sp. nov. from Mt. Kilimanjaro, Tanzania, and five species from the Udzungwa Mts, Tanzania: A. longipala Enghoff sp. nov., A. major Enghoff sp. nov., A. stereosathe Enghoff sp. nov., A. submajor Enghoff sp. nov., and A. udzungwensis Enghoff sp. nov.
Chiraziulus is a highly disjunct, hitherto monotypic genus of cambalid millipedes, geographically isolated in Iran by more than 7000 km from its presumed closest relatives in East Asia and North America. Recent fieldwork in caves of Iran has provided several specimens of this genus, allowing the description of Chiraziulus troglopersicus sp. nov. The intraspecific variability of the type species, C. kaiseri Mauriès, 1983, is illustrated with scanning electron micrographs. Chiraziulus is characterized by exceedingly long microtrichose gonopod flagella which from their insertion points on the posterior face of the anterior gonopod coxites first point distad instead of basad or basad-posteriad as in most other flagelliferous Cambalidea (and Julida), then traverse a groove on the mesal surface of the anterior gonopod coxites, making a full (360°) loop. The same feature is also illustrated for the first time in the genus Cambala. The patterns and prevalence of the infection with a species of ectoparasitic fungus of the genus Rickia (order Laboulbeniales) in the type material of C. kaiseri is described. An updated review of the cave-adapted fauna of Iran is given.
Twenty new species of the millipede genus Chaleponcus Attems, 1914, are described from the Udzungwa Mountains: C. netus sp. nov., C. quasimodo sp. nov., C. malleolus sp. nov., C. scopus sp. nov., C. nikolajscharffi sp. nov., C. mwanihanensis sp. nov., C. basiliscus sp. nov., C. krai sp. nov., C. nectarinia sp. nov., C. circumvallatus sp. nov., C. ibis sp. nov., C. vandenspiegeli sp. nov., C. vilici sp. nov., C. teres sp. nov., C. hamerae sp. nov., C. termini sp. nov., C. gracilior sp. nov., C. mwabvui sp. nov., C. howelli sp. nov. and C. tintin sp. nov. Together with C. dabagaensis Kraus, 1958, they constitute the Chaleponcus dabagaensis-group, well characterized by apparently apomorphic gonopodal characters, presumably monophyletic, and the first example of a major radiation within the Udzungwas. All species are restricted to altitudes >1390 m, all but one were found in only one, rarely two forest reserves, and the vast majority of specimens were collected in montane forest. Chaleponcus gracilior sp. nov. was collected in four forest reserves, often in secondary habitats where other species were only exceptionally found. Co-occurrence of multiple species, inter-specific differences in body size and unusual tarsal setation of a few species tentatively suggest adaptive radiation.
Dolichoiulus typhlocanaria sp. nov., D. oromii sp. nov. and D. longunguis sp. nov. are described from caves and the mesovoid shallow stratum (MSS) on Gran Canaria. The genus Anagaiulus Enghoff, 1992 is synonymized under Dolichoiulus Verhoeff, 1900, resulting in Dolichoiulus blancatypa (Enghoff, 1992) comb nov.