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Research Article
Mites of the genus Bryobia (Acari, Tetranychidae): taxonomic notes on some species and a diagnostic key to the world species
expand article infoJawwad Hassan Mirza‡, Nasreldeen Ahmed Elgoni‡, Muhammad Kamran‡, Fahad Jaber Alatawi‡
‡ King Saud University, Riyadh, Saudi Arabia

Corresponding author: Fahad Jaber Alatawi ( falatawi@ksu.edu.sa )

Academic editor: Fernando Jacinavicius
© 2025 Jawwad Hassan Mirza, Nasreldeen Ahmed Elgoni, Muhammad Kamran, Fahad Jaber Alatawi.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Mirza JH, Elgoni NA, Kamran M, Alatawi FJ (2025) Mites of the genus Bryobia (Acari, Tetranychidae): taxonomic notes on some species and a diagnostic key to the world species. ZooKeys 1239: 51-73. https://doi.org/10.3897/zookeys.1239.149111
ZooBank: urn:lsid:zoobank.org:pub:4B36CD7E-C1C3-4391-AD46-43A8769AB8E8
Open Access

Abstract

The present study aimed to develop taxonomic keys to the world species of the genus Bryobia, categorized into three subgenera: Allobia Livschits & Mitrofanov, Bryobia s. str. Koch, and Lyobia Livschits & Mitrofanov. Published descriptions, redescriptions, and illustrations of a total of 149 world species were thoroughly analyzed. The taxonomic notes on the status of the species in the species groups of each subgenus are discussed in detail. The variability of morphological characters found among different populations of a species is discussed. As a result, 116 species of the genus Bryobia were classified in three diagnostic keys, with 22, 43, and 51 species assigned to the three subgenera Allobia, Bryobia, and Lyobia, respectively. The population of B. neoribis Tuttle & Baker from Utah, USA, should be re-identified through type examination due to differences from the original description of the species. Additionally, taxonomic notes are provided on the status of the remaining 33 species, and arguments are provided on suggested synonyms among them.

Key words:

Allobia, Lyobia, neoribis, praetiosa, species groups, subgenera

Introduction

The genus Bryobia Koch, 1836, is the largest in the subfamily Bryobiinae (Pritchard and Baker 1955) and comprises 149 described species reported globally (Migeon and Dorkeld 2025). These mites are phytophagous and include some of the most notorious pests (Jeppson et al. 1975). The clover mite, B. praetiosa Koch, 1836, is a famous member of the genus, infesting different economic fruit, grain, and ornamental crops, and is distributed worldwide (Jeppson et al. 1975).

Historically, Bryobia species were once divided into seven species groups based on the presence of a row of stout setae on leg femur I (Eyndhoven 1956). Livschits and Mitrofanov (1971) introduced a comprehensive analysis of the genus, provided new species synonymies, and, based on the combination of eight morphological characters, proposed five subgenera in the genus, while the sixth subgenus was added by Mitrofanov (1973). Recently, Mirza et al. (2024) comprehensively re-evaluated those morphological characters for generic differentiation and proposed three subgenera in the genus Bryobia: Bryobia Koch s. str., Allobia Livschits & Mitrofanov, and Lyobia Livschits & Mitrofanov. These subgenera were diagnosed based on the presence or absence of duplex setae (tactile seta with a sensory solenidion) on leg tarsi III and IV (Fig. 1a, b). Additionally, the species of each subgenus were categorized into three species groups based on the position of the inner sacral setae f1 (Mirza et al. 2024). A total of eight species described by Meyer (1974, 1987), which possess pad-like true claws on leg I, were also discussed over the contradiction with the diagnosis of the tribe Bryobiini (Mirza et al. 2024).

Figure 1. 

Duplex setae on leg tarsi III and IV a duplex setae present on both leg tarsi III-IV in Bryobia (Bryobia) praetiosa Koch, 1836 (redrawn from Livschits and Mitrofanov 1971) b duplex setae absent on leg tarsus III in Bryobia (Lyobia) rubrioculus (Scheuten, 1857) (redrawn from Vacante 1983).

There are various morphological characters that have been misinterpreted (e.g., the position of inner and outer sacrals), while others have been mistakenly considered as differences to distinguish species, rather than as intraspecific variations (i.e., body length, length of propodosomal lobes, number of setae on leg segments, length of leg segments). This raised the issue of species complexes, and the perfect example is the praetiosa species complex (Pritchard and Baker 1955). Different regional keys have been published over time from around the world, including those from Africa, Asia, the USSR, the USA, and Europe (Meyer 1974, 1987, 1992; Eyndhoven and Vacante 1985; Livschits and Mitrofanov 1971; Ehara 1999; Auger et al. 2015; Çobanoğlu et al. 2021; Stathakis et al. 2022). However, in the absence of a world key to Bryobia species, it would be difficult to grasp the true species identity. This study, based entirely on published literature, aims to distinguish true morphological differences from intraspecific variations to validate species statuses, develop taxonomic keys for the world’s Bryobia species within the three subgenera proposed by Mirza et al. (2024), and provide taxonomic notes on the status of certain species.

Materials and methods

The published morpho-taxonomic literature of 149 world species of the genus Bryobia was collected using the websites of different research journals and spider mite web databases (Migeon and Dorkeld 2006–2025). All the published literature related to the taxonomy and systematics of the Bryobia species were equally considered. The classification proposed by Mirza et al. (2024) is followed for the subgenera and species groups. The species descriptions, redescriptions, illustrations, taxonomic revisions, and regionally prepared identification keys were critically investigated to develop three dichotomous keys for the three subgenera of the genus Bryobia to identify the species.

Results and discussion

Family Tetranychidae Donnadieu

Subfamily Bryobiinae Berlese

Bryobiini Reck

Diagnosis.

True claws uncinate and empodium pad-like.

Bryobia Koch, 1836

Type species.

Bryobia praetiosa Koch, 1836: 8.

Diagnosis

(based on females). As defined by Arabuli et al. (2019) and Mirza et al. (2024).

There are four species, B. apsheronica Khalilova, 1953, B. desertorum Hassan, Afifi & Nawar, 1986, B. ribis Thomas, 1896, and B. weyerensis Packard, 1889, not included in any subgenus or species group due to inadequate and insufficient literature, as also reported by Mirza et al. (2024). In the very brief descriptive statements of B. weyerensis, the original author provided the two completely different generic names to which this species may belong, “Bryiobia ? (or Penthaleus)” (Packard 1889). The former three species require re-description based on type examination to be added to the respective subgenus and species group.

Allobia Livschits & Mitrofanov, 1971

Type species.

Bryobia pritchardi Rimando, 1962: 9.

Diagnosis

(based on females). As defined by Mirza et al. (2024).

Key to the 22 species of the subgenus Allobia

Species groups definition is based on Mirza et al. 2024.

1 Dorsocentral setae f1 present centrally, aligned with other dorsocentral setae, the distance f1-f1 is always shorter than f2-f2 (Fig. 2a) abbatielloi species group 3
– Dorsocentral setae f1 present laterally or sub laterally 2
2 Dorsocentral setae f1 present laterally along the margin and the distance f1-f1 is always greater than f2-f2 (Fig. 2b) pritchardi species group 4
– Dorsocentral setae f1 present sub laterally, neither aligned with other dorsocentral setae nor present marginally, and the distance f1-f1 could be shorter or longer than f2-f2 (Fig. 2c) deserticola species group 21
3 Propodosoma with distinct, 4 well-developed lobes (Fig. 3a) B. (A.) querci Hatzinikolis & Panou, 1997
– Propodosomal lobes absent (Fig. 3b) B. (A.) abbatielloi (Smiley & Baker, 1995)
4 Empodium I with 1 pair of tenent hairs (Fig. 4a) 5
– Empodium I with > 1 pairs of tenent hairs (Fig. 4b) 16
5 Genu I with ≤ 6setae 6
– Genu I with 7 or 8 setae 9
6 Genu I with 6 setae; femur I with 9 setae B. (A.) beaufortensis Meyer, 1992
– Genu I with 4 or 5 setae 7
7 Propodosomal lobes well developed; peritremes ending in an enlarge anastomose (Fig. 5a) B. (A.) marcandrei Hatzinikolis & Panou, 1996
– Propodosomal lobes weakly developed 8
8 Peritremes ending in simple bulb (Fig. 5b); genu I with 5 setae B. (A.) ylikiensis (Hatzinikolis & Emmanouel, 1993)
– Peritremes ending in an ovate anastomosis; genu I with 4 setae B. (A.) giannitsensis Hatzinikolis & Panou, 1996
9 Median propodosomal lobes well developed; femur III with 3 setae B. (A.) relhaniae Meyer, 1992
– Median propodosomal lobes weakly developed or fused into a single lobe (Fig. 3c, d) 10
10 Femur II with ≥ 11 setae 11
– Femur II with 8–10 setae 13
11 Propodosoma with 8 setae 12
– Propodosoma with 7 setae; peritremes anastomosed B. (A.) aegyptiacus (Zaher, Gomaa & El-Enany, 1982)
12 Peritremes terminate in simple bulb B. (A.) nigromontana Meyer, 1992
– Peritremes terminate in a chamber consisting of a few lobes; stylophore with deep depression B. (A.) geyeri Meyer, 1974
13 Femur II with 9 setae B. (A.) caricae Hatzinikolis & Emmanouel, 1991
– Femur II with 8 setae 14
14 Tibia III with 7 setae B. (A.) macedonica Hatzinikolis & Panou, 1996
– Tibia III with 9 setae 15
15 Tarsus III without solenidion, peritremes elongate anastomose B. (A.) pritchardi Rimando, 1962
– Tarsus III with a solenidion, peritremes simple B. (A.) meyerae Zaher, Gomaa & El-Enany, 1982
16 Propodosoma with incomplete reticulation medially; peritremes end in simple bulb B. (A.) angolensis Meyer, 1987
– Propodosoma without reticulation 17
17 Peritremes end in small anastomosis 18
– Peritremes end in simple bulb 20
18 Median propodosomal lobes fused into a single lobe 19
– Median propodosomal lobes well incised and developed; palp tarsus with 7 setae B. (A.) imbricata Meyer, 1974
19 Palp tarsus with 6 setae B. (A.) monechmae Meyer, 1974
– Palp tarsus with 7 setae B. (A.) tuberosa Meyer, 1974
20 Femur I with 7 setae; stylophore deeply incised B. (A.) coatesi Meyer, 1974
– Femur I with > 7 setae; stylophore rounded B. (A.) incana Meyer, 1992
21 Empodium I with a pair of tenant hairs; peritremes ending in a small anastomosis B. (A.) deserticola Meyer, 1989
– Empodium I with 2 pairs of tenant hairs; peritremes ending in a simple bulb B. (A.) birivularis Meyer, 1989

Notes on the species of the subgenus Allobia

The subgenus Allobia includes 28 species (Mirza et al. 2024) although only 22 valid species are included in the key above. Among the remaining six species, five species described by Meyer (1974, 1987) have pad-like true claws on leg I. Mirza et al. (2024) provided a detailed discussion on how this character state contradicts the diagnosis of the Bryobiini tribe. In the present study, the sixth species B. (A.) orycustodia Meyer (in Meyer & Ueckermann, 1989) from the species group pritchardi is also considered among those five species of Meyer in which leg I true claws are also pad-like. These six species were not added to the diagnostic key for the time being as this requires an update of the diagnoses of all tribes of subfamily Bryobiinae based on the shape of leg I true claws.

Figure 2. 

Position of setae f1 and relative distances of f1–f1 vs f2–f2 (shown by dashed line) a setae f1 present centrally in Bryobia (Allobia) abbatielloi (Smiley & Baker, 1995) (redrawn from Smiley and Baker 1995) b setae f1 present laterally in Bryobia (Allobia) pritchardi Rimando, 1962 (redrawn from Rimando 1962) c setae f1 present sub laterally in Bryobia (Bryobia) artemisiae Bagdasarian, 1951 (redrawn from Livschits and Mitrofanov 1971).

Figure 3. 

Development of propodosomal lobes a well developed in Bryobia (Bryobia) praetiosa Koch, 1836 (redrawn from Livschits and Mitrofanov 1971) b absent in Bryobia (Allobia) abbatielloi (Smiley & Baker, 1995) (redrawn from Smiley and Baker 1995) c three lobes with median lobes fused in Bryobia (Bryobia) bakeri (Zaher, Gomaa & El-Enany, 1982) (redrawn from Zaher et al. 1982) d three lobes in which median lobe is weakly developed in Bryobia (Allobia) geyeri Meyer, 1974 (redrawn from Meyer 1974).

Figure 4. 

Number of tenent hairs on leg I empodium a one pair of tenent hairs in Bryobia (Bryobia) strunkovae Mitrofanov, 1968 (redrawn from Mitrofanov 1968) b more than one pair of tenent hairs in Bryobia (Bryobia) borealis Oudemans, 1930 (redrawn from Mathys 1962).

Figure 5. 

Shape of peritremes a enlarged, anastomose in Bryobia (Allobia) marcandrei Hatzinikolis & Panou, 1996 (redrawn from Hatzinikolis and Panou 1996) b simple bulb in Bryobia (Allobia) birivularis Meyer, 1989 (redrawn from Meyer and Ueckermann 1989).

Species group abbatielloi

There are only two species existing in the species group abbatielloi, B. (A.) abbatielloi (Smiley & Baker, 1995) and B. (A.) querci Hatzinikolis & Panou, 1997 (Mirza et al. 2024). The species B. (A.) querci was distinguished by the presence of f2 setae in line with other dorsocentral setae c1, d1, and e1 (Hatzinikolis and Panou 1997). This position of seta f2 is incorrectly described in this species, based on the nomenclature of Lindquist (1985). Hence, the seta f2 (outer sacral) described by Hatzinikolis and Panou (1997) is actually seta f1 (inner sacral) and vice versa.

Bryobia s. str. Koch, 1836

Type species.

Bryobia praetiosa Koch, 1836: 8.

Diagnosis

(based on females). As defined by Mirza et al. (2024).

Key to the 43 species of the subgenus Bryobia

Species groups definition is based on Mirza et al. 2024

1 Fourth pair of dorsocentral setae f1 present centrally, aligned with another 3 pairs of dorsocentral setae neoephedrae species group B. (B.) neoephedrae (Gutierrez & Bolland, 1998)
– Fourth pair of dorsocentral setae f1 present sublaterally where the distance f1-f1 is shorter than f2-f2 osterloffi species group 2
– Fourth pair of dorsocentral setae f1 present laterally, along the margin and the distance f1-f1 always greater than f2-f2 praetiosa species group 8
2 Femur IV with ≥ 5 setae 4
– Femur IV with ≤ 5 setae 3
3 Genu II with 5 setae B. (B.) artemisiae Bagdasarian, 1951
– Genu II with 3 setae B. (B.) serifiotica Hatzinikolis, Papadoulis & Kapaxidi, 2007
4 Femur IV with 7 setae B. (B.) abyssiniae Fashing & Ueckermann, 2016
– Femur IV with 5 setae 5
5 Femur III with 4 or 5 setae 6
– Femur III with 6 or 7 setae 7
6 Genu II with 8 setae B. (B.) petrilunara Meyer, 1987
– Genu II with 5 or 6 setae B. (B.) burkei Meyer, 1987
7 Genu IV with 6 setae B. (B.) osterloffi Reck, 1947
– Genu IV with 4 or 5 setae B. (B.) variabilis Manson, 1967
8 Propodosoma with 7 setae B. (B.) bakeri (Zaher, Gomaa & El-Enany, 1982)
– Propodosoma with 8 setae 9
9 Femur IV with 2 or 3 setae 10
– Femur IV with > 3 setae 13
10 Genu I with 8 setae B. (B.) meteoritica Hatzinikolis & Panou, 1996
– Genu I with 4 setae 11
11 Femur II with 6 setae B. (B.) reckiana Mitrofanov & Strunkova, 1968
– Femur II with 5 setae 12
12 Genu III with 3 setae B. (B.) montana Mitrofanov, 1973
B. (B.) nitrariae He & Tan, 1993
– Genu III with 2 setae B. (B.) tadjikistanica Livschits & Mitrofanov, 1968
13 Femur IV with 6 setae 14
– Femur IV with < 6 setae 17
14 Genu II with 6 setae 16
– Genu II with 4 or 5 setae 15
15 Femur I with 18 setae; genu I with 6 setae B. (B.) xiningensis Ma & Yuan, 1981
– Femur I with ≥ 20 setae; genu I with > 6 setae B. (B.) vasiljevi Reck, 1953
16 Dorsal integument densely granulates without striae B. (B.) agioriticus Hatzinikolis & Emmanouel, 1996
– Propodosoma with irregular discontinuous fine striae, hysterosoma mostly transverse with irregular fine striae medially B. (B.) alberensis Auger & Migeon, 2023 (in Auger et al. 2023)
17 Genu II with 7 or 8 setae 38
– Genu II with < 7 setae 18
18 Femur I with ≥ 14 setae 21
– Femur I with ≤ 13 setae 19
19 Empodium I with a pair of tenant hairs 20
– Empodium I with > 1 pair of tenant hairs; dorsocentral setae c1 and d1 crossing basis of next setae B. (B.) hengduanensis Wang & Cui, 1991
20 Tibiae I and II with 11 or 12 and 9 setae, respectively B. (B.) strunkovae Mitrofanov, 1968
– Tibiae I and II with 16 and 8 setae, respectively B. (B.) ziziphorae Strunkova & Mitrofanov, 1983
21 Tibia I with ≥ 21 setae B. (B.) macrotibialis Mathys, 1962
– Tibia I with ≤ 20 setae 22
22 Dorsal setae c2 and c3 are in the same horizontal line 27
– Dorsal setae c2 and c3 distinctly not in the same horizontal line 23
23 Tarsi III and IV each with 13 setae B. (B.) gigas Auger, Arabuli & Migeon, 2014
– Tarsi III and IV each with > 13 setae 24
24 Genua III and IV with 3 and 4 setae, respectively B. (B.) qilianensis Ma & Yuan, 1981
– Genua III and IV each with 6 setae 25
25 Femora III and IV each with 4 or 5 setae 26
– Femora III and IV with 7 and 5 setae, respectively B. (B.) latisetae Wang, 1985
26 Femora III and IV each with 4 setae B. (B.) exserta Wang, 1985
– Femora III and IV each with 5 setae B. (B.) graminum (Schrank, 1781)
B. (B.) monticola Wang, 1985
27 Genua I and II with 4 and 3 setae, respectively 28
– Genu I with 7 or 8 setae, genu II with 5 or 6 setae 29
28 Stylophore anteriorly rounded, true claws of leg II-IV with 2 rows of tenent hairs ……………… B. (B.) magallanica Gonzalez, 1977
– Stylophore anteriorly slightly notched, true claws of leg II-IV with 4–8 tenent hairs B. (B.) glacialis Berlese, 1913
29 Tibia I with 12–16 setae 32
– Tibia I with 17–20 setae 30
30 Tarsus I with 20 setae 31
– Tarsi I and II with 31 and 19 setae respectively B. (B.) qinghaiensis Ma & Yuan, 1981
31 Femur I with 23 setae; tarsus II with 15 setae B. (B.) cyclamenae Hatzinikolis & Panou, 1996
– Femur I with 19 setae; tarsus II with 18 setae B. (B.) platani Hatzinikolis & Panou, 1997
32 Empodium I with 2 rows of tenant hairs B. (B.) borealis Oudemans, 1930
– Empodium I with a pair of tenant hairs 33
33 Dorsal body setae palmate (Fig. 6a); femur IV with 4 setae; tarsus IV with 14 setae B. (B.) fuegina Gonzalez, 1977
– Dorsal body setae not as above 34
34 Propodosoma without lateral projection; tarsal claws II-IV each with > 1 pair of tenant hairs 35
– Propodosoma with lateral projection 36
35 Tarsus I with a pair of tenent hairs B. (B.) cagani Çobanoğlu, Ueckermann & Cilbircioğlu, 2021
– Tarsus I with > 1 pair of tenent hairs B. (B.) urticae Sayed, 1946
36 Stylophore rounded B. (B.) praetiosa Koch, 1836
B. (B.) kissophila Eyndhoven, 1955
– Stylophore notched 37
37 Tibia I with 14–16 setae; femur II with ≥ 10 setae B. (B.) watersi Manson, 1967
– Tibia I with 13 setae; femur II with 8 setae B. (B.) attica Hatzinikolis & Emmanouel, 1990
38 Genu II with 7 setae B. (B.) emmanoueli Hatzinikolis & Panou, 1996
– Genu II with 8 setae B. (B.) nikitensis Livschits & Mitrofanov, 1969

Notes on the species of the subgenus Bryobia

The subgenus Bryobia includes 53 species (Mirza et al. 2024), although only 43 species are included in the key above. Among the remaining ten, two species belong to the species group praetiosa, B. geigeriae Meyer, 1974, and B. karooensis Meyer, 1974, which are excluded from the key due to ambiguity in the leg I true claw morphology as debated by Mirza et al. (2024). The two species B. (B.) calida Karg, 1985 and B. (B.) lagodechiana Reck, 1953 could not be assigned to any species group due to insufficient information available on the position of the inner sacral seta f1. The status of the remaining six species excluded from the above key is discussed below.

Figure 6. 

Shape of dorsal setae in adult female a palmate in Bryobia (Lyobia) alveolata Auger & Flechtmann, 2009 (redrawn from Auger et al. 2009) b fan-shaped in Bryobia (Lyobia) kakuliana Reck, 1956 (redrawn from Livschitz and Mitrofanov 1971) c spatulate in Bryobia (Allobia) birivularis Meyer, 1989 (redrawn from Meyer and Ueckermann 1989) d lanceolate in Bryobia (Lyobia) gushariensis Livschits & Mitrofanov, 1971 (redrawn from Livschits and Mitrofanov 1971) e setiform serrate in Bryobia (Lyobia) cinereae Auger & Migeon, 2014 (redrawn from Auger and Migeon 2014).

Species group praetiosa

The species B. (B.) montana Mitrofanov, 1973 was originally described from Tadjikistan on the host plant Astragalus sp., while the species B. (B.) nitrariae He & Tan, 1993 was reported from China on the host plant Nitraria sibirica. These two species are similar in all morphological characters, including leg chaetotaxy. The only difference is in the number of setae on tarsus I for both species, 20 vs 18, respectively. The descriptions of both species provided leg setal counts as the total number, including sensory and tactile setae. It is important to note that He and Tan (1993) differentiated B. (B.) nitrariae from B. (B.) tadjikistanica Livschits & Mitrofanov, 1968, which is also morphologically close to B. (B.) montana. The two species, B. (B.) tadjikistanica and B. (B.) montana, share the same type locality, Tadjikistan. There are also minor differences between B. (B.) nitrariae and B. (B.) tadjikistanica in the shape of their spermathecae and true claws. The two species B. (B.) montana and B. (B.) nitrariae key out near each other. Examining the type specimens would help to clarify their statuses.

The three species, B. (B.) graminum (Schrank, 1781), B. (B.) monticola Wang, 1985, and B. (B.) neopraetiosa Meyer, 1974 are also morphologically close. They have been reported from Germany (on Poaceae sp.), China (on Poaceae sp.), and South Africa (on multiple hosts), respectively. The leg chaetotaxy for B. (B.) neopraetiosa is neither described nor illustrated in detail (except for femur I, genua I and II, and tibia I), while that of B. (B.) graminum and B. (B.) monticola has few variations on leg tarsal segments. Based on the available descriptions, re-descriptions, and illustrations, it could be suggested that B. (B.) monticola and B. (B.) neopraetiosa should be synonymized with B. (B.) graminum. Similarly, the species B. (B.) exserta Wang, 1985 was reported from China on Artemisia sp. and was distinguished from B. (B.) praetiosa Koch, 1836 based on minor morphological variations, including body length, propodosomal lobe lengths, leg genu I segment comparative lengths. Bryobia (B.) exserta also morphologically resembles the three species discussed above. It is impossible to decide the synonymy of B. (B.) exserta, whether it should be synonymized with B. (B.) graminum or B. (B.) praetiosa. The species B. (B.) praetiosa is the type species of the genus Bryobia, while B. (B.) graminum, one of the oldest species described, was moved to the genus Bryobia by Oudemans (1929). Mitrofanov et al. (1987) synonymized B. (B.) praetiosa with B. (B.) graminum, but previously, Pritchard and Baker (1955) considered synonymizing B. (B.) praetiosa with B. (B.) graminum and suggested further detailed studies. However, these two species still remain valid (Migeon and Dorkeld 2025).

Two species, B. (B.) qinghaiensis Ma & Yuan, 1981 and B. (B.) yunnanensis Ma & Yuan, 1981, are described from China, from the Palearctic and Oriental regions, respectively. They are morphologically similar to each other, apart from some setal variations on leg tarsal and tibial segments, and were differentiated from B. (B.) praetiosa and B. (B.) qinghaiensis, respectively, based on a few minor differences. These species resemble B. (B.) praetiosa, the type of the genus. Note that the concept of apraetiosa species complex still exists, and there are a considerable numbers of populations described under the name of praetiosa, or otherwise, from different localities of the world. Each of those descriptions and illustrations provided various degrees of chaetotaxies and body measurements, which further complicate the true identification of B. (B.) praetiosa. Pritchard and Baker (1955) provided an excellent debate on the overall situation of the praetiosa complex. It appears that this complex and its synonyms will continue to grow.

The species B. (B.) batrae Baker & Tuttle, 1994 was described from the USA, occurring on the host plant Stellaria media. This species cannot be added to the key as it was very briefly described and illustrated. Baker and Tuttle (1994) also did not compare it with any related species. The species B. (B.) japonica Ehara & Yamada, 1968, also cannot be included as it was also very briefly described. The authors did compare it with B. (B.) sarothamni and B. (B.) tadjikistanica based on the absence of dorsal lobes. These two species belong to the subgenus Allobia (Bryobia) (Mirza et al. 2024).

Tuttle and Baker (1976) described B. (B.) neoribis with a duplex on both leg tarsi III and IV. However, their 1994 original description of the species on Acer negundo from Utah, USA stated an absence of duplex on leg tarsus IV. Based on the current designations, the latter species/population belongs to the subgenus Lyobia (Bryobia). The authors further stated that this species was similar to the European B. (B.) ribis Thomas, 1896. The latter is poorly described and has been suggested as a synonym of B. (B.) praetiosa (Pritchard and Baker 1955). This population of B. (B.) neoribis should be reidentified based on type material examination to reach a valid species designation.

The two species, B. (B.) neoribis sensu Tuttle and Baker (1976) and B. (B.) ribis are morphologically close. Mathys (1957) provided detailed morphological analysis and bioecological aspects of the species B. (B.) ribis and other Bryobia species found in the French part of Switzerland. The species B. (B.) neoribis was differentiated from B. (B.) ribis based on the number of setae on the femur I (24 vs 16) and variations in body and setal lengths. Tuttle and Baker (1976) did not provide a comprehensive description of the species, preventing detailed comparison and validation with other Bryobia species. Similarly, there is no detailed description and illustration of B. (B.) ribis. Mathys (1957) stated that complementary morphological differences could be found in the larval stage of B. (B.) ribis. This raises doubts over the validity of B. (B.) neoribis as only the female stage was briefly described. It would require a comprehensive set of specimens from the type locality to validate the status of the B. (B.) neoribis. For the time being, both species are excluded from the diagnostic key.

The species B. (B.) xizangensis Wang, 1985 was described from China from an unknown host plant. This species was originally differentiated from B. (L.) longisetis Reck, 1947 and was described with one or two pairs of tenent hairs on leg empodium I. Based on the findings of the present study, this species could be morphologically close to B. (B.) hengduanensis Wang & Cui, 1991 due to one pair of tenent hairs present on empodium I, but differentiated based on the length of dorsal body hairs, short vs long, crossing the bases of setae next in line, respectively. Considering the two pairs of tenent hairs on empodium I, B. (B.) xizangensis is similar to B. (B.) ziziphorae Strunkova & Mitrofanov, 1983, but is easily differentiated based on the development of propodosomal lobes, strongly developed and deep incision between the inner and outer lobes vs. weakly developed with small lobes, respectively.

Lyobia Livschits & Mitrofanov, 1971

Type species.

Bryobia rubrioculus (Scheuten) 1857: 104.

Diagnosis

(based on females). As defined by Mirza et al. (2024).

Key to the 51 species of the genus Lyobia

Species groups definition is based on Mirza et al. 2024.

1 Dorsocentral setae f1 present centrally, aligned with dorsocentral setae c1 eurotiae species group 3
– Dorsocentral setae f1 present laterally or sublaterally 2
2 Dorsocentral setae f1 present sublaterally, distance f1-f1 shorter than f2-f2 sarothamni species group 4
– Dorsocentral setae f1 present laterally, distance f1-f1 longer than f2-f2 rubrioculus species group 10
3 Dorsal body setae sit on distinct tubercles; tibia II with 6 tactile setae B. (L.) pamirica Mitrofanov, 1973
– Dorsal body setae sit on indistinct tubercles; tibia II with 9 tactile setae B. (L.) eurotiae Mitrofanov, 1973
4 Leg empodium I with 1 pair of tenent hairs B. (L.) chrysocomae Meyer, 1974
– Leg empodium I with > 1 pair of tenent hairs 5
5 Dorsocentral setae longer than the distance to bases of setae next in line 6
– Dorsocentral setae distinctly shorter than the distance to bases of setae next in line 7
6 Palp of tarsus distinctly longer than tibial claw palp B. (L.) perinsignis Eyndhoven & Vacante, 1985
– Palp of tarsus equal to tibial claw palp B. (L.) nasrvasensis Bagdasarian, 1960
7 Stylophore rounded anteriorly 8
– Stylophore emarginate anteriorly 9
8 Dorsal body setae lanceolate, broad distally B. (L.) sarothamni Geijskes, 1939
– Dorsal body setae slender B. (L.) annatensis Manson, 1967
9 Palptibial claw bidentate B. (L.) polymorpha Auger & Migeon, 2023 (in Auger et al. 2023)
– Palptibial claw simple B. (L.) spica Pritchard & Keifer, 1958
10 Propodosoma with reticulation pattern 11
– Propodosoma without reticulation pattern 14
11 Dorsal body setae fan-shaped (Fig. 6b) or spatulate or subspatulate (Fig. 6c) 12
– Dorsal body setae palmate; opisthosoma with 7 large oval dimple-like depressions with rounded reticulations B. (L.) alveolata Auger & Flechtmann, 2009
12 Opisthosoma with 3 pairs of oval depressions 13
– Opisthosoma without oval depressions; empodium I with 2 rows of tenent hairs B. (L.) dianthi Mitrofanov & Sharonov, 1983
13 Propodosoma with 2 large oval lateral depressions; empodium I with 3 pairs of tenent hairs B. (L.) hadizeni Barbar, Parker & Auger, 2022
– Propodosoma without oval depressions; empodium I with 1 pair of tenent hairs B. (L.) mercantourensis Auger & Migeon, 2014
14 Most of dorsal body setae spatulate 15
– Dorsal body setae not as above 38
15 Empodium I with 1 pair of tenent hairs 16
– Empodium I with ≥ 1 pairs of tenent hairs 25
16 Dorsocentral setae c1 and d1 elongate crossing half distance between next setae or reaching basis of next setae 17
– Dorsocentral setae c1 and d1 short not crossing half distance between next setae 18
17 First and second pairs of propodosomal setae v1 and v2 equal in length; c1 and d1 reaching or almost reaching bases of next setae B. (L.) dubinini Bagdasarian, 1960
– First pair of propodosomal setae v1 distinctly shorter than second pair v2; c1 and d1 reaching half distance between next setae B. (L.) longisetis Reck, 1947
18 Stylophore notched 19
– Stylophore rounded 23
19 Second and fourth pairs of propodosomal setae v2 and sc2 spatulate 20
– Second and fourth pairs of propodosomal setae v2 and sc2 palmate; tibia I with 15 setae B. (L.) kassioticus Hatzinikolis & Panou, 1997
20 Area posterior seta e1 with U-shaped striation; propodosoma granulate with irregular longitudinal striae B. (L.) siliquae Hatzinikolis & Emmanouel, 1991
– Area posterior setae e1 with irregular or transverse striae, not U-shaped 21
21 Second pair of propodosomal lobes well developed 22
– Second pair of propodosomal lobes absent or poorly developed; femur II with 10 setae B. (L.) syriensis Barbar & Auger, 2020
22 Median propodosomal lobes well developed B. (L.) baroni Auger, Arabuli & Migeon, 2022
– Median propodosomal lobes weakly developed or fused B. (L.) populi Wang & Zang, 1984
23 Femur II with 12 setae B. (L.) cerasi Hatzinikolis & Emmanouel, 1991
– Femur II with < 12 setae 24
24 Dorsum granulate with irregular striae; femur II with 9 setae B. (L.) dikmenensis Eyndhoven & Vacante, 1985
– Dorsum granulate without striae; femur II with 8 setae B. (L.) piliensis Hatzinikolis & Emmanouel, 1996
25 Median propodosomal lobes expanded and slightly overlapping; stylophore rounded B. (L.) berlesei Eyndhoven, 1957
– Median propodosomal lobes not overlapping 26
26 First and second pairs of propodosomal setae v1 and v2 are equal in length 35
– First pair of propodosomal setae v1 shorter than second pair v2 27
27 Setae c3 in line with setae c1 and c2 28
– Setae c3 not in line with setae c1 and c2 31
28 Tibia I with 9 setae; genua I and II each with 4 setae B. (L.) nothofagi Gonzalez, 1977
– Tibia I with 12 setae or more; genua I and II each with ≥ 4 setae 29
29 Tibia I with 2 sensory setae 30
– Tibia I with 3 sensory setae; femora II and III with 8 and 5 setae, respectively B. (L.) mirmoayedii Khanjani, Gotoh & Kitashima, 2008
30 Femora II and III with 10 and 7 setae, respectively B. (L.) cooremani Eyndhoven & Vacante,
– Femora II and III with 7 and 4 setae, respectively B. (L.) vaneyndhoveni Vacante, 1983
31 Setae c3 below setae c2 B. (L.) eharai Pritchard & Keifer, 1958
– Setae c3 above setae c2 32
32 Tarsus I with 8 sensory setae 33
– Tarsus I with 7 sensory setae; genu I with 6 setae 34
33 Femora I and IV with 13–15 and 4 or 5 setae, respectively B. (L.) cavalloroi Vacante & Eyndhoven, 1986
– Femora I and IV with 10–12 and 6 setae, respectively B. (L.) ulicis Eyndhoven, 1959
34 Dorsum integument granulate with irregular striae B. (L.) pyrenaica Eyndhoven & Vacante, 1985
– Dorsum integument granulated B. (L.) strombolii Vacante, 1983
35 Setae c3 located above setae c2 36
– Setae c3 located below setae c2; empodia I with 2 pairs of tenent hairs B. (L.) chongqingensis Ma & Yuan, 1981
36 Genu I with 7 or 8 setae; femora II and III with 9 and 7 setae, respectively B. (L.) aetnensis Vacante, 1983
– Genu I with < 6 setae 37
37 Genu I with 6 setae; femur III with 6 setae B. (L.) dekocki Eyndhoven & Vacante, 1985
– Genu I with 4 or 5 setae; femur III with 7 setae B. (L.) vandaelei Vacante, 1983
38 Dorsal body setae palmate or fan-shaped 39
– Dorsal body setae lanceolate or slender 46
39 Peritremes with simple bulb distally 40
– Peritremes anastomose distally 42
40 Dorsal body setae palmate; empodium I with 1 pair of tenent hairs B. (L.) convolvulus Tuttle & Baker, 1964
– Dorsal body setae fan-shaped 41
41 Empodium I with 1 pair of tenent hairs B. (L.) oblonga Livschits & Mitrofanov, 1968
– Empodium I with ≥ 1 pairs of tenent hairs B. (L.) kakuliana Reck, 1956
42 Dorsocentral setae c1-c1, d1-d1, e1-e1 very close to each other B. (L.) angustisetis Jakobashvili, 1958
– Dorsocentral setae c1-c1, d1-d1, e1-e1 widely spaced 43
43 Femur I with 23 setae; tibia I with 14 setae B. (L.) parietariae Reck, 1947
– Femur I with not more than 21 setae 44
44 Tibia I with 12 setae B. (L.) centaureae Livschits & Mitrofanov, 1972
– Tibia I with 15–16 setae 45
45 Femur II with 9 setae B. (L.) rubrioculus (Scheuten, 1857)
– Femur II with 11 setae B. (L.) tiliae (Oudemans, 1928)
46 Dorsal setae lanceolate (Fig. 6d) 47
– Dorsal setae slender, long at least reaching (Fig. 6e), empodium I with 2 rows of tenent hairs B. (L.) cinereae Auger & Migeon, 2014
47 Tibiae III-IV each with 9 setae 48
– Tibiae III-IV each with < 9 setae; femur I with 12 setae or fewer 49
48 Femur I with 20 or 22 setae B. (L.) gushariensis Livschits & Mitrofanov, 1972
– Femur I with 13 or 18 setae B. (L.) obihsaphedi Mitrofanov, 1968
49 Femur I with 12 setae; tibiae III-IV each 7 setae B. (L.) livschitzi Mitrofanov & Strunkova, 1968
– Femur I with 8 or 9 setae; tibiae III-IV each with < 7 setae 50
50 Setae on femora I-IV 9–7–5–3; tibiae III-IV each with 4 setae B. (L.) astragali Strunkova & Mitrofanov, 1983
– Setae on femora I-IV 8–7–4–2; tibiae III-IV with 3 and 5 setae, respectively B. (L.) bucharica Strunkova & Mitrofanov, 1983

Notes on the species of the subgenus Lyobia

The subgenus Lyobia includes 58 species (Mirza et al. 2024) but the key to only 51 species is provided above. The species B. (L.) ericoides Meyer, 1974, belonging to the species group eurotiae, is excluded from the key due to leg I true claw morphology. The status of the remaining six species, all belonging to the species group rubrioculus, are discussed below.

Species group eurotiae

The two species in this species group B. (L.) eurotiae Mitrofanov, 1973 and B. (L.) pamirica Mitrofanov, 1973, are morphologically similar and share the type host plant (Eurotia sp.), type locality (Tadjikistan), and date of collection (23 July 1967). These two species share most morphological characteristics, including similar body length and width, lacking propodosomal lateral lobes, setae v2 longer than v1, slender setiform setae, length of leg I equal to body length, number of tenant hairs on leg empodia I-IV, and most of the leg chaetotaxy. The morphological characters which differentiate B. (L.) eurotiae from B. (L.) pamirica include state of propodosomal lobes (completely absent vs inner lobes joined from the middle, forming a cone), dorsal setal tubercles (indistinct vs distinct), leg chaetotaxy of femora I-III (9-7-4 vs 8-6-3), genua I and II (8-5 vs 4-4), and tibia II (9 vs 6), respectively. The differences in leg chaetotaxy mentioned above should be re-examined and could be considered as variations. The original description of B. (L.) eurotiae provides leg chaetotaxy in which tibia I has 24 setae. It appears that the setal count of tibia I was missed, and the setal counts for tarsus I were provided. It could be assumed that there are 24 setae on tarsus I, which was also described for B. (L.) pamirica because the chaetotaxy of tibiae II-IV is similar in both species. Similarly, the setae f1 were described to be present sublaterally in B. (L.) pamirica, while they are illustrated as aligned with dorsocentral setae c1. Hence, the setae f1 are present centrally or subcentrally in B. (L.) pamirica, similar to B. (L.) eurotiae. Although there is evidence for the possible synonymy of these two species, it is important to re-examine the type specimens to reach a definitive conclusion.

Species group sarothamni

There are seven species in this species group (Mirza et al. 2024). The morphology of propodosomal lobes has been described with variations. For instance, B. (L.) sarothamni Geijskes, 1939, was originally described from the Netherlands, with the presence of four propodosomal lobes in the form of tubercles (Geijskes 1939). Pritchard and Baker (1955) distinguished the English population of B. (L.) sarothamni with a complete absence of “cephalic projections”. Baker and Tuttle (1994) reported the presence of the propodosomal projection, where outer ones were as broad as long, and 1/3 as long as the inner pair. This situation is similar to that in the praetiosa species complex (in the subgenus Bryobia (B.)). It is recommended to approach the species in this species group with extreme caution, and morphological variations should be completely understood before describing new taxa.

Species group rubrioculus

There are 48 species included in this species group (Mirza et al. 2024). The species B. (L.) cinereae Auger & Migeon, 2014 was placed in the species group sarothamni (Mirza et al. 2024). However, in the present study, it is included in the species group rubrioculus due to the marginal position of sacral f1 and f2 setae. This species is morphologically close to B. (L.) belliloci Auger, Arabuli & Migeon, 2015; however, the morphological differences are debatable. It has been stated that setae d1 clearly surpass the bases of e1 in B. (L.) belliloci (illustrated as just passing) while setae d1 just reach the base of setae e1 in B. (L.) cinereae (Auger and Migeon 2014). There are other morphological characters which were used to differentiate B. (L.) belliloci from B. (L.) cinereae including the depth of the inner lobe incisions (but illustrated as exactly same for both species), peritremal distal enlargement length (both anastomosing but length has 7 μm difference), length of internal seta l’1 on femur I, lengths and shapes of coxal setae 1b and 1c (discrepancies in the description and illustrations of B. (L.) belliloci). These characters may reflect variations in the morphologies, especially when both species have the same host plant, Genista cinerae, and are both reported from France (Auger and Migeon 2014; Auger et al. 2015). The species B. (L.) belliloci is excluded from the key, and perhaps further studies may suggest it as a junior synonym of B. (L.) cinereae.

The four species B. (L.) tiliae (Oudemans, 1928; Germany), B. (L.) rubrioculus (Scheuten, 1857; Germany), B. (L.) lonicerae Reck, 1956 (Georgia), and B (L.) ulmophila Reck, 1947 (Georgia), are very similar to each other in all morphological aspects including leg morphology. The species B. (L.) rubrioculus has been described and illustrated from different regions of the world and number of species have been synonymized under it (Migeon and Dorkeld 2025). Frommer and Jorgensen (1972) studied the morphological and behavioral variations with host specificity of B. (L.) rubrioculus and distinctly separated this species from B. (L.) praetiosa. The two species B. (L.) lonicerae and B. (L.) ulmophila were morphologically compared with B. (L.) redikorzevi that is considered a synonym of B. (L.) rubrioculus by Frommer and Jorgenson (1972). Wainstein (1960) considered B. (L.) ulmophila as synonym of B. (L.) redikorzevi. The species B. (L.) tiliae was originally described as a type species of the genus Schmiedleinia Oudemans, 1928, based on the larval specimens collected from the host plant Tiliae sp. in Germany (Oudemans 1928). The genus was later synonymized with the genus Bryobia, and the species tiliae was considered as the larvae of B. praetiosa (Oudemans 1930). Bagdasarian (1957) described the species B. (L.) tiliae from Armenia on the same host plant, Tiliae sp. It was later considered a synonym of the species described by Oudemans (1928) (Wainstein 1960). In that synonymy, B. (L.) tiliae was considered to be morphologically close to B. (L.) ulmophila and B. (L.) redikorzevi (Bagdasarian 1957) but distinguished based on the number of setae on leg femur I. Both of the latter two species have been considered as a synonym of B. (L.) rubrioculus. The leg chaetotaxy alone would not be sufficient to confidently validate the identity of the species. In light of this debate, it would be difficult to reach any definitive conclusion regarding the validity of these three species, and their synonymy with B. (L.) rubrioculus requires further investigation.

Eyndhoven and Vacante (1985) described 13 species belonging to the berlesei species group, eight of which were described for the first time. Among them, five species B. (L.) pandayi Eyndhoven & Vacante, 1985, B. (L.) pyrenaica Eyndhoven & Vacante, 1985, B. (L.) pelerentsi Eyndhoven & Vacante, 1985, B. (L.) dikmenensis Eyndhoven & Vacante, 1985, and B. (L.) provincialis Eyndhoven & Vacante, 1985, have variable morphological characters. The three species B. (L.) pandayi, B. (L.) pyrenaica, and B. (L.) pelerentsi were considered close to each to other and the differential character designated as “Each species has its own host plant” (Eyndhoven and Vacante 1985: 400). In all other morphological aspects, these three species resemble each other, and it is difficult to differentiate them. The remarks for these species were stated as “For general remarks see Bryobia pandayi”. It is important to mention that a species having its own host plant does not necessitate its validity. The species B. (L.) pandayi was reported from Calicotome spinosa. The same host plant (Calicotome sp.) harbors almost seven Bryobia taxa (Migeon and Dorkeld 2025). Interestingly, B. (L.) pelerentsi is also reported from Calicotome sp. (Eyndhoven and Vacante 1985). Hence, with this argument, the synonymy of B. (L.) pyrenaica and B. (L.) pelerentsi with B. (L.) pandayi appears undeniable. Similarly, both species, B. (L.) dikmenensis and B. (L.) provincialis are reported from the same host plant (Genistus sp.) and were morphologically designated close to each other by Eyndhoven and Vacante (1985). The only morphological difference described was that the second and third dorsocentrals were smaller than the other dorsal body setae in B. (L.) dikmenensis, while of similar length in B. (L.) provincialis. However, this contradicts what has been described for these setae based on 14 specimens (Eyndhoven and Vacante 1985). This places the status of these species as doubtful, and there is an urgent need for re-analysis of the morphological characters of these species.

Conclusions

In conclusion, the present study provides a comprehensive taxonomic status of the species of Bryobia through detailed literature-based morphological analysis. The diagnostic keys to the majority of Bryobia species will undoubtedly prove useful for acarologists. The taxonomic notes on some species and the variability in morphological characters found among different populations of a species deepen our understanding of morphological diversity in the genus. It is important to note that there are four species: B. apsheronica Khalilova, 1953, B. desertorum Hassan, Afifi & Nawar, 1986, B. ribis Thomas, 1896, and B. weyerensis Packard, 1889 that are not included in any subgenus or species group due to inadequate and insufficient literature, as also reported by Mirza et al. (2024). The species B. weyerensis may not even belong to the family Tetranychidae, while the former three species should be redescribed. Although some species have been suggested to be synonymized with closely related species, a valid taxonomic and systematic decision should be backed and supported by careful examination of the type specimens. In the scenario where types have been lost, a collection should be initiated to revisit the type locality.

Acknowledgments

The authors would like to thank Carlos Holger Wenzel Flechtmann for sharing the complete literature of all Bryobia species. The authors would like to extend their appreciation to the two respected reviewers whose constructive comments improved the overall quality of the research and to Nathalie Yonow, the copy editor, who thoughtfully edited our paper.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was funded by Researchers Supporting Project [number RSPD2025R807], King Saud University, Riyadh, Saudi Arabia.

Author contributions

Conceptualization, JHM and FJA; methodology, JHM, NAE and FJA; validation, FJA and MK; formal analysis, JHM and NAE; investigation, JHM and NAE; data curation, JHM and NAE; writing—original draft preparation, JHM and NAE; writing—review and editing, FJA and MK; funding acquisition, FJA. All authors have read and agreed to the published version of the manuscript.

Author ORCIDs

Jawwad Hassan Mirza https://orcid.org/0000-0002-1726-4331

Nasreldeen Ahmed Elgoni https://orcid.org/0009-0007-7698-1377

Muhammad Kamran https://orcid.org/0000-0001-6084-203X

Fahad Jaber Alatawi https://orcid.org/0000-0002-6824-2650

Data availability

All of the data that support the findings of this study are available in the main text.

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