Abstract
Phoresy is an important dispersal strategy in pseudoscorpions, yet associations with Diptera have not been comprehensively revised for more than two decades. An updated synthesis of pseudoscorpion-Diptera phoresy is presented based on newly collected European material, a revision of previously studied Slovak material, a critical reassessment of worldwide published records, and available fossil evidence. In total, 11 previously unreported cases of pseudoscorpion phoresy from four European countries are documented, including the first records of Dactylochelifer degeerii (C.L. Koch, 1835) from France and of Pselaphochernes scorpioides (Hermann, 1804) from Slovenia. Seven dipteran species and three dipteran families are newly recorded worldwide as hosts of pseudoscorpion phoresy, and one case of multiple phoresy was recorded from Slovakia. Altogether, 172 records spanning the period from 1761 to 2025 were compiled and evaluated. Compared with previous syntheses, the number of known dipteran hosts was increased to 74 species from 30 families, and pseudoscorpion phoronts to 39 species from seven families. This study provides the most comprehensive overview of pseudoscorpion-Diptera phoresy to date and establishes a robust framework for future taxonomic, ecological, and evolutionary studies.
Key words: Dispersal, distribution, fly, host specificity, phoront
Introduction
Even though arachnids are generally incapable of active long-distance dispersal, they occur in almost all terrestrial habitats. Their spread is primarily mediated by external vectors, with short-range dispersal facilitated by other animals and long-distance dispersal mainly achieved through air currents. Owing to these mechanisms, arachnids, together with insects, are often among the first animals to colonise newly available habitats (Szymkowiak et al. 2007). Dispersal strategies, such as phoresy or ballooning, are well-documented in spiders, mites, and pseudoscorpions and are typically associated with specific morphological and trophic adaptations, habitat preferences, and the availability and type of suitable carriers (Szymkowiak et al. 2007).
Phoresy is a dynamic, interspecific, and temporary relationship in which a phoront attaches to a host for the time required to migrate from one habitat to another, with dispersal being the primary outcome of this interaction (Camerik 2010). Records of phoretic pseudoscorpions date back to the eighteenth century (Poda 1761), yet the reasons underlying passive dispersal via phoresy in pseudoscorpions remain poorly known. Two hypotheses have been proposed to explain phoresy in pseudoscorpions (summarised in Poinar et al. 1998). The first hypothesis – that transport by other arthropods is accidental, motivated by hunger, and occurs because pseudoscorpions are incapable of consuming their hosts – had been rejected by Zeh and Zeh (1992b). The second hypothesis considers phoresy to be an adaptive behaviour functioning primarily for dispersal and is currently accepted. Phagophily does not represent the primary motivation for phoretic associations (Zeh and Zeh 1992c). Although some species may prey on mites present on their hosts or use host bodies as strategic sites for mating interactions, dispersal remains the principal function of phoresy in pseudoscorpions (Zeh and Zeh 1992c).
At a broader scale, phoresy has been reported across a wide range of animal taxa. A synthesis by Bartlow and Agosta (2021) showed that phoretic dispersal has been documented in animals belonging to 13 phyla, 25 classes, and 60 orders, predominantly involving small invertebrates. Fossil evidence, mainly preserved in amber, indicates that phoresy is an ancient phenomenon, with records particularly well documented for mites, springtails, and pseudoscorpions (Magowski 1995; Poinar et al. 1998; Penney et al. 2012). Some mite records date back approximately 85 million years (Magowski 1995), whereas pseudoscorpion phoresy is known from fossils up to 40 million years old (Poinar et al. 1998).
Within Pseudoscorpiones, phoresy has been documented in 12 of 26 recent families: Atemnidae, Cheiridiidae, Cheliferidae, Chernetidae, Chthoniidae, Garypinidae, Geogarypidae, Larcidae, Neobisiidae, Sternophoridae, Syarinidae, and Withiidae (e.g., Ressl 1965; Aguiar and Bührnheim 1998b; Poinar et al. 1998; Harms and Dunlop 2017), with Garypinidae restricted to fossil material. Phoretic pseudoscorpions have been reported from a wide range of hosts, including mammals, birds, ten insect orders (Coleoptera, Diptera, Hemiptera, Hymenoptera, Lepidoptera, Mecoptera, Neuroptera, Odonata, Orthoptera, and Plecoptera), as well as arachnids such as harvestmen, spiders, and scorpions (Aguiar and Bührnheim 1998a; Poinar et al. 1998; Harvey et al. 2015; Christophoryová et al. 2017b; Hetešová and Christophoryová 2022; Warburg et al. 2023b). Neuroptera were not included among phoretic hosts in the comprehensive overview by Poinar et al. (1998), but were subsequently reported by Aguiar and Bührnheim (1998b). As a result, Neuroptera are occasionally overlooked in later studies relying primarily on the earlier summary.
Among the various host groups, Diptera represent one of the most frequently reported phoretic hosts of pseudoscorpions (Poinar et al. 1998). This prominence can be attributed to their high abundance, ecological diversity, and frequent contact with microhabitats occupied by pseudoscorpions, such as decaying organic matter, tree hollows, bark, and nests. Many dipteran species are capable of active flight over considerable distances, making them effective vectors for passive dispersal. In addition, the apparent importance of Diptera as phoretic hosts is partly influenced by sampling bias, as flies are commonly collected using methods that increase the likelihood of detecting phoretic associations.
The most comprehensive synthesis of pseudoscorpion phoresy involving Diptera remains that of Poinar et al. (1998), which compiled records across all arthropod host groups. In the study, phoretic associations between pseudoscorpions and Diptera comprised 37 dipteran species (32 genera) from 17 families hosting 23 pseudoscorpion species (16 genera) from four families. Despite being more than a quarter of a century old, this work continues to be widely cited and serves as a primary reference for pseudoscorpion–Diptera phoretic associations. However, subsequent use of this compilation has revealed inconsistencies, duplications, and taxonomic inaccuracies, reflecting both the state of knowledge at the time and later advances in pseudoscorpion and dipteran systematics. Moreover, numerous records documented in earlier syntheses, such as Beier (1948), as well as data published after Poinar et al. (1998), were not included. These limitations highlight the need for an updated and critically revised synthesis.
During our research, we obtained valuable new material documenting pseudoscorpion phoresy on Diptera from Europe. Accordingly, we present a synthesis of pseudoscorpion–Diptera host–phoront associations based on (i) newly collected material, (ii) the revision of previously studied Slovak material, (iii) a critical reassessment of published records worldwide, and (iv) available fossil evidence.
Materials and methods
New material examined
Pseudoscorpions were collected individually or using pyramid trap and Malaise traps at 11 localities in Czechia, France, Slovakia, and Slovenia (Fig. 1). Specimens were studied as temporary slide mounts prepared by immersion in lactic acid for clearing. After examination, they were rinsed in water and returned to 75% ethanol. All specimens were examined using a Leica DM1000 compound microscope with an ICC50 camera module (LAS EZ v. 3.4.0). Measurements were taken from digital images using AxioVision 40LE (v. 4.6.3.0). Digital photographs were taken using a Canon EOS 5D Mark II camera attached to a Zeiss Axio Zoom V16 stereomicroscope. Image stacks were created manually and merged using Zerene Stacker software (v.1.4). Spatial data used in the map were converted from original coordinates and visualised in QGIS (v. 3.36.2). Figures were edited in Adobe Photoshop CC (v. 25.6.0).
Figure 1.
New records of pseudoscorpion phoresy in Europe. Locality codes are given in Materials and methods. Country abbreviations: CZ – Czechia, FR – France, SI – Slovenia, SK – Slovakia.
Pseudoscorpions were identified using the key in Krajčovičová et al. (2022) and data from Christophoryová et al. (2023). The latter study (Christophoryová et al. 2023) employed an interactive approach that combined molecular DNA barcoding, cytogenetic analysis, and multivariate statistical methods to investigate species boundaries within the genus Lamprochernes. These results are particularly relevant for specimens identified as L. chyzeri (Tömösváry, 1883), as the study revealed the presence of a cryptic species, L. abditus Christophoryová, Krajčovičová, Šťáhlavský, Španiel & Opatova, 2023, which is morphologically indistinguishable from L. chyzeri and can be reliably separated only using molecular and cytogenetic evidence. Because the examined specimens were not preserved in pure ethanol, molecular analyses could not be performed; therefore, the presence of L. abditus in the study area cannot be excluded. Consequently, specimens of this taxon in both the newly collected material and the revised Slovak material are herein referred to as Lamprochernes cf. chyzeri.
Several colleagues assisted with the identification of dipteran hosts and are acknowledged in the Results - New material examined section. One host specimen from locality 3 was identified only to the genus level because the material was poorly preserved and lacked diagnostic coloration. In some cases, the attachment site of phoretic pseudoscorpions was noted by the collector as the host’s leg; however, subsequent detachment of the specimens from the flies prevented more precise localisation of the attachment site on the leg (localities 3, 5, and 11).
Pseudoscorpions and the majority of Diptera hosts are deposited in the zoological collection of the Department of Zoology, Comenius University in Bratislava, Slovakia (depository codes FO 40/6–FO 50/6). Hosts from localities 1 and 8 are deposited in the Silesian Museum, Opava, Czechia.
Revision of published data on phoresy from Slovakia
Published records on pseudoscorpion–dipteran phoresy from Slovakia (Mašán and Krištofík 1992; Christophoryová et al. 2011, 2017a, 2018, 2021; Červená et al. 2019; Hetešová and Christophoryová 2022) were revised in detail, as original material or supporting documentation was accessible to the authors. Several hosts, phoronts, or their sexes were additionally identified.
Published data on the phoresy of Pseudoscorpiones and Diptera worldwide
Published data on the phoresy of Pseudoscorpiones and Diptera were reviewed from the earliest available records to the beginning of 2026. The literature survey was conducted with particular emphasis on two major revisionary works (Beier 1948; Poinar et al. 1998), which served as reference frameworks for reassessing earlier and subsequent records. In addition, all accessible publications were examined, including the catalogue (WPC 2026) as well as papers by key specialists in pseudoscorpion research, in order to retrieve all published information on phoresy.
The most recent comprehensive revision of published data on pseudoscorpion–dipteran phoresy was provided by Poinar et al. (1998). Their dataset, which summarised records by listing the host, phoront, continent, and literature source, was critically re-examined in the present study, with each original publication cited therein being individually reviewed, and the geographic information refined by replacing continent-level records with the precise country of occurrence.
Pseudoscorpion taxa were verified against WPC (2026), and issues related to synonymy, invalid names, and subsequent taxonomic revisions were resolved accordingly (see Notes).
Host identifications of Diptera were updated to their currently accepted names (see Notes), in many cases following consultation with dipteran specialists.
The same methodological approach was applied to the processing and evaluation of fossil data.
Results and discussion
New material examined
In total, 11 previously unreported cases of phoresy were documented in the present study from four European countries (Fig. 1; Table 1, data marked as “Present study”). The highest number of records was obtained from Slovakia (five) and France (four). Four pseudoscorpion species belonging to two families (Chernetidae and Cheliferidae) were found attached to the legs of 11 dipteran hosts representing nine families (Fanniidae, Limoniidae, Lonchaeidae, Muscidae, Scatopsidae, Sciomyzidae, Tipulidae, Ulidiidae, and Xylophagidae). One record from Slovakia involved multiple phoresy (Table 1). Dactylochelifer degeerii was recorded for the first time from France, and both the genus and species Pselaphochernes scorpioides were recorded for the first time from Slovenia. In addition, seven dipteran species were documented for the first time as hosts of pseudoscorpion phoresy (Ceroxys urticae, Limonia nubeculosa, Neolimonia dumetorum, Pherbellia annulipes, Scatopse notata, Tipula vernalis, Xylophagus ater) together with three dipteran families newly reported as hosts (Scatopsidae, Sciomyzidae, Xylophagidae) (Table 1).
Table 1.
Extant records of phoresy between Diptera and Pseudoscorpiones.
| Host taxa | Phoront taxa | Country | References | |
|---|---|---|---|---|
| Anthomyiidae | ||||
| Anthomyia pluvialis (Linnaeus, 1758) | Chernetidae | Lamprochernes cf. chyzeri (Tömösváry, 1883) | SK | Hetešová and Christophoryová (2022) |
| Anthomyia Meigen, 1803 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*1 | DE | Gerstaecker (1862) |
| Delia floralis (Fallén, 1824)2 | Chernetidae | Allochernes peregrinus Lohmander, 1939 | SE | Lohmander (1939a, 1939b) |
| Delia platura (Meigen, 1826) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SK | Christophoryová et al. (2018) |
| Eutrichota affinis (Stein, 1898)3 | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | US | Muchmore (1971) |
| Eutrichota apicalis (Stein, 1898)4 | Chernetidae | Hesperochernes Chamberlin, 1924 | US | Muchmore (1971) |
| Calliphoridae | ||||
| Calliphora vicina Robineau-Desvoidy, 1830*5 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*5 | DE | Leydig (1867, 1881, 1893) |
| Calliphora vicina Robineau-Desvoidy, 1830*6 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)6 | GB-ENG | Pickard-Cambridge (1892); Kew (1901) |
| Calliphora vomitoria (Linnaeus, 1758)7 | Cheliferidae | Chelifer cancroides (Linnaeus, 1758)7 | GB-ENG | Donovan (1797) |
| Calliphora vomitoria (Linnaeus, 1758)8 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*8 | GB-ENG | Kirby and Spence (1826) |
| Calliphora Robineau-Desvoidy, 1830*9 | Not specified | Not specified | AR | Berg (1893) |
| Lucilia caesar (Linnaeus, 1758)10 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SK | Mašán and Krištofík (1992) |
| Lucilia sericata (Meigen, 1826) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | ES | Castillo-Miralbes (2002) |
| Stevenia Robineau-Desvoidy, 183011 | Chernetidae | Pselaphochernes lacertosus (L. Koch, 1873) | ES | Zaragoza and Ruiz de la Cuesta Santiago (2017) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Judson (1979) |
| Chyromyidae | ||||
| Chyromya flava (Linnaeus, 1758) | Chernetidae | Lamprochernes cf. chyzeri (Tömösváry, 1883) | SK | Hetešová and Christophoryová (2022) |
| Clusiidae | ||||
| Clusiodes albimanus (Meigen, 1830) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | SK | Hetešová and Christophoryová (2022) |
| Culicidae | ||||
| Aedes sticticus (Meigen, 1838)12 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | CZ | Minář (1966) |
| Anopheles freeborni Aitken, 1939 | Cheiridiidae | Cheiridium Menge, 1855 | US-CA | Meyer et al. (1985) |
| Culex tarsalis Coquillett, 1896 | Cheiridiidae | Cheiridium Menge, 1855 | US-CA | Meyer et al. (1985) |
| Culicini [tribe] Meigen, 1818 | Larcidae | Larca lata (Hansen, 1885) | AT | Ressl (1965) |
| Not specified | Chernetidae | Parachernes (Parachernes) inpai Mahnert, 1979 | BR | Aguiar and Bührnheim (1998b) |
| Not specified | Larcidae | Larca chamberlini Benedict & Malcolm, 1978 | US-CA | Benedict and Malcolm (1977) |
| Dolichopodidae | ||||
| Dolichopus acuticornis Wiedemann, 181713 | Syarinidae | Syarinus strandi (Ellingsen, 1901)13 | FI | Kaisila (1949) |
| Drosophilidae | ||||
| Drosophila funebris (Fabricius, 1787) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Vachon (1940) |
| Drosophila hydei Sturtevant, 1921 | Chernetidae | Lamprochernes savignyi (Simon, 1881) | AU | Harvey (1987) |
| Drosophila melanogaster Meigen, 1830 | Chernetidae | Lamprochernes savignyi (Simon, 1881) | AU | Harvey (1987) |
| Drosophila aff. repleta Wollaston, 1858 | Chernetidae | Parachernes (P.) beieri Muchmore, 199914 | EC | Mahnert (1987) |
| Empididae | ||||
| Hilara Meigen, 1822 | Chernetidae | Allochernes peregrinus Lohmander, 1939 | SK | Christophoryová et al. (2017a) |
| Fanniidae | ||||
| Fannia canicularis (Linnaeus, 1761) | Chernetidae | Americhernes aff. incertus Mahnert, 1979 | BR | Lira et al. (2014) |
| Fannia canicularis (Linnaeus, 1761)*15 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*15 | GB-ENG | Macrae (1869) |
| Fannia canicularis (Linnaeus, 1761)16 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*16 | DE | Grimpe (1921) |
| Fannia canicularis (Linnaeus, 1761)17 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)17 | Not specified | Berland (1932) |
| Fannia canicularis (Linnaeus, 1761) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1978) |
| Fannia canicularis (Linnaeus, 1761) | Chernetidae | Lamprochernes savignyi (Simon, 1881) | AU | Harvey (1987) |
| Fannia canicularis (Linnaeus, 1761) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | SK | Present study |
| Fannia canicularis (Linnaeus, 1761) | Chernetidae | Sphenochernes camponoti (Beier, 1970) | BR | Lira and Tizo-Pedroso (2017) |
| Fannia pusio (Wiedemann, 1830) | Chernetidae | Sphenochernes camponoti (Beier, 1970) | BR | Lira and Tizo-Pedroso (2017) |
| Fannia yenhedi Albuquerque, 1957 | Chernetidae | Sphenochernes camponoti (Beier, 1970) | BR | Lira and Tizo-Pedroso (2017) |
| Heleomyzidae | ||||
| Amoebaleria defessa (Osten-Sacken, 1877) | Chernetidae | Chelodamus Chamberlin, 192518 | US | Muchmore (1971) |
| Amoebaleria Garrett, 192119 | Chernetidae | Hesperochernes tamiae Beier, 1930 | US | Muchmore (1971) |
| Scoliocentraa Loew, 1862 | Not specified | Not specified | N America | Marshall (2012) |
| Tephrochlamys rufiventris (Meigen, 1830)*20 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*20 | FR | Burr (1919) |
| Hybotidae | ||||
| Leptopeza flavipes (Meigen, 1820) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | SK | Červená et al. (2019) |
| Leptopeza flavipes (Meigen, 1820) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | AT | Červená et al. (2019) |
| Leptopeza flavipes (Meigen, 1820) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | AT | Červená et al. (2019) |
| Limoniidae | ||||
| Achyrolimonia decemmaculata (Loew, 1873)21 | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | GB-ENG | Hinton (1954) |
| Achyrolimonia decemmaculata (Loew, 1873)22 | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | GB-ENG | Jones (1978) |
| Achyrolimonia decemmaculata (Loew, 1873) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | ES | Mederos and Zaragoza (2017) |
| Limonia nubeculosa Meigen, 1804 | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | SI | Present study |
| Neolimonia dumetorum (Meigen, 1804) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | FR | Present study |
| Not specified | Cheliferidae | Dactylochelifer degeerii (C.L. Koch, 1835)23 | AT | Ressl (1995) |
| Not specified | Not specified | Not specified | NL | Jong et al. (2024) |
| Lonchaeidae | ||||
| Lonchaea carpathica Kovalev, 1974 | Chernetidae | Chernes cimicoides (Fabricius, 1793) | CZ | Christophoryová and Blažej (2025) |
| Lonchaea chorea (Fabricius, 1781) | Chernetidae | Lamprochernes chyzeri (Tömösváry, 1883) | GB-ENG | Jones (1978) |
| Lonchaea chorea (Fabricius, 1781) | Chernetidae | Lamprochernes cf. chyzeri (Tömösváry, 1883) | SK | Present study |
| Lonchaea chorea (Fabricius, 1781)24 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)24 | GB-ENG | Ed (1834) |
| Lonchaea chorea (Fabricius, 1781)25 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)25 | GB-ENG | Graham-Smith (1916) |
| Lonchaea chorea (Fabricius, 1781) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1978) |
| Lonchaea laticornis Meigen, 1826 | Cheliferidae | Rhacochelifer similis Beier, 1932 | TN | Vachon (1940) |
| Lonchaea Fallén, 1820 | Chernetidae | Lamprochernes cf. chyzeri (Tömösváry, 1883) | SK | Christophoryová et al. (2021) |
| Lonchaea Fallén, 1820 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1978) |
| Lonchaea Fallén, 1820 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Judson (1979) |
| Lonchaea Fallén, 1820 | Chernetidae | Lamprochernes savignyi (Simon, 1881)26 | GB-ENG | Jones (1978) |
| Micropezidae | ||||
| Grallipeza gracilis Hennig, 1934 | Not specified | Not specified | EC | Marshall (2012) |
| Rainieria calceata (Fallén, 1820) | Chernetidae | Lamprochernes chyzeri (Tömösváry, 1883) | CH | DeVore-Scribante (1999) |
| Scipopus Enderlein, 1922 | Chernetidae | Americhernes oblongus (Say, 1821) | PA | Martínez et al. (2020) |
| Not specified | Not specified | Not specified | CR | Szymkowiak et al. (2007) |
| Muscidae | ||||
| Hydrotaea ignava (Harris, 1780)27 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)27 | GB-ENG | Graham-Smith (1916) |
| Hydrotaea meteorica (Linnaeus, 1758)28 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)28 | GB-ENG | Anonymous (1834) |
| Hydrotaea similis Meade, 1887 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SK | Mašán and Krištofík (1992) |
| Musca autumnalis De Geer, 1776 | Chernetidae | Lamprochernes cf. chyzeri (Tömösváry, 1883) | SK | Present study |
| Musca autumnalis De Geer, 177629 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)29 | GB-ENG | Graham-Smith (1916) |
| Musca autumnalis De Geer, 1776 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1978) |
| Musca domestica Linnaeus, 1758 | Cheliferidae | Chelifer cancroides (Linnaeus, 1758) | Not specified | Anonymous (1879) |
| Musca domestica Linnaeus, 1758*30 | Cheliferidae | Chelifer cancroides (Linnaeus, 1758) | ES | Franganillo (1913) |
| Musca domestica Linnaeus, 1758 | Cheliferidae | Chelifer cancroides (Linnaeus, 1758) | Not specified | Beier (1951) |
| Musca domestica Linnaeus, 1758 | Cheliferidae | Chelifer cancroides (Linnaeus, 1758) | IE | Jones (1978) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Chernes cimicoides (Fabricius, 1793) | CZ | Stecker (1875) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Dinocheirus panzeri (C.L. Koch, 1836)31 | BE | Preudhomme de Borre (1873) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Dinocheirus panzeri (C.L. Koch, 1836)32 | AT | Ressl (1965) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes chyzeri (Tömösváry, 1883) | DK | Meinertz (1964) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes cf. chyzeri (Tömösváry, 1883) | SK | Christophoryová et al. (2011) |
| Musca domestica Linnaeus, 1758*33 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)33 | AT | Poda (1761) |
| Musca domestica Linnaeus, 1758*34 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*34 | GB-ENG | Adams (1787) |
| Musca domestica Linnaeus, 1758*35 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)35 | Europe | Hermann (1804) |
| Musca domestica Linnaeus, 1758*36 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*36 | GB-ENG | Leach (1817) |
| Musca domestica Linnaeus, 175837 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*37 | GB-ENG | Anonymous (1831, 1832) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)38 | GB-ENG | Anonymous (1834) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)38 | GB-ENG | Lukis (1834) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)38 | GB-ENG | Moore (1834) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*1 | GB-ENG | Clarke (1858) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*39 | GB-ENG | Stainton (1865) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)40 | GB-ENG | Stevens (1866) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)40 | GB-ENG | Stevens (1866) |
| Musca domestica Linnaeus, 1758*36 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*36 | GB-ENG | M’Intire (1869) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*41 | GB-ENG | Bisshopp (1871) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)42 | CH, DE, FR | Koch (1873) |
| Musca domestica Linnaeus, 1758*43 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)43 | GB-ENG | Anonymous (1875); Newman (1875) |
| Musca domestica Linnaeus, 1758*44 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)44 | GB-ENG | Dale (1878) |
| Musca domestica Linnaeus, 1758*45 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)45 | FR | Simon (1879) |
| Musca domestica Linnaeus, 1758*46 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)46 | FR | Courtois de Langlade (1883); Mégnin (1886) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Pickard-Cambridge (1885) |
| Musca domestica Linnaeus, 1758*48 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)48 | GB-ENG | Pickard-Cambridge (1892) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)38 | DE | Hess (1894) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)49 | CH | Müller and Schenkel (1895); Lessert (1911) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)38 | IT | Fenizia (1902) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*50 | GB-ENG | Hill (1905); Hickson (1905) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Pocock (1905) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)51 | CH | Frey-Gessner (1906); Lessert (1911) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Jackson (1907) |
| Musca domestica Linnaeus, 1758*45 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)45 | DE | Tullgren (1907) |
| Musca domestica Linnaeus, 1758*45 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)45 | DE | Ellingsen (1908) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Jackson (1908) |
| Musca domestica Linnaeus, 1758*48 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)48 | GB-SCT | Godfrey (1909) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Hewitt (1910, 1914) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)52 | CH | Lessert (1911) |
| Musca domestica Linnaeus, 1758*53 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)53 | IE 53 | Carpenter (1912) |
| Musca domestica Linnaeus, 1758*45 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)45 | GB-ENG | Ellingsen (1913) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Graham-Smith (1916) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Graham-Smith (1916) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Graham-Smith (1916) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Graham-Smith (1916) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)54 | ES | Navás (1918) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)55 | FR | Burr (1919) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*41 | DE | Grimpe (1921) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*41 | DE | Grimpe (1921) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*41 | DE | Grimpe (1921) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*41 | DE | Grimpe (1921) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*41 | DE | Grimpe (1921) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*41 | DE | Grimpe (1921) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*41 | DE | Grimpe (1921) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)54 | IT | Caporiacco (1923) |
| Musca domestica Linnaeus, 1758*45 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)45 | ES | Navás (1925) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)52 | IT | Beier (1929) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)56 | FR | Berland (1932) |
| Musca domestica Linnaeus, 1758*30 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Vachon (1934a) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Vachon (1934b) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)54 | former USSR | Redikorzev (1936) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | AT | Beier (1939) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SE | Lohmander (1939a) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SE | Lohmander (1939a) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | AT | Beier (1948) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | KG | Redikorzev (1949) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Vachon (1953) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Vachon (1954) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | AT | Ressl and Beier (1958) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | AT | Ressl (1965) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1970) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | AT | Ressl (1970) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1978) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Judson (1979) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DK | Andersen (1987) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DK | Andersen (1987) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DK | Andersen (1987) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DE | Carl (1994) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DE | Bellmann (1997) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SK | Christophoryová et al. (2017a) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes savignyi (Simon, 1881)57 | IE | Stephens (1910); Kew (1916) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes savignyi (Simon, 1881)58 | IE 58 | Kew (1916) |
| Musca domestica Linnaeus, 175859 | Chernetidae | Lamprochernes savignyi (Simon, 1881)59 | JP | Morikawa (1960) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes savignyi (Simon, 1881) | AU | Harvey (1987) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes savignyi (Simon, 1881) | AU | Harvey (1987) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Lamprochernes Tömösváry, 188360 | US | Muchmore (1971) |
| Musca domestica Linnaeus, 1758 | Neobisiidae | Neobisium (Neobisium) carcinoides (Hermann, 1804)61 | GB-ENG | Jackson (1907) |
| Musca domestica Linnaeus, 1758 | Neobisiidae | Neobisium (N.) sylvaticum (C.L. Koch, 1835) | Europe | Poinar et al. (1998) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Pselaphochernes lacertosus (L. Koch, 1873) | ES | Domínguez et al. (2008) |
| Musca domestica Linnaeus, 1758 | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | GB-ENG | Jones (1978) |
| Musca domestica Linnaeus, 1758 | Not specified | Not specified62 | US-PA | Leidy (1877); Muchmore (1971) |
| Musca domestica Linnaeus, 1758 | Not specified | Not specified63 | N America | Webster (1897); Muchmore (1971) |
| Musca domestica Linnaeus, 1758 | Not specified | Not specified | NZ | Edwards (1955) |
| Musca Linnaeus, 1758 | Chernetidae | Lamprochernes savignyi (Simon, 1881) | ZA | Beier (1953) |
| Musca? Linnaeus, 1758 | Chernetidae | Pselaphochernes Beier, 1932 | US | Muchmore (1971) |
| Muscina stabulans (Fallén, 1817) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | FR | Vachon (1953) |
| Not specified | Chernetidae | Dinocheirus serratus (Moles, 1914)64 | US-CA | Chamberlin (1952) |
| Not specified65 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)65 | GH 65 | Ellingsen (1913) |
| Stomoxys calcitrans (Linnaeus, 1758) | Chernetidae | Lamprochernes cf. chyzeri (Tömösváry, 1883) | SK | Christophoryová et al. (2017a) |
| Stomoxys calcitrans (Linnaeus, 1758)66 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)66 | GB-ENG | Graham-Smith (1916) |
| Stomoxys calcitrans (Linnaeus, 1758) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Vachon (1940) |
| Stomoxys calcitrans (Linnaeus, 1758) | Chernetidae | Lamprochernes nodosus (Schrank, 1803)54 | BE | Leclercq (1945) |
| Stomoxys calcitrans (Linnaeus, 1758) | Chernetidae | Lamprochernes savignyi (Simon, 1881)67 | IE | Kew (1916) |
| Stomoxys calcitrans (Linnaeus, 1758) | Chernetidae | Lamprochernes savignyi (Simon, 1881)26 | GB-ENG | Jones (1978) |
| Stomoxys Geoffroy, 1762 | Chernetidae | Lamprochernes savignyi (Simon, 1881) | ZA | Beier (1964) |
| Stomoxys Geoffroy, 1762 | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804)68 | GB-ENG | Graham-Smith (1916) |
| Neriidae | ||||
| Odontoloxozus longicornis (Coquillett, 1904) | Chernetidae | Dinocheirus arizonensis (Banks, 1901) | US-AZ | Zeh and Zeh (1992b) 69 |
| Pantophthalmidae | ||||
| Pantophthalmus tabaninus Thunberg, 1819 | Chernetidae | Semeiochernes armiger (Balzan, 1892) | PA | Zeh and Zeh (1992a) |
| Pantophthalmus tabaninus Thunberg, 1819 | Chernetidae | Semeiochernes armiger (Balzan, 1892)70 | BR | Santos et al. (2005) |
| Not specified | Chernetidae | Semeiochernes armiger (Balzan, 1892)71 | BR | Mahnert (1987) |
| Sarcophagidae | ||||
| Not specified | Chernetidae | Anthrenochernes stellae Lohmander, 1939 | SE | Gärdenfors and Wilander (1995) 72 |
| Scatopsidae | ||||
| Scatopse notata (Linnaeus, 1758) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | FR | Present study |
| Scenopinidae | ||||
| Scenopinus fenestralis (Linnaeus, 1758)73 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Vachon (1940) |
| Sciomyzidae | ||||
| Pherbellia annulipes (Zetterstedt, 1846) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | SK | Present study |
| Sphaeroceridae | ||||
| Limosina silvatica (Meigen, 1830)74 | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | GB-ENG | Cuthbertson (1982) |
| Not specified | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | Europe | Weygoldt (1966, 1969) |
| Stratiomyidae | ||||
| Artemita amenides (Walker, 1849) | Chernetidae | Parachernes (P.) setiger Mahnert, 1979 | BR | Guimarães et al. (2025) |
| Dicranophora bispinosa (Wiedemann, 1830)75 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*75 | FR | Vachon (1940) |
| Sargus bipunctatus (Scopoli, 1763) | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | DE | Ziegler (2004) |
| Sargus iridatus (Scopoli, 1763)76 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1978) |
| Sargus iridatus (Scopoli, 1763) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Judson (1979) |
| Syrphidae | ||||
| Brachyopa bicolor (Fallén, 1817) | Chernetidae | Chernes hahnii (C.L. Koch, 1839) | AT | Červená et al. (2019) |
| Brachypalpus laphriformis (Fallén, 1816) | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*1 | DE | Gerstaecker (1862) |
| Eristalis arbustorum (Linnaeus, 1758) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1978) |
| Myathropa florea (Linnaeus, 1758) | Chernetidae | Anthrenochernes stellae Lohmander, 1939 | DE | Ssymank and Muster (2010) |
| Myolepta dubia (Fabricius, 1805) | Chernetidae | Pselaphochernes lacertosus (L. Koch, 1873) | ES | Ricarte et al. (2016) |
| Volucella zonaria (Poda, 1761) | Chernetidae | Lamprochernes nodosus (Schrank, 1803)77 | FR | Berland (1932); Vachon (1940) |
| Not specified78 | Chernetidae | Lamprochernes savignyi (Simon, 1881) | ZA | Beier (1958) |
| Tabanidae | ||||
| Phorcotabanus cinereus (Wiedemann, 1821) | Chernetidae | Parachernes (P.) inpai Mahnert, 1979 | BR | Aguiar and Bührnheim (1998b) |
| Phorcotabanus cinereus (Wiedemann, 1821) | Chernetidae | Parachernes (P.) melanopygus Beier, 1959 | BR | Aguiar and Bührnheim (1998b) |
| Phorcotabanus cinereus (Wiedemann, 1821) | Chernetidae | Parachernes (P.) plumosus (With, 1908) | BR | Aguiar and Bührnheim (1998b) |
| Stenotabanus cretatus Fairchild, 1961 | Chernetidae | Parachernes (P.) melanopygus Beier, 1959 | BR | Aguiar and Bührnheim (1998b) |
| Stenotabanus cretatus Fairchild, 1961 | Chernetidae | Parachernes (P.) plumosus (With, 1908) | BR | Aguiar and Bührnheim (1998b) |
| Tabanus amapaensis Fairchild 1961 | Chernetidae | Parachernes (P.) melanopygus Beier, 1959 | BR | Aguiar and Bührnheim (1998b) |
| Tabanus occidentalis Linnaeus, 1758 | Chernetidae | Parachernes (P.) inpai Mahnert, 1979 | BR | Aguiar and Bührnheim (1998b) |
| Tabanus trivittatus Fabricius, 1805 | Chernetidae | Parachernes (P.) plumosus (With, 1908) | BR | Aguiar and Bührnheim (1998b) |
| Tabanus Linnaeus, 1758 | Not specified | Not specified | AR | Berg (1893) |
| Not specified | Chernetidae | Parachernes (P.) argentatopunctatus (Ellingsen, 1910) | GY | Beier (1948) |
| Tachinidae | ||||
| Exorista larvarum (Linnaeus, 1758)79 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)79 | GB-ENG | Anonymous (1834) |
| Zelia vertebrata (Say, 1829) | Not specified | Not specified | N America | Marshall (2012) |
| Not specified80 | Chernetidae | Hesperochernes pallipes (Banks, 1893)80 | US-CO | Banks (1895) |
| Tanypezidae | ||||
| Neotanypeza Hendel, 1903 | Not specified | Not specified | CR | Marshall (2012) |
| Tipulidae | ||||
| Ctenophora pectinicornis (Linnaeus, 1758) | Chernetidae | Anthrenochernes stellae Lohmander, 1939 | SE | Gärdenfors and Wilander (1995) |
| Ctenophora pectinicornis (Linnaeus, 1758) | Chernetidae | Lamprochernes nodosus (Schrank, 1803)81 | DE | Wagner (1892); Berland (1932) |
| Ctenophora pectinicornis (Linnaeus, 1758) | Not specified | Not specified | FR | Vachon (1953) |
| Dictenidia bimaculata (Linnaeus, 1761) | Chernetidae | Chernes cimicoides (Fabricius, 1793) | GB-ENG | Bloxham and Smart (2001) |
| Dictenidia bimaculata (Linnaeus, 1761) | Chernetidae | Chernes cimicoides (Fabricius, 1793) | CZ | Christophoryová and Krásenský (2022) |
| Tipula pabulina Meigen, 1818 | Chernetidae | Allochernes peregrinus Lohmander, 1939 | CH | DeVore-Scribante (1999) |
| Tipula unicincta Doane, 1901 | Syarinidae | Syarinus obscurus (Banks, 1893) | US | Muchmore (1971) |
| Tipula vernalis Meigen, 1804 | Cheliferidae | Dactylochelifer degeerii (C.L. Koch, 1835) | FR | Present study |
| Tipula Linnaeus, 1758 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)82 | GB-ENG | Jenyns (1846) |
| Tipula Linnaeus, 1758 | Syarinidae | Syarinus obscurus (Banks, 1893)83 | US | Hagen (1879); Muchmore (1971) |
| Zelandotipula Alexander, 1922 | Chernetidae | Not specified | BR | Matthiesen and Hahn (1981) |
| Not specified | Cheliferidae | Dactylochelifer degeerii (C.L. Koch, 1835)84 | AT | Ressl (1995) |
| Not specified | Chernetidae | Hesperochernes molestus Hoff, 1956 | US-TX | Muchmore (1992) |
| Ulidiidae | ||||
| Ceroxys urticae (Linnaeus, 1758) | Chernetidae | Lamprochernes cf. chyzeri (Tömösváry, 1883) | CZ | Present study |
| Physiphora alceae (Preyssler, 1791)85 | Chernetidae | Lamprochernes muscivorus Redikorzev, 1949 | TM | Redikorzev (1949) |
| Physiphora alceae (Preyssler, 1791)85 | Chernetidae | Lamprochernes muscivorus Redikorzev, 1949 | UZ | Redikorzev (1949) |
| Physiphora alceae (Preyssler, 1791)85 | Chernetidae | Lamprochernes muscivorus Redikorzev, 1949 | UZ | Redikorzev (1949) |
| Physiphora alceae (Preyssler, 1791)86 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)86 | HU | Loew (1845) |
| Physiphora alceae (Preyssler, 1791)87 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)87 | AT | Schiner (1872) |
| Physiphora alceae (Preyssler, 1791)88 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | AT | Ressl (1965) |
| Physiphora alceae (Preyssler, 1791)88 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | AT | Ressl (1970) |
| Physiphora alceae (Preyssler, 1791)88 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Clements (1987) |
| Physiphora alceae (Preyssler, 1791)88 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | RO | Georgescu and Capuse (1996) |
| Physiphora alceae (Preyssler, 1791) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SK | Christophoryová et al. (2018) |
| Physiphora alceae (Preyssler, 1791) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SK | Christophoryová et al. (2018) |
| Physiphora alceae (Preyssler, 1791) | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SK | Červená et al. (2019) |
| Physiphora alceae (Preyssler, 1791) | Not specified | Not specified89 | FR | Livory (2021) |
| Physiphora clausa Macquart, 184390 | Chernetidae | Lamprochernes savignyi (Simon, 1881) | MU | Beier (1953) |
| Physiphora Fallén, 1810 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Present study |
| Ulidia erythrophthalma Meigen, 1826 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)91 | AT | Löw (1866) |
| Xylophagidae | ||||
| Xylophagus ater Meigen, 1804 | Chernetidae | Chernes cimicoides (Fabricius, 1793) | SK | Present study |
| Diptera | ||||
| Not specified | Cheliferidae | Chelifer cancroides (Linnaeus, 1758) | FR | Moniez (1893) |
| Not specified | Cheliferidae | Chelifer loewi Hagen, 187992 | PA | Hagen (1879) |
| Not specified | Chernetidae | Chernes sanborni Hagen, 186893 | N America | Hagen (1867, 1868); Muchmore (1971) |
| Not specified | Chernetidae | Chernes sanborni Hagen, 186894 | US-MA | Hagen (1879) |
| Not specified | Chthoniidae | Ephippiochthonius tetrachelatus (Preyssler, 1790) | HU | Novák (2024) |
| Not specified | Chernetidae | Hesperochernes Chamberlin, 1924 | US | Muchmore (1971) 95 |
| Not specified | Chernetidae | Lamprochernes chyzeri (Tömösváry, 1883) | GB-ENG | Jones (1978) |
| Not specified | Chernetidae | Lamprochernes chyzeri (Tömösváry, 1883) | AT | Ressl (1995) |
| Not specified | Chernetidae | Lamprochernes chyzeri (Tömösváry, 1883) | CH | DeVore-Scribante (1999) |
| Not specified | Chernetidae | Lamprochernes chyzeri (Tömösváry, 1883), Lamprochernes nodosus (Schrank, 1803) | DE | Drogla and Lippold (2004) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803)82 | GB-ENG | Jenyns (1846) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803)*1 | AT | Löw (1867) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803)54 | FR | Moniez (1893) |
| Not specified96 | Chernetidae | Lamprochernes nodosus (Schrank, 1803)96 | GB-ENG | Kew (1901) 96 |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-ENG | Kew (1904) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | GB-SCT | Godfrey (1907, 1908) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | SE | Tullgren (1911); Lohmander (1939a) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | ES | Navás (1918) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803)47 | ZA | Godfrey (1921) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | FR | Vachon (1954) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DE | Zielke (1969) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DE | Zielke (1969) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DE | Zielke (1969) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Jones (1978) |
| Not specified97 | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | GB-ENG | Judson (1987) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | CH | DeVore-Scribante (1999) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | DE | Drogla and Lippold (2004) |
| Not specified | Chernetidae | Lamprochernes nodosus (Schrank, 1803) | IR | Nassirkhani and Shoushtari (2016) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881)98 | GB-ENG | Ellingsen (1907) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881)98 | GB-ENG | Ellingsen (1907) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881)99 | GB-ENG | Kew (1911) 99 |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881)58 | IE 58 | Kew (1916) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881) | SD | Beier (1946); Beier (1948)100 |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881) | SD | Beier (1953) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881) | ZA | Beier (1958) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881) | IL | Beier (1963) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881) | ZA | Beier (1964) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881) | NZ | Beier (1969) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881)26 | GB-ENG | Judson (1979) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881) | IE | Harvey (1987) |
| Not specified | Chernetidae | Lamprochernes savignyi (Simon, 1881) | IL | Warburg et al. (2023a) |
| Not specified | Chernetidae | Parachernes (P.) melanopygus Beier, 1959 | BR | Aguiar and Bührnheim (1998b) |
| Not specified | Chernetidae | Phymatochernes crassimanus Mahnert, 1979 | BR | Aguiar and Bührnheim (1998b) |
| Not specified | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804)101 | GB-ENG | Kew (1929) |
| Not specified | Chernetidae | Pselaphochernes scorpioides (Hermann, 1804) | DE | Drogla and Lippold (2004) |
| Not specified | Not specified | Not specified63 | US | Knab (1897); Muchmore (1971) |
| Not specified | Not specified | Not specified | NZ | Edwards (1955) |
Numbers in superscript refer to the Notes, and the asterisk (*) indicates the interpretation of Beier (1948), see Results. Abbreviations: AR – Argentina; AT – Austria; AU – Australia; BE – Belgium; BR – Brazil; CH – Switzerland; CR – Costa Rica; CZ – Czech Republic; DE – Germany; DK – Denmark; EC – Ecuador; ES – Spain; FI – Finland; FR – France; GB-ENG – United Kingdom, England; GB-SCT – United Kingdom, Scotland; GH – Ghana; GY – Guyana; HU – Hungary; IE – Ireland; IL – Israel; IR – Iran; IT – Italy; JP – Japan; KG – Kyrgyzstan; MU – Mauritius; N America – North America; NL – Netherlands; NZ – New Zealand; PA – Panama; RO – Romania; SD – Sudan; SE – Sweden; SI – Slovenia; SK – Slovakia; TM – Turkmenistan; TN – Tunisia; US – United States of America; US-AZ – United States of America, Arizona; US-CA – United States of America, California; US-CO – United States of America, Colorado; US-MA – United States of America, Massachusetts; US-PA – United States of America, Pennsylvania; US-TX – United States of America, Texas; UZ – Uzbekistan; ZA – South Africa.
1. Czechia • 1 ♀ of Lamprochernes cf. chyzeri attached to femur III of 1 ♀ of Ceroxys urticae (Ulidiidae); Lysá nad Labem, Hrabanovská černava National Nature Monument; 50.2188°N, 14.8358°E; 186 m a.s.l.; 13 Jun.–8 Jul. 2024; Jindřich Roháček leg.; wetland, lowland fen, Malaise trap; host Jindřich Roháček det., deposited in the Silesian Museum, Opava, Czechia; FO 40/6 (Fig. 2A).
Figure 2.
Cases of phoresy from Czechia, Slovakia, and Slovenia. A. Ceroxys urticae and Lamprochernes cf. chyzeri (locality 1); B. Pherbellia annulipes and Pselaphochernes scorpioides (locality 6); C. Fannia canicularis and P. scorpioides (locality 7); D. Xylophagus ater and Chernes cimicoides (locality 8); E. Musca autumnalis and L. cf. chyzeri (locality 9); F. Lonchaea chorea and L. cf. chyzeri (locality 10); G, H. Limonia nubeculosa and P. scorpioides (locality 11). Scale bars: 1 mm.
2. France • 1 ♀ of Pselaphochernes scorpioides attached to the base of femur III of 1 ♂ of Scatopse notata (Scatopsidae); Bénouville; 49.2444, -0.2865; 23 m a.s.l.; 15 Oct. 2023; Vincent Mathy leg.; family house garden, compost bin walls, individual sampling; host Vincent Mathy det.; FO 41/6 (Fig. 3A, B).
Figure 3.
Cases of phoresy from France. A, B. Scatopse notata and Pselaphochernes scorpioides (locality 2); C, D. Physiphora sp. and Lamprochernes nodosus (locality 3); E, F. Neolimonia dumetorum and P. scorpioides (locality 4); G, H. Tipula vernalis and Dactylochelifer degeerii (locality 5). Scale bars: 1 mm.
3. France • 1 ♀ of Lamprochernes nodosus phoretic on the leg of 1 ♀ of Physiphora sp. (Ulidiidae); Mauges-sur-Loire; 47.3311, -1.0361; 56 m a.s.l.; 29 Aug. 2016; Olivier Gabory leg.; inside a house, individual sampling; host Jindřich Roháček det.; FO 42/6 (Fig. 3C, D).
4. France • 1 ♀ of P. scorpioides attached to trochanter I of 1 ♂ of Neolimonia dumetorum (Limoniidae); Vernet-les-Bains; 42.5437°N, 2.3842°E; 690 m a.s.l.; 22 May 2023; Julien Tchilinguirian leg.; pine forest, camping ground, individual sampling; host Levente-Péter Kolcsár det.; FO 43/6 (Fig. 3E, F).
5. France • 1 ♀ of Dactylochelifer degeerii phoretic on the leg of 1 ♀ of Tipula vernalis (Tipulidae); Vimont; 49.1490, -0.1709; 5 m a.s.l.; 9 May 2021; Antoine Racine leg.; swamp area, individual sampling; host Antoine Racine det.; FO 44/6 (Fig. 3G, H).
6. Slovakia • 1 ♀ of P. scorpioides attached between trochanter and femur III of 1 ♀ of Pherbellia annulipes (Sciomyzidae); Bratislava, Botanical Garden; 48.1465°N, 17.0753°E; 144 m a.s.l.; 3–12 Jul. 2024; Simona Chamulová, Jana Christophoryová leg.; botanical garden, Malaise trap; host Jindřich Roháček det.; FO 45/6 (Fig. 2B).
7. Slovakia • 1 ♀ of P. scorpioides attached to the base of femur III of 1 ♂ of Fannia canicularis (Fanniidae); Bratislava, Police Academy; 48.2030°N, 17.1720°E; 136 m a.s.l.; 17 Apr. 2024; Lucia Švecová leg.; grassy area, pyramid trap; host Miroslav Barták det.; FO 46/6 (Fig. 2C).
8. Slovakia • 1 ♀ of Chernes cimicoides attached between trochanter and femur III of 1 ♂ of Xylophagus ater (Xylophagidae); Cerová vrchovina Highlands, Fenek Protected Area; 48.1847°N, 20.0416°E; 240 m a.s.l.; 3–25 May 2022; Jindřich Roháček, Jan Ševčík, Michal Tkoč leg.; alder forest, Malaise trap; host Jindřich Roháček det., deposited in the Silesian Museum, Opava, Czechia; FO 47/6 (Fig. 2D).
9. Slovakia • 1 ♂ of L. cf. chyzeri attached between trochanter and femur II of 1 ♂ of Musca autumnalis (Muscidae); Ivanka pri Dunaji, Bažantnica; 48.1942°N, 17.2675°E; 130 m a.s.l.; 12 May 2022; Ján Samay, Ľubomír Vidlička leg.; overgrown meadow among trees, Malaise trap; host Erikas Lutovinovas det.; FO 48/6 (Fig. 2E).
10. Slovakia • 2 ♀♀, 1 ♂ of L. cf. chyzeri attached to femora III of 1 ♀ of Lonchaea chorea (Lonchaeidae); Kežmarok, Čajka Ranch; 49.1587°N, 20.4528°E; 630 m a.s.l.; 3–5 Aug. 2022; Ema Hetešová leg.; ranch area, Malaise trap; host Ian MacGowan det.; FO 49/6 (Fig. 2F).
11. Slovenia • 1 ♀ of P. scorpioides attached to the leg of 1 ♂ of Limonia nubeculosa (Limoniidae); Otoška Jama, Veliki Otok; 45.7918°N, 14.1970°E; 543 m a.s.l.; 1 Aug. 2024; Pavol Purgat, Lucia Švecová leg.; cave entrance, individual sampling; host Levente-Péter Kolcsár det.; FO 50/6 (Fig. 2G, H).
Revision of published data on phoresy from Slovakia (Table 1)
Christophoryová et al. (2017a): In Allochernes peregrinus, the host had originally been reported only as Diptera. In the present study, a female of Hilara (Empididae; det. Miroslav Barták) was identified based on photographic documentation. The specimen is deposited in the collection of Dr Viera Stloukalová, but could not be located. In addition, three further phoretic records had originally been identified only to the genus Lamprochernes with their hosts recorded merely as Diptera. In the July sample, the host was identified as a female of Stomoxys calcitrans (Muscidae; det. Miroslav Barták), and the phoront as Lamprochernes cf. chyzeri (det. J. Christophoryová). In the August sample, the pseudoscorpions were identified as L. nodosus (det. J. Christophoryová), with the host identified as a male of Musca domestica (Muscidae; det. Miroslav Barták).
Christophoryová et al. (2018): The host originally identified only to the superfamily Muscoidea was reidentified as Delia platura (Anthomyiidae; det. Miroslav Barták, Samuel Krčmárik).
Červená et al. (2019): The host previously assigned to Empididae was reidentified as a female of Leptopeza flavipes (Hybotidae; det. Miroslav Barták, Jindřich Roháček).
Christophoryová et al. (2021): The host was lost during examination by a specialist and could therefore be identified only to the genus level based on photographic documentation.
Hetešová and Christophoryová (2022): The host belonging to Chyromyidae was identified as a female of Chyromya flava (det. Jindřich Roháček).
Published data on the phoresy of Pseudoscorpiones and Diptera worldwide
Altogether, 172 published records as well as new records (marked as “Present study”) documenting phoretic associations between pseudoscorpions and dipterans, spanning the period from 1761 to 2025, were compiled and evaluated (Table 1).
The present study expands the original dataset of Poinar et al. (1998) by incorporating 100 additional relevant publications published up to and including the year 1998 (Adams 1787; Hermann 1804; Leach 1817; Kirby and Spence 1826; Anonymous 1831, 1832, 1834, 1875, 1879; Ed 1834; Lukis 1834; Moore 1834; Loew 1845; Jenyns 1846; Clarke 1858; Stainton 1865; Stevens 1866; Hagen 1867, 1868; Löw 1867; M’lntire 1869; Bisshopp 1871; Koch 1873; Newman 1875; Stecker 1875; Leidy 1877; Dale 1878; Simon 1879; Courtois de Langlade 1883; Pickard-Cambridge 1885, 1892; Mégnin 1886; Moniez 1893; Hess 1894; Müller and Schenkel 1895; Knab 1897; Kew 1901, 1904, 1911, 1916, 1929; Fenizia 1902; Hickson 1905; Hill 1905; Pocock 1905; Frey-Gessner 1906; Ellingsen 1907, 1908, 1913; Godfrey 1907, 1908, 1909, 1921; Jackson 1907, 1908; Tullgren 1907, 1911; Hewitt 1910, 1914; Stephens 1910; Lessert 1911; Carpenter 1912; Franganillo 1913; Navás 1918, 1925; Grimpe 1921; Caporiacco 1923; Beier 1929, 1939, 1946, 1951, 1953, 1958, 1963, 1964, 1969; Vachon 1934a, 1934b, 1953, 1954; Lohmander 1939a; Leclercq 1945; Kaisila 1949; Redikorzev 1949; Hinton 1954; Edwards 1955; Ressl and Beier 1958; Meinertz 1964; Ressl 1965, 1970, 1995; Benedict and Malcolm 1977; Judson 1979, 1987; Meyer et al. 1985; Andersen 1987; Harvey 1987; Muchmore 1992; Georgescu and Capuse 1996; Bellmann 1997).
In subsequent years, additional data on phoresy became available, and records published from 1998 onwards were compiled in the present study, resulting in the inclusion of 31 further publications (Aguiar and Bührnheim 1998b; DeVore-Scribante 1999; Bloxham and Smart 2001; Castillo-Miralbes 2002; Drogla and Lippold 2004; Ziegler 2004; Santos et al. 2005; Szymkowiak et al. 2007; Domínguez et al. 2008; Ssymank and Muster 2010; Christophoryová et al. 2011, 2017a, 2018, 2021; Marshall 2012; Lira et al. 2014; Nassirkhani and Shoushtari 2016; Ricarte et al. 2016; Lira and Tizo-Pedroso 2017; Mederos and Zaragoza 2017; Zaragoza and Ruiz de la Cuesta Santiago 2017; Červená et al. 2019; Martínez et al. 2020; Livory 2021; Christophoryová and Krásenský 2022; Hetešová and Christophoryová 2022; Warburg et al. 2023a; Jong et al. 2024; Novák 2024; Christophoryová and Blažej 2025; Guimarães et al. 2025).
Records marked with an asterisk (*) in Table 1 indicate cases in which host or phoront identifications follow the interpretation of Beier (1948), representing the most recent traceable taxonomic treatment of the originally cited material; these identifications have not been confirmed in WPC (2026). Missing data from published records are marked as “Not specified” in Table 1. Table 1 includes numbered Notes (1–101) documenting revisions and corrections to published records, including changes in host and phoront names, updates to family assignments, spelling errors in original sources (marked as [sic]), additions of data based on photographic documentation, correction of erroneous information, adjustments of country records, identification of duplicate data, and related amendments.
Hosts illustrated in Bellmann (1997) and Ziegler (2004) were identified from photographs as Musca domestica and Sargus bipunctatus, respectively, by a dipteran specialist (det. Alexssandro Camargo). The host from Szymkowiak et al. (2007) was identified only on the family level as Micropezidae (det. Nathalie Fial, Nikolaus Szucsich).
Reassessment of records summarised by Poinar et al. (1998)
The dataset compiled by Poinar et al. (1998), representing the most comprehensive synthesis of pseudoscorpion–dipteran phoresy at the time, was critically re-evaluated in the present study. Although several of the following publications were included in Poinar et al. (1998), not all available data on phoresy were extracted, and additional records were identified through re-examination of the original publications:
Hagen (1879): records of Syarinus obscurus (reported as Obisium) associated with Tipula; and Chernes sanborni associated with Diptera.
Graham-Smith (1916): records of Lamprochernes nodosus associated with Musca domestica.
Burr (1919): record of L. nodosus associated with M. domestica.
Berland (1932): records of L. nodosus associated with M. domestica and Fannia canicularis.
Vachon (1940): records of L. nodosus associated with Stomoxys calcitrans.
Muchmore (1971): Two records of Hesperochernes associated with Diptera.
Jones (1978): records of Chelifer cancroides associated with M. domestica; Lamprochernes chyzeri associated with Diptera; and L. nodosus associated with F. canicularis, M. domestica, Lonchaea, and Diptera.
Mahnert (1987): record of Parachernes beieri associated with Drosophila aff. repleta.
Addendum in Gärdenfors and Wilander (1995): record of Anthrenochernes stellae associated with Sarcophagidae.
The following records compiled in Poinar et al. (1998) were found to contain inaccuracies when re-examined against the original sources:
Data on Lamprochernes nodosus phoretic on Stomoxys calcitrans were attributed to Lukis (1831) and Schiner (1872). However, Lukis (1831) represents only a response to correspondence from Anonymous (1831) and provides no original data. Moreover, Schiner (1872) reported the host only as Chloria demandata (currently Physiphora alceae). The host S. calcitrans listed in Poinar et al. (1998) is therefore not supported by the original sources.
Records from Zielke (1969) and Macrae (1869) were cited as L. nodosus on Fannia canicularis. The record from Macrae (1869) is supported, whereas Zielke (1969) indicates different associations (see Table 1).
The phoront in Webster (1897) was listed as Chernes sanborni on Musca domestica. However, according to Muchmore (1971), cited therein, this identification is not supported; the phoront is therefore treated as not specified.
The record of Lamprochernes savignyi on Tabanidae is attributed to Ellingsen (1913); however, this association could not be verified in the original paper.
The record attributed to Vachon (1954) for L. nodosus on Drosophila funebris is not supported by the original source (here treated as Vachon 1954), in which this host association is not reported.
Data on L. savignyi from New Zealand, reported as phoretic on unidentified Tipulidae and attributed to Muchmore (1971), could not be verified in the original source.
Data on the phoresy of L. nodosus on Diptera attributed to Redikorzev (1936) are imprecise. The original publication contains only an illustration depicting L. nodosus phoretic on M. domestica from the former USSR and does not provide supporting records from Asia.
Lamprochernes nodosus on Diptera from the Pacific was cited based on Guilmette Jr et al. (1970). However, the original material was obtained using a suction trap, and no host taxon was specified; consequently, this record was not considered evidence of phoresy on Diptera in the present study.
The phoresy of Chelifer cancroides on M. domestica in Europe was attributed to Nonidez (1917). However, this does not represent original data from Nonidez (1917) but a citation from Franganillo (1913).
The record from Donovan (1797) appears twice, under C. cancroides and L. nodosus, which is most likely an error (see Note 7 for details).
The host family cited from Löw (1866) and Clements (1987) is given as Otitidae; this family is currently merged with Ulidiidae.
The host cited from Macrae (1869) is misspelled as Fannia canalicularis [sic] (should be Fannia canicularis), and the family is given as Muscidae, whereas it is currently treated as Fanniidae.
The host Sargus iridatus cited from Jones (1978) is placed in Syrphidae, whereas it correctly belongs to Stratiomyidae.
The host cited from Jones (1978) is given as the subgenus Achyrolimonia within the genus Limonia; it is currently treated as the genus Achyrolimonia.
The host Hydrotaea ignava cited from Graham-Smith (1916) is listed under its former name and family (see Note 27 for details).
The pseudoscorpion cited from Hagen (1879) is given as Lamprochernes loewi (Hagen, 1879), but this name should be treated as Chelifer loewi, a nomen nudum.
The host cited from Burr (1919) is given as Tephrochlamis [sic] canescens (should be Tephrochlamys canescens (Meigen, 1830)).
The host genus cited from Zeh and Zeh (1992c) as Odontolorozuz [sic] should be corrected to Odontoloxozus Enderlein, 1922 (treated here as Zeh and Zeh 1992b).
The pseudoscorpion cited from Jones (1978) as Lamprochernes scorpioides represents an incorrect name combination and should be L. chyzeri.
The first comprehensive synthesis of published data on pseudoscorpion phoresy was provided by Beier (1948), who compiled numerous published records and added only three new observations (see Table 1). Poinar et al. (1998) aimed to include these new data in their synthesis; however, re-examination shows that some records treated as new in Poinar et al. (1998) are based on secondary citations of earlier publications or require correction. These cases are outlined below:
The record of Lustrochernes argentinus (Thorell, 1877) on Lucilia Robineau-Desvoidy, 1830 from South America, attributed to Beier (1948), appears to be based on a misinterpretation of the original source. In Beier (1948: 452), the host is Lucilius Kuwert, 1891 (= Passalus Fabricius, 1792; Coleoptera).
Data on Allochernes wideri (currently Lamprochernes nodosus) phoretic on Ulidia erythrophthalma were treated as new data from Beier (1948). However, in Beier (1948), this record is cited from Löw (1866) and therefore does not represent a new observation.
Data on Syarinus obscurus phoretic on Tipula from North America were treated as new data from Beier (1948). However, in Beier (1948), this record is cited from Hagen (1879), with additional data later provided by Muchmore (1971).
Records attributed to Beier (1948) list L. nodosus and Chelifer cancroides phoretic on Musca meteorica (currently Hydrotaea meteorica) and on Tachina larvarum (currently Exorista larvarum). According to the WPC (2026), these records should be referred exclusively to L. nodosus, as C. cancroides represents a misidentification. Furthermore, these data are not original to Beier (1948) but were cited therein from Anonymous (1834).
Data from Beier (1948) on L. savignyi from Africa (Egypt, Sudan) were supplemented with the host Eutrichota apicalis (reported as Pegomya apicalis). However, in the original source (Beier 1948), the host is mentioned only as Diptera. In addition, Europe is listed as a region for this phoretic association, although the original publication does not support this.
Data on L. nodosus phoretic on M. domestica from Austria reported by Beier (1948) were not included.
Two species (L. nodosus and Chernes cimicoides) are mentioned with the host Ctenophora pectinicornis, attributed to Wagner (1892). However, only data referring to L. nodosus should be considered (see Note 81 for details).
Notes
1 Pseudoscorpion as Chelifer Geoffroy, 1762; in Beier (1948), as Lamprochernes nodosus.
2 Host as Hylemyia floralis (Fallén, 1824).
3 Host as Pegomya [sic] affinis Stein, 1898 (should be Pegomyia affinis).
4 Host as Pegomya [sic] apicalis (Stein, 1898) (should be Pegomyia apicalis).
5 Originally as “Schmeißfliege” and “Bücherskorpion” = Calliphoridae and Chelifer cancroides. In Beier (1948), the host is specified as Calliphora erythrocephala (Meigen, 1826), and the pseudoscorpion is redetermined as Lamprochernes nodosus.
6 Host originally reported as flies; one specimen from Mr Campbell was caught on the leg of a blow-fly, on a window in October 1887 (Kew 1901); the host was later specified as Calliphora erythrocephala (Beier 1948). Pseudoscorpion as Chernes nodosus Schrank, 1803.
7 Host as Musca vomitoria Linnaeus, 1758; pseudoscorpion as Phalangium crancroides [sic] (should be Phalangium cancroides (Linnaeus, 1758)). In Beier (1948), the record is listed as L. nodosus and Chelifer cancroides, with hosts given as Musca domestica and M. vomitoria. Due to these inconsistencies, only the original combination (C. cancroides on M. vomitoria) was used.
8 Host originally as M. vomitoria and pseudoscorpion as C. cancroides. In Beier (1948), the host is specified as Calliphora vomitoria, and the pseudoscorpion is redetermined as L. nodosus.
9 Host only as Calliphoridae; specified as Calliphora in Beier (1948).
10 Genus of the host as Lucia [sic] (should be Lucilia).
11 Host family as Rhinophoridae, currently placed in Calliphoridae.
12 Genus of the host as Aëdes (now Aedes).
13 Host as Dolichopus acutirostris [sic] (should be Dolichopus acuticornis); pseudoscorpion as Microcreagris strandi (Ellingsen, 1901). The attachment to the fly may have occurred in the net, as the specimen was presumably collected using a hand net.
14 Pseudoscorpion as Parachernes (Argentochernes) nigrimanus Beier, 1948; junior homonym, replaced by Parachernes (P.) beieri.
15 Host as fly; pseudoscorpion as Chelifer latreillii Leach, 1817. In Beier (1948), the host is specified as Fannia canalicularis [sic] (should be Fannia canicularis), and the pseudoscorpion is redetermined as Lamprochernes nodosus.
16 Host as Homalomyia canicularis (Linnaeus, 1761). Pseudoscorpion originally as C. cancroides; in Beier (1948) as L. nodosus.
17 Host as Homalomyia canicularis; pseudoscorpion as Chelifer (Chernes) nodosus Schrank, 1803.
18 Pseudoscorpion as Pseudozaona Beier, 1932; junior synonym of Chelodamus Chamberlin, 1925, synonymised by Muchmore (1984).
19 Host as Amoebalaria [sic] (should be Amoebaleria).
20 Host as Heteromyza flavipes Zetterstedt, 1838; in Beier (1948) as Tephrochlamis [sic] canescens (should be Tephrochlamys canescens (Meigen, 1830)). Pseudoscorpion as Chernes Menge, 1855; in Beier (1948) as Lamprochernes nodosus (Schrank, 1803).
21 Host as Dicranomyia decemmaculata (Loew, 1873).
22 Host as subgenus Achyrolimonia within the genus Limonia.
23 Pseudoscorpion as Dactylochelifer latreillii latreillii (Leach, 1817).
24 Host as Lonchaea vaginalis Fallén, 1820. Pseudoscorpion as C. cancroides; in Beier (1948) and WPC (2026) as L. nodosus.
25 Host as L. vaginalis. Pseudoscorpion as Chernes nodosus.
26 Pseudoscorpion as Lamprochernes godfreyi (Kew, 1911). Chelifer (Chernes) godfreyi Kew, 1911 is a junior synonym of Lamprochernes savignyi Simon, 1881, synonymised by Muchmore (1975).
27 Host changed from Ophyra leucostoma (Wiedemann, 1817) (Anthomyiidae) to Hydrotaea ignava (Harris, 1780) (Muscidae). Pseudoscorpion as Chernes nodosus.
28 Host as Musca meteorica Linnaeus, 1758. Pseudoscorpion as C. cancroides in the original paper; treated as a misidentification in WPC (2026).
29 Host as Musca corvina Fabricius, 1781; pseudoscorpion as Chernes nodosus.
30 Host only as a fly; specified as M. domestica in Beier (1948).
31 Pseudoscorpion as Chelifer panzeri Koch, 1836.
32 Pseudoscorpion as Toxochernes panzeri (C.L. Koch, 1836).
33 Host as a fly; pseudoscorpion as Acarus cancroides Linnaeus, 1758. This record is treated as a misidentification and referred to L. nodosus (Beier 1948; WPC 2026); the host is specified as M. domestica in Beier (1948).
34 Host as a fly; pseudoscorpion without precise identification. In Beier (1948), treated as L. nodosus on M. domestica.
35 Host as fly; in Beier (1948) treated as M. domestica. Pseudoscorpion as Chelifer parasita Hermann, 1804; a junior synonym of L. nodosus, synonymised by Simon (1879).
36 Host as flies; pseudoscorpion not mentioned in the original paper. In Beier (1948), treated as L. nodosus on M. domestica.
37 Host as Musca carnaria Linnaeus, 1758 (Anonymous 1831); pseudoscorpion not specified. Corrected to M. domestica and C. cancroides (as Chélifer cancröìdes) (Anonymous 1832). In Beier (1948), the phoront is treated as L. nodosus.
38 Pseudoscorpion as C. cancroides in the original paper; treated as a misidentification in Beier (1948) and WPC (2026).
39 Pseudoscorpion as Chelifers in the original paper; in Beier (1948) as L. nodosus.
40 Pseudoscorpion as Chelifers in the original paper. This record is treated as a misidentification and referred to L. nodosus (Beier 1948; WPC 2026).
41 Pseudoscorpion as C. cancroides; in Beier (1948), as L. nodosus.
42 Pseudoscorpion as Chelifer reussii C.L. Koch, 1843; a junior synonym of L. nodosus, synonymised by Simon (1879).
43 Host as a fly; pseudoscorpion as C. cancroides. In Beier (1948), referred to L. nodosus on M. domestica; the pseudoscorpion identification is confirmed in WPC (2026).
44 In the original paper, the host is mentioned only as flies; in Beier (1948), it is specified as M. domestica. Pseudoscorpion as Obisium muscorum Leach, 1817, treated as a misidentification and referred to L. nodosus (Beier 1948; WPC 2026).
45 Host as flies; pseudoscorpion as Chelifer nodosus Schrank, 1803. In Beier (1948), the host is specified as M. domestica.
46 Host as flies. Pseudoscorpion not specified in the original paper (Courtois de Langlade 1883); later cited, with the phoront given as C. cancroides in Mégnin (1886). This record is treated as a misidentification and referred to L. nodosus (Beier 1948; WPC 2026). Host as M. domestica in Beier (1948).
47 Pseudoscorpion as Chernes nodosus Schrank, 1803.
48 Host originally reported as flies; pseudoscorpion as Chernes nodosus. In Beier (1948), several fly records were specified as M. domestica (see Note 6).
49 Pseudoscorpion as Chelifer nodosus. The specimens cited in this work belong only partly to L. nodosus (three specimens), whereas one specimen belongs to L. chyzeri; for details, see Lessert (1911).
50 Pseudoscorpion in the original paper as Pseudoscorpiones (Chelifer?), as L. nodosus in Beier (1948).
51 Pseudoscorpion originally as Chelifer cancroides (Linnaeus, 1758), but Lessert (1911) suggested that it should be referred to Chelifer (Lamprochernes) nodosus Schrank, 1803.
52 Pseudoscorpion as Chelifer (Lamprochernes) nodosus Schrank, 1803.
53 The pseudoscorpion is given as Chernes nodosus on flies. Beier (1948) redetermined the host as M. domestica but incorrectly cited Kew (1916). The record in Kew (1916) is itself cited from Carpenter (1912) (citation no. 24 on p. 78), to whom the original data should be attributed. Beier (1948) also incorrectly stated England as the country of origin; the correct locality is Ireland.
54 Pseudoscorpion as Chelifer nodosus Schrank, 1803.
55 Pseudoscorpion as Chernes reussii (C.L. Koch, 1843), a junior synonym of L. nodosus, synonymised by Simon (1879).
56 Pseudoscorpion as Chelifer (Chernes) nodosus.
57 Pseudoscorpion as C. cancroides in Stephens (1910), but in Kew (1916) redefined as Chelifer (C.) godfreyi, later synonymised with L. savignyi by Muchmore (1975).
58 Pseudoscorpion as Chelifer (C.) godfreyi, later synonymised with L. savignyi by Muchmore (1975). Beier (1948) incorrectly stated England as the country of origin, whereas the correct locality is Ireland.
59 Host as Musca vicina Macquart, 1851. Pseudoscorpion as Muscichernes katoi Morikawa, 1960, a junior synonym of L. savignyi, synonymised by Harvey (1987).
60 Pseudoscorpion as Pycnochernes Beier, 1932, a junior synonym of Lamprochernes, synonymised by Muchmore (1975).
61 Pseudoscorpion as Obisium muscorum.
62 Pseudoscorpion as C. cancroides; in Beier (1948) as Neochernes sanborni (Hagen, 1869). Muchmore (1971) stated that this record cannot be identified as Chernes sanborni Hagen, 1868, and is therefore treated here as Not specified.
63 In the original paper, the phoront is mentioned only as a pseudoscorpion; in Beier (1948), it is given as N. sanborni. Muchmore (1971) stated that this record cannot be identified as Chernes sanborni Hagen, 1868, and is therefore treated here as Not specified.
64 Pseudoscorpion as Dinocheirus sicarius Chamberlin, 1952, a junior synonym of Dinocheirus serratus (Moles, 1914), synonymised by Muchmore (1997).
65Beier (1948) mentioned the host only as Diptera, whereas in the original data, it is specified as a muscid (Ellingsen 1913). Pseudoscorpion as Chelifer nodosus in the original paper from Gold Coast, West Africa (present-day Ghana).
66 Pseudoscorpion as Chernes nodosus. Data on phoresy based on Plate XXX.
67 Pseudoscorpion as Chelifer (C.) godfreyi, later synonymised with L. savignyi by Muchmore (1975).
68 Pseudoscorpion as Chernes scorpioides (Hermann, 1804).
69 The paper also includes data from experiments; however, only field data were considered here.
70 Pseudoscorpion Semeiochernes armiger Balzan, 1892 – the author and year should be given in parentheses.
71 Pseudoscorpion as Semeiochernes militaris Beier, 1932; in Aguiar and Bührnheim (1998b: 458), a host is added and the species is given as S. armiger.
72 Data mentioned in the Addendum.
73 Host as Omphrale fenestralis (Linnaeus, 1758).
74 Host as Leptocera sylvatica [sic] (Meigen, 1830) (should be Leptocera sylvatica).
75 Host as Hoplites [sic] bispinosus Macq. (should be Hoplistes bispinosus (Wiedemann, 1830)). Pseudoscorpion not identified in the original paper; treated as L. nodosus in Beier (1948).
76 The host family as Stratiomydae [sic] (should be Stratiomyidae).
77 Pseudoscorpion as Chelifer (Chernes) nodosus in Berland (1932).
78 Host given as “Serphid fly”, a misspelling referring to a syrphid fly (hoverfly).
79 Host as Musca larvarum Linnaeus, 1758 (Muscidae), now Exorista larvarum (Tachinidae). Pseudoscorpion as C. cancroides in the original paper; treated as a misidentification in WPC (2026).
80 Host as “Dexidae” (and “Dexiidae”) refers to the tachinid subfamily Dexiinae (Tachinidae), not to Dixidae; these spellings represent historical variants of Dexiinae rather than a misidentification. Pseudoscorpion as Chelanops pallipes Banks, 1893.
81 Pseudoscorpion given as C. cimicoides = C. hahnii in Wagner (1892); in Berland (1932) as Chelifer (Chernes) nodosus. In WPC (2026), this record is treated as a misidentification and referred to L. nodosus.
82 Pseudoscorpion as Chelifer geoffroyi Leach, 1817, a junior synonym of Chernes cimicoides, synonymised by Simon (1879). In WPC (2026), this record is treated as a misidentification and referred to L. nodosus.
83 Pseudoscorpion as genus Obisium Illiger, 1798; as Syarinus obscurus (Banks, 1893) in Muchmore (1971).
84 Pseudoscorpion as Dactylochelifer latreillii (Leach, 1817).
85 Host as Chrysomyza demandata (Fabricius, 1798).
86 Host as Ulidia demandata (Fabricius, 1798). Pseudoscorpion as Chelifer corallinus Loew, 1845, later synonymised with L. nodosus by Tömösváry (1883).
87 Host as Chloria demandata (Fabricius, 1798). Pseudoscorpion as Chelifer hahnii C.L. Koch, 1839, treated as a misidentification in WPC (2026) and referred to L. nodosus.
88 Host as Physiphora demandata (Fabricius, 1798).
89 Pseudoscorpion as “probably Chernetidae”, treated here as Not specified.
90 Host as Chrysomyza aenea (Fabricius, 1794).
91 Pseudoscorpion as Chelifer wideri C.L. Koch, 1843, treated as a misidentification in WPC (2026) and referred to as L. nodosus.
92 Pseudoscorpion as Chelifer loewi – nomen nudum.
93 Pseudoscorpion as chelifers in Hagen (1867, 1868); treated as Neochernes sanborni by Muchmore (1971), currently as Chernes sanborni.
94 Pseudoscorpion as Chernes santorni [sic] (should be Chernes sanborni).
95 Pseudoscorpion females found at the bottom of the Malaise trap were not considered phoretic.
96 Host as a fly; pseudoscorpion as Chernes nodosus. In Beier (1948: 442), another record of L. nodosus on Calliphora is mentioned, but this could not be found in the original paper.
97 Several fly species are listed under Diptera.
98 Pseudoscorpion as Chelifer nodosus. In Beier (1948) and WPC (2026), the record referred to Lamprochernes godfreyi. Chelifer (Chernes) godfreyi is a junior synonym of Lamprochernes savignyi, synonymised by Muchmore (1975).
99 Pseudoscorpion as Chelifer (Chernes) godfreyi Kew, 1911 on flies. Chelifer (Chernes) godfreyi is a junior synonym of Lamprochernes savignyi, synonymised by Muchmore (1975).
Data about Chernes nodosus from Kew (1911) are just citations of other papers. Beier (1948) treats these under L. nodosus, but these records are based solely on previously published sources and do not constitute new data.
100 The records from Beier (1946, 1948) for Sudan were merged into a single record (one female from a fly trap, February).
101 Pseudoscorpion as Chelifer scorpioides Hermann, 1804.
Fossil records
The fossil data were compared with Poinar et al. (1998) and supplemented with three studies published subsequently. Table 2 includes, where provided in the original publications, information on host and phoront taxa, amber type, and reference. Missing data from published records are marked as “Not specified” in Table 2. Notes on discrepancies and differences among records are presented in Table 2. Altogether, five pseudoscorpion records were documented, of which three were identified to species or genus level. These phoronts were attached to the legs of five dipteran taxa representing three families, with some hosts remaining unidentified; only two dipteran species could be identified to species level (Table 2). Regarding amber provenance, most records originate from Baltic amber (Table 2), indicating that phoretic associations between pseudoscorpions and their dipteran hosts were already established by at least the Paleogene.
Table 2.
Fossil records of phoresy between Pseudoscorpiones and Diptera.
| Host taxa | Phoront taxa | Amber type | References | |
|---|---|---|---|---|
| Limoniidae | ||||
| Trentepohlia immemorata Podenas & Poinar, 2001 | Cheliferidae | Hysterochelifer manpauch Córdova-Tabares, Riquelme & Villegas-Guzmán, 2024 | Mexican amber | Córdova-Tabares et al. (2024) |
| Not specified1 | Not specified | Not specified | Baltic amber | Schlee and Glöckner (1978) |
| Tipulidae | ||||
| Not specified | Garypinidae | Garypinus electri Beier, 1937 | Baltic amber | Judson (2004) |
| Xylophagidae 2 | ||||
| Chrysothemis speciosa Loew, 1850 | Not specified | Not specified | Baltic amber | Ross (1997) |
| Not specified3 | Cheliferidae | Electrochelifer Beier, 1937 | Bitterfeld amber | Ahrens et al. (2019) |
Notes: 1 In Poinar et al. (1998) as Tipulidae and Chernetidae; 2 Host family originally as Rachiceridae; 3 Host only as probably Nematocera.
Conclusion
In summary, following a revision of published data and the addition of new European records, pseudoscorpions were recorded phoretically on dipteran hosts belonging to 74 species (59 genera) from 30 families, with the majority of records associated with Musca domestica (Muscidae). Although 47 records are identified only at the level of Diptera, pseudoscorpions participating in phoretic associations with Diptera represent 39 species (21 genera) from seven families, most of which belong to the genus Lamprochernes. However, these data are likely biased, as earlier studies predominantly reported L. nodosus attached to the house fly, a pattern further reinforced by the extensive redeterminations of specimens by Beier (1948). This taxonomic uncertainty complicates the interpretation of historical data and underscores the need for careful reassessment of species identities in phoretic records. Geographically, the records originate from 44 countries, although some data were reported only at the continental level. Fossil evidence from amber further indicates that pseudoscorpion–Diptera phoretic associations represent a long-standing ecological interaction, already established by at least the Paleogene and persisting into the present day. Compared with Poinar et al. (1998), the number of dipteran hosts increased by 37 species (27 genera) and 13 families, while the number of pseudoscorpion phoronts increased by 16 species (five genera) and three families. This synthesis provides the most comprehensive overview to date of pseudoscorpion–Diptera phoretic associations and establishes a robust framework for future taxonomic, ecological, and evolutionary studies.
Acknowledgements
We are very grateful to all collectors of pseudoscorpions and to the dipteran specialists listed in this article who assisted with the identification of host species. We especially thank our colleagues Jiří Roháček and Samuel Krčmárik for valuable discussions on Diptera. We are grateful to Alica Christophoryová for her technical assistance with the figures. We thank our students Ema Hetešová and Simona Chamulová for their assistance with selected data. The authors received research support from the Scientific Grant Agency of the Ministry of Education of the Slovak Republic, Grant No. VEGA 1/0817/25.
Funding Statement
Scientific Grant Agency of the Ministry of Education of the Slovak Republic, Grant No. VEGA 1/0817/25
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Use of AI
No use of AI was reported.
Funding
This work was supported by the Scientific Grant Agency of the Ministry of Education of the Slovak Republic, Grant No. VEGA 1/0817/25.
Author contributions
Conceptualization: LV, JC. Data curation: LV, CH, VM, JC. Formal analysis: JC. Methodology: LV, JC, VM. Resources: LV, JC, CH, VM. Visualization: JC. Writing - original draft: JC. Writing - review and editing: VM, LV, CH.
Author ORCIDs
Jana Christophoryová https://orcid.org/0000-0002-3746-1367
Vincent Mathy https://orcid.org/0000-0002-8448-3112
Christoph Hörweg https://orcid.org/0000-0002-8208-0758
Lucia Vičanová https://orcid.org/0009-0005-9558-8437
Data availability
All of the data that support the findings of this study are available in the main text.
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Data Availability Statement
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