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Some cryptic Korean karst creatures: revalidation of the pseudoscorpion genus Spelaeochthonius (Pseudoscorpiones: Pseudotyrannochthoniidae) and description of two new species from Korea

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... From a biological perspective the genus is interesting because most of the species have been collected from high altitude habitats with harsh climatic conditions that include regular frost and snow as well as long winters; habitats that are too cold for most other pseudoscorpions. Biogeographically, the genus seems to belong to an ancient Laurasian radiation of pseudotyrannochthoniids with 16 setae on the carapace, as its closest morphological relatives occur either in the warm temperate forests of western North America (Oregon, Washington, California and Utah) (Muchmore 1967;Benedict & Malcolm 1970) or are exclusively confined to caves in Japan and Korea (You et al. 2022;also Fig. 1). ...
... Amongst the three pseudotyrannochthoniid genera known to occur in Asia today, the speciose and morphologically diverse genus Allochthonius includes species in which these spines sit on a common protuberance (a ''spray '' or ''fan'' according to Chamberlin 1929a) at the apical margin of the leg coxa, with each spine being tripartite, relatively short and with a spatulate apical end (Fig. 3). The newly reinstated genus Spelaeochthonius Morikawa, 1954 from caves in Japan and Korea has 9-11 coxal blades that are inserted directly on the subapical margin of the pedipalp coxa, never sit on a common protuberance, and each spine is rather long and has plumose or tassel-shaped tips (e.g., Morikawa 1954Morikawa , 1956Morikawa , 1960You et al. 2022;Fig. 5). ...
... Muchmore (1967) did not mention Centrochthonius or any of the literature pertaining to this genus, and the synonymy of Spelaeochthonius with Pseudotyrannochthonius -suggested as provisional by Muchmore (1967) himself -was abandoned by Schwarze et al. (2022) who rediagnosed Centrochthonius for species with only 4-5 coxal blades that are situated subapically on the antero-mesal margin of leg coxa I, and individual spines that are short, tripartite and have an acute tip. Schwarze et al. (2022) also emphasized the importance of the number of carapaceal setae in Holarctic pseudotyrannochthoniids and noted the presence of only 16 setae on the carapace in Centrochthonius, a similar configuration to Spelaeochthonius from caves in Japan and Korea [currently with nine valid species, see You et al. 2022], and to the three species from western U.S.A. that presently remain misplaced in Pseudotyrannochthonius but are rather related to Centrochthonius and Spelaeochthonius. ...
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The first new fossil pill millipede, order Glomerida, description in more than 150 years is based on two specimens, one male and one female, found in Baltic amber. Both specimens were studied with light- and micro-CT technology, allowing a detailed reconstruction of the posterior telopods and numerous other morphological characters. Based on these characters, the specimens can be securely placed in the megadiverse, mainly Asiatic genus Hyleoglomeris Verhoeff, 1910 as a new species, Hyleoglomeris groehni n. sp. The species H. groehni n. sp. is the oldest and only known fossil member of the Glomerida that is sufficiently described to allow for placement in a family or genus. The discovery of a fossil Hyleoglomeris in Baltic amber has biogeographic implications as the genus is now speciose in Asia, with the nearest relatives found in the Caucasus and some caves in Serbia and Greece. With the discovery of a fossil European Hyleoglomeris species, the cave Hyleoglomeris species are interpreted as relicts, surviving underground as living witnesses of a different time when Hyleoglomeris was still extant in Europe. Other cave Glomerida in Europe, as well as morphologically aberrant pill millipede species endemic to the Canary Islands, should be checked for potential placement in Hyleoglomeris.
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Lagynochthonius fragilis n. sp. is described from a limestone cave in the Hong Chong karst of Kien Giang Province, southern Vietnam, which is currently threatened by quarrying activities. This is the first record of a troglomorphic species of Lagynochthonius Beier, 1951 from continental Asia. The presence of chemosensory setae on the dorsum of the chelal palm is interpreted as a synapomorphy of the tribe Tyrannochthoniini Chamberlin, 1962. The New Zealand genus Maorichthonius Chamberlin, 1925 is transferred from the Chthoniini Daday, 1888 to the Tyrannochthoniini. The genus Tyrannochthoniella Beier, 1966, also endemic to New Zealand, is assigned to the tribe Chthoniini Daday, 1888. The genus Stygiochthonius Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001, from southern Spain, is synonymized with Paraliochthonius Beier, 1956 (n. subj. syn.). Five new combinations are proposed: Lagynochthonius ovatus Vitali-di Castri, 1984 (ex Tyrannochthonius); Paraliochthonius barrancoi (Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001) (ex Stygiochthonius); P. curvidigitatus (Mahnert, 1997) (ex Lagynochthonius); P. setiger (Mahnert, 1997) (ex Tyrannochthonius); and P. superstes (Mahnert, 1986) (ex Tyrannochthonius). A key is given to the genera of the Tyrannochthoniini. The parallel evolution in several groups of pseudoscorpions of a characteristic chelal morphology, here termed lagyniform, is discussed. New designations are proposed for the spot-sensilla of the chelal fingers. The so-called ‘sensorium’ near the tip of the fixed chelal finger of Lagynochthonius species is shown to be a modified tooth that has migrated dorsally from the dental margin. The new term rallum is introduced as a replacement for the inappropriate term ‘flagellum’, as applied to the cheliceral blades of pseudoscorpions. The term bothridial vestibulum is introduced for the internal cuticular sheath at the base of the bothridia of the trichobothria.Lagynochthonius fragilis n. sp. est décrit d’une grotte calcaire de la province de Kien Giang, au sud du Vietnam, actuellement menacée par une exploitation de carrière. Elle est la première espèce troglomorphe du genre Lagynochthonius Beier, 1951 connue de l’Asie continentale. La présence des soies chemosensorielles sur la main de la pince est interprétée comme une synapomorphie de la tribu des Tyrannochthoniini Chamberlin, 1962. Le genre néo-zélandais Maorichthonius Chamberlin, 1925 est transféré des Chthoniini Daday à la tribu des Tyrannochthoniini. Le genre Tyrannochthoniella Beier, 1966, également endémique de la Nouvelle Zélande, est attribué à la tribu des Chthoniini Daday, 1888. Le genre Stygiochthonius Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001, du sud de l’Espagne, est mis en synonymie avec Paraliochthonius Beier, 1956 (n. syn. subj.). Cinq combinaisons nouvelles sont proposées : Lagynochthonius ovatus Vitali-di Castri, 1984 (ex Tyrannochthonius) ; Paraliochthonius barrancoi (Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001) (ex Stygiochthonius) ; P. curvidigitatus (Mahnert, 1997) (ex Lagynochthonius) ; P. setiger (Mahnert, 1997) (ex Tyrannochthonius) ; et P. superstes (Mahnert, 1986) (ex Tyrannochthonius). Une clé de détermination des genres de Tyrannochthoniini est fournite. L’évolution parallèle des facies caractéristiques de la pince, ici qualifié de “ lagyniforme ”, est évoquée chez plusieurs groupes de pseudoscorpions. Desnouveaux sigles sont proposés pour les sensilles punctiformes des doigts de la pince. Il est démontré que le “ sensorium ” à l’extrémité du doigt fixe de la pince des espèces de Lagynochthonius est une dent modifiée qui a migré dorsalement dès la marge dentale. Le terme inapproprié de “ flagelle ”, dans le sens de son application aux lames chélicèriennes des pseudoscorpions, est remplacé par rallum. Le terme nouveau vestibule trichobothriale est introduit pour la gaine cuticulaire à la base des bothridies des trichobothries.
Article
The south-western division of Australia is the only biodiversity hotspot in Australia and is well-known for extreme levels of local endemism. Climate change has been identified as a key threat for flora and fauna, but very few data are presently available to evaluate its impact on invertebrate fauna. Here, we derive a molecular phylogeography for pseudoscorpions of the genus Pseudotyrannochthonius that in the south-west are restricted to regions with the highest rainfall. A dated molecular phylogeny derived from six gene fragments is used for biogeographic reconstruction analyses, spatial mapping, environmental niche-modelling, and to infer putative species. Phylogenetic analyses uncover nine clades with mostly allopatric distributions and often small linear ranges between 0.5 and 130 km. Molecular dating suggests that the origins of contemporary diversity fall into a period of warm/humid Palaeogene climates, but splits in the phylogeny coincide with major environmental shifts, such as significant global cooling during the Middle Miocene. By testing several models of historical biogeography available for the south-west, we determine that Pseudotyrannochthonius is an ancient relict lineage that principally follows a model of allopatric speciation in mesic zone refugia, although there are derivations from this model in that some species are older and distribution patterns more complex than expected. Ecological niche models indicate that drier and warmer future climates will lead to range contraction towards refugia of highest rainfall, probably mimicking past variations that have generated high diversity in these areas. Their conservation management will be crucial for preserving the unique biodiversity heritage of the south-west.
Chapter
Why certain animals lose features believed essential, like eyes, bodily color, and robustness, to live permanently underground has long intrigued biologists and laymen. Many of these features evolved independently and shared among diverse groups living in caves including both terrestrial and aquatic cavernicoles. The degree of change often correlates with the level of association of the species to caves. This association allowed development of a classification scheme to help understand the evolutionary ecology of cave communities. The refined scheme, called the Schiner-Racovitza system, is based on both morphology and ecology. The categories are troglobionts and stygobionts (animal species that obligately live underground on land or in water, respectively), troglophiles (animals that can live and reproduce in both underground and surface habitats), and trogloxenes (animals that regularly visit caves for food or refuge). Common adaptations to cave life involve morphology, behavior, and physiology. In addition to the conspicuous losses, many compensatory traits have evolved, such as longer appendages, longer and more slender body, more and larger sensory structures, and specialized mouthparts and tarsi. Modified behavioral traits include reduction in circadian rhythm, reduced dispersal ability, slower but nearly continuous activity, and modified mating behavior. Physiological adaptations include low metabolism rate, dietary changes, resistance to starvation, modified water balance mechanisms, tolerance to high CO2 and low O2, and increased longevity. Cave-adapted animals also display greater K-selection with fewer and larger eggs and reduced life cycle.
Article
Two new troglomorphic pseudoscorpion species, Bisetocreagris maomaotou sp. nov. (Family Neobisiidae) and Tyrannochthonius chixingi sp. nov. (Family Chthoniidae) are described from one cave in the tower karst of northern Guangxi Province, China. This cave is located at close proximity to a village and an adjacent urban area. As with many caves in the South China Karst, this feature occurs at an elevation slightly above agriculture and rural activities; thus, we suggest it may be partially buffered from human activities in the lowland areas. We discuss the likelihood of narrow range endemism and provide research and conservation recommendations to guide future management of these two species.
Article
Pseudoscorpions are one of the common arthropod in soil mesofauna but are infrequently studied in East Asia. The fauna in South Korea is not adequate enough, and practical faunistic survey of pseudoscorpions have not been conducted in last 20 years. In this article, the current pseudoscorpion fauna in South Korea is reviewed, and the results of the survey in Chungcheongnam-do Province, the DNA barcoding (mitochondrial DNA cytochrome c oxidase I sequencing) and molecular phylogenetic analysis are shown. A total 64 specimens, including four species, Allochthonius (Allochthonius) buanensis, Bisetocreagris japonica, Bisetocreagris turkestanica (first record from South Korea), and Microbisium pygmaeum, were collected. The result of molecular phylogenetic analysis based on the cytochrome c oxidase I sequences (427 bp) implied the existence of a possible cryptic species within A. (A.) buanensis. Faunistic survey and DNA barcoding of South Korean species can greatly contribute to the understanding of East Asian pseudoscorpion systematics. © 2018 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA)
Chapter
This chapter presents the procedure and results of the cave evaluation process integrated into geoheritage inventorying in South Korea. Since 2003, the Korean Government has investigated and evaluated caves in order to define which ones should be protected under the existing legal framework of the country. Geological criteria were prepared by cave scientists and applied to all studied caves. A total of 1022 caves have been assessed to date and the Cultural Heritage Administration has legally protected the ones with significant geoheritage value. This project will continue until the whole inventory in South Korea is completed. The results of this inventory will also allow the protection of caves in Korea from development pressure in the future and will provide valuable information for forthcoming scientific research.
Article
The main Korean mountain range [the so-called “Baekdudaegan” (BDDG)] is thought to have served as a glacial refugium, mainly for the boreal and temperate flora of northeastern Asia. To test this hypothesis, we have critically reviewed (i) the genetic literature focused on plants for which Korean populations have been studied or Korean endemic plants, and (ii) palaeoecological studies with data from the Korean Peninsula and/or from neighboring regions. Consistent with this hypothesis, Korean populations showed higher intrapopulation genetic diversity (and, in some cases, with latitudinal decreases of genetic variation), harbored ancestral haplotypes, and/or exhibited significant amounts of unique haplotypes/alleles. Palaeoecological data support the genetic studies, indicating that the BDDG sustained an assemblage of boreal and temperate forests at the Last Glacial Maximum (LGM). Given its floristic, biogeographic, and cultural value (it is regarded as a “holy mountain system” in Korea), the BDDG merits high priority for conservation. In order to guarantee the preservation of the BDDG, we suggest enlarging the current BDDG Mountains Reserve in South Korea, whereas in North Korea the accelerated deforestation should be stopped and new protected areas should be set up. Cooperation between the two Koreas is also needed, and may constitute an important step towards more ambitious initiatives such as the creation of the “Ecological Corridor of Northeast Asia”.
Article
Two new pseudoscorpion species, Pseudotyrannochthonius cheni sp. n. and Allochthonius (A.) fanjingshan sp. n. (family Pseudotyrannochthoniidae Beier, 1932) are described and illustrated. P. cheni sp. n. represents the first record of this genus from China and is also the first epigean (surface) species to be recorded from eastern Asia because all other species (from Korea and Japan) occur in subterranean habitats. A. (A.) fanjingshan sp. n. is the eighth species in this genus from China and reflects a wider distribution of the genus in this country. Detailed diagnosis, descriptions and illustrations are presented together with distribution maps and some habitat information.
Article
Fully troglobitic pseudoscorpions are rare in the Afrotropical Region, and we explored the identity and phylogenetic relationships of specimens of a highly modified troglobite of the family Gymnobisiidae in the dark zone of the Wynberg Cave system, on Table Mountain, South Africa. This large pseudoscorpion - described as Gymnobisium inukshuk Harvey & Giribet, sp. nov. - lacks eyes and has extremely long appendages, and has been found together with other troglobitic fauna endemic only to this cave system. Phylogenetic analyses using the nuclear ribosomal genes 18S rRNA and 28S rRNA and the mitochondrial protein-encoding gene cytochrome c oxidase subunit I unambiguously place the new species with other surface Gymnobisium from South Africa. This placement receives strong support and is stable to analytical treatments, including static and dynamic homology, parsimony and maximum likelihood, and data removal for ambiguously aligned sites. This species is the first troglobitic species of the family and one of the most highly modified pseudoscorpions from the Afrotropical Region. http://zoobank.org/urn:lsid:zoobank.org:pub:5227092B-A64B-4DB3-AD90-F474F0BA6AED.
Article
A new pseudoscorpion species, Allochthonius exornatus sp. n., belonging to the family Pseudotyrannochthoniidae Beier, 1932, is described from China. Detailed diagnosis, descriptions and illustrations are presented, and a key to the genus Allochthonius Chamberlin, 1929 is provided.
Article
Two new pseudoscorpion species of Allochthonius are described: Allochthonius (Allochthonius) liaoningensis sp. n. and A. (Urochthonius) brevitus sp. n. The latter represents species of the subgenus Allochthonius (Urochthonius) found in China for the first time.
Article
Lagynochthonius fragilis n. sp. is described from a limestone cave in the Hong Chong karst of Kien Giang Province, southern Vietnam, which is currently threatened by quarrying activities. This is the first record of a troglomorphic species of Lagynochthonius Beier, 1951 from continental Asia. The presence of chemosensory setae on the dorsum of the chelal palm is interpreted as a synapomorphy of the tribe Tyrannochthoniini Chamberlin, 1962. The New Zealand genus Maorichthonius Chamberlin, 1925 is transferred from the Chthoniini Daday, 1888 to the Tyrannochthoniini. The genus Tyrannochthoniella Beier, 1966, also endemic to New Zealand, is assigned to the tribe Chthoniini Daday, 1888. The genus Stygiochthonius Carabajal Márquez, Garcia Carrillo & Rodríguez Fernândez, 2001, from southern Spain, is synonymized with Paraliochthonius Beier, 1956 (n. subj. syn.). Five new combinations are proposed: Lagynochthonius ovatus Vitali-di Castri, 1984 (ex Tyrannochthonius); Paraliochthonius barrancoi (Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001) (ex Stygiochthonius); P. curvidigitatus (Mahnert, 1997) (ex Lagynochthonius); P. setiger (Mahnert, 1997) (ex Tyrannochthonius); and P. superstes (Mahnert, 1986) (ex Tyrannochthonius). A key is given to the genera of the Tyrannochthoniini. The parallel evolution in several groups of pseudoscorpions of a characteristic chelal morphology, here termed lagyniform, is discussed. New designations are proposed for the spot-sensilla of the chelal fingers. The so-called 'sensorium' near the tip of the fixed chelal finger of Lagynochthonius species is shown to be a modified tooth that has migrated dorsally from the dental margin. The new term rallum is introduced as a replacement for the inappropriate term 'flagellum', as applied to the cheliceral blades of pseudoscorpions. The term bothridial vestibulum is introduced for the internal cuticular sheath at the base of the bothridia of the trichobothria. Pseudoscorpion, Tyrannochthoniini, taxonomy, morphology, endangered species, cave, Vietnam, New Zealand, Spain, sensilla
Article
Cave-dwelling pseudoscorpions (Arachnida, Pseudoscorpiones) from Brazil. - Pseudoscorpions collected in about 100 caves have been studied and 25 species of seven families identified. Two new genera - Spelaeobochica (Bochicidae) (type species allodentatus sp. n.) and Spelaeochernes (Chernetidae) (type species dentatus sp. n., with 7 other new species) - are described, together with 10 additional new species in the genera Pseudochthonius (Chthoniidae), Ideoroncus (Ideoroncidae), Progarypus (Olpiidae), Cheiridium (Cheiridiidae) and Zaona (Chernetidae: previously known from its type species from Florida only). Geogarypus itapemirinensis Feio is redescribed and transferred to the genus Cheiridium.
Article
The Last Glacial Maximum (LGM) at 21,000 years before present is a marked climate event with thick ice sheets over North America and Europe and associated sea level drop of about 120 m. We used the most updated coupled climate model results to analyze the climate change over Korea for the LGM. Eight coupled models (CCSM, CNRM, COSMOS, FGOALS, IPSL, MIROC, MPI, and MRI) were used to analyze the LGM climate. With LGM boundary conditions, surface air temperature decreases almost everywhere in all seasons with the largest cooling in winter and least cooling in summer. Overall, in the LGM, the multi-model annual-mean cooling over Korea is 5.85 C°, which is consistent with proxy evidence using pollen reconstructions. Associated with the surface cooling, precipitation decreases in general, with the largest reduction in winter by 56%, but in summer precipitation increases by 12% in the LGM over Korea. Overall, precipitation decreases by about 14% in the LGM over Korea. The surface cooling tends to increase surface pressure almost everywhere over Asia. The sea level pressure increase is especially larger in high latitudes and this leads to the easterly wind anomaly in northern part of Korea. On the other hand, in South Korea, the increase in surface pressure leads to the westerly wind anomaly. The increase in surface pressure associated with surface cooling leads to the anticlockwise wind anomalies in the LGM over Korea.
Article
Two new species of pseudotyrannochthoniine pseudoscorpions are described from western Oregon and northern Utah. These, together with Allochthonius incognitus Schuster, 1966 are placed in the genus Pseudotyrannochthonius, which has heretofore been known only from South America and Australia.
Article
The pseudoscorpion genus Allochthonius Chamberlin, 1929, belonging to the family Pseudotyrannochthoniidae Beier, 1932, is reported from China, and the subgeneric characters of Allochthonius (Allochthonius) are reviewed in detail. Three new species are diagnosed, described and illustrated under the names Allochthonius (A.) fuscus sp. nov., A. (A.) wui sp. nov. and A. (A.) trigonus sp. nov. A distribution map and a key to the species of subgenus Allochthonius (A.) are provided. In addition, Centrochthonius sichuanensis Schawaller, 1995 is transferred to Allochthonius, forming the new combination A. (A.) sichuanensis (Schawaller).
Article
: The middle Devonian (Givetian–Eifelian) pseudoscorpion Dracochela deprehendor Schawaller, Shear and Bonamo is redescribed from the type material and an additional palpal fragment. Dracochela differs from extant pseudoscorpions in having numerous spinules on the leg tarsi, the femur at least as long as the patella on the posterior legs, the stem of the arolia thick, most blades of the serrulae only weakly fused and in lacking a spinneret on the chelicera. The blades of the cheliceral rallum are shown to have been arranged in two rows, as in most Heterosphyronida. The cheliceral serrulae are compared with analogous structures in other arachnids (Notostigmata, Opiliones, Palpigradi, Schizomida and Scorpiones), and it is concluded that the panctenal state (all lamellae attached to finger) is plesiomorphic relative to the hemictenal state (apical lamellae raised), which has evolved independently in Heterosphyronida and Neobisioidea. The trichobothriotaxy of the chela of Dracochela is shown to be similar to that of the extant family Pseudotyrannochthoniidae. The growth of the chelal fingers followed the same pattern as that seen in modern pseudoscorpions, with most of the increase in length occurring at the base of the fingers. The family Dracochelidae Schawaller, Shear and Bonamo is treated as a plesion and assigned to the stem-group of Pseudoscorpiones. The ordinal name Chelonethi Thorell is restricted to crown-group pseudoscorpions, and the superordinal name Pseudoscorpiones Latreille is adopted for the total-group (i.e. stem-group plus Chelonethi).
Article
In Korea, only a few cave spiders have so far reported, so little is known as their fauna. The author investigated on the caves, various part of this country to have taken up the study of these spiders, detailed report will be made in future, but for the present, has dealt with as follow; 1) As a result of the historical review of the study on cave spiders have been recorded 39 species of 27 genera belonging to 15 families, including 16 new species in Korea. 2) It were recorded that the name and location of 154 caves; 121 limeston caves, 33 lava caves which surveyed by the author. 3) A list of cave spiders made out about 100 species 60 genera belonging to 23 families. However there are considerable number of species which have not yet been identified and waiting for descriptions particularly in the troglobiontic species. Especially the Leptonetid, Nesticid and Cybaeid species are very important matrials for discussing the variatoin and lineage of species through isolation. 4) Species common to foreign countries are restricted to a few; 25 species of Japan, 2 species of Europe. There are not exist troglobiontic species, and most of them are trogloxenous ones.
Article
As a result of this report, the pseudoscorpion family Chthoniidae in Korea comprises nine species in four genera. All species are collected mainly from the litter layer in wooded forests. The two new species, Tyrannochthonius suppressalis n. sp. and Tyrannochthonius spinatus n. sp., described herein occur on islands in the Yellow Sea.