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Research Article
Species groups, subgroups, and key to world species of the genus Schizotetranychus Trägårdh, 1915 (Acari, Prostigmata, Tetranychidae)
expand article infoMuhammad Kamran‡, Abdul Hafeez§, Fahad Jaber Alatawi‡, Carlos Holger Wenzel Flechtmann|
‡ King Saud University, Riyadh, Saudi Arabia
§ Ghazi University, Dera Ghazi Khan, Pakistan
| Universidade de São Paulo, São Paulo, Brazil

Corresponding author: Muhammad Kamran ( murafique@ksu.edu.sa )

Academic editor: Vladimir Pesic
© 2024 Muhammad Kamran, Abdul Hafeez, Fahad Jaber Alatawi, Carlos Holger Wenzel Flechtmann.
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: Kamran M, Hafeez A, Alatawi FJ, Flechtmann CHW (2024) Species groups, subgroups, and key to world species of the genus Schizotetranychus Trägårdh, 1915 (Acari, Prostigmata, Tetranychidae). ZooKeys 1211: 131-150. https://doi.org/10.3897/zookeys.1211.127353
ZooBank: urn:lsid:zoobank.org:pub:F1E24C8D-23B5-444C-8526-91EF52BF27DC
Open Access

Abstract

After a comprehensive taxonomic assessment of descriptions/ illustrations of all known (118) species of the spider mite genus Schizotetranychus Trägårdh (Acari: Prostigmata: Tetranychidae), five species groups are proposed based on the number of tactile setae on tibia II in female, i.e., vermiculatus with four setae (four spp.), schizopus with five setae (52 spp.), spireafolia with six setae (10 spp.), asparagi with seven setae (20 spp.), and bambusae with eight setae on tibia II (22 spp.). The species group schizopus is further divided into three species subgroups based on tactile setae on tibia I: schizopus with eight/ nine setae (21 spp.), andropogoni with seven setae (26 spp.), and taquarae with six tactile setae excluding the solenidion on tibia I (five spp.). Eight Schizotetranychus species were not assigned to any species group because of brief descriptions and/ or illustration and without information on the number of tactile setae on tibiae I and II. Moreover, two Schizotetranychus species, S. gausus Baker & Pritchard and S. luculentus Tseng that have six setae/ structures including a spinneret and a solenidion on the palp tarsus, are provisionally transferred to the genus Stigmaeopsis Banks. Finally, keys to species groups and subgroups of the world species of Schizotetranychus are provided.

Key words

Asparagi, schizopus, tactile setae, taquarae, vermiculatus

Introduction

The genus Schizotetranychus Trägårdh (Acari, Prostigmata, Tetranychidae) was erected by Trägårdh (1915) based on the shape of leg empodia, i.e., divided deeply into two claw-like structures and having ten pairs of dorsal hysterosomal setae. It is one of the largest genera of spider mites containing 118 species, widely distributed in the world (Migeon and Dorkeld 2006–2024). Schizotetranychus species are phytophagous on different plant species and some species are considered as pests of agricultural crops, i.e., Schizotetranychus andropogoni (Hirst) and Schizotetranychus asparagi (Oudemans, 1928) are widespread in United States and Europe causing serious infestations to pineapple plants (Jeppson et al. 1975; Hoy 2011).

Schizotetranychus species identity has been challenging due to the inadequate number of diagnostic characters, minute differences in male aedeagus morphology, and interspecific similarities in females of many species. Specimens of both sexes are usually required for accurate identification of Schizotetranychus species (Pritchard and Baker 1955; Meyer 1974, 1987; Jeppson et al. 1975; Flechtmann 2012). Pritchard and Baker (1955) and Meyer (1974, 1987) provided only diagnostic keys of Schizotetranychus species occurring in some regions. Lastly, Flechtmann (2012) arbitrarily organized the Schizotetranychus species into 17 groups only based on female morphology; however, the identification characters used are confusing. Additionally, a diagnostic key to world Schizotetranychus species is not available. The present study aimed to classify the species of the genus Schizotetranychus into species group and subgroups based on consistent diagnostic morphological characters, and to develop diagnostic keys to species groups and subgroups of world species of the genus Schizotetranychus.

Materials and methods

The taxonomic literature of all known Schizotetranychus species were critically studied, and the diagnostic characters were compared. The generic characters of Schizotetranychus and Stigmaeopsis were carefully analyzed for possible new combinations. The strength of each morphological character was evaluated for its suitability at the specific level. The consistency of tibia I and II setal counts were carefully evaluated for the construction of species and sub-species group. The key to species of the genus Schizotetranychus is provided based on persistent and fixed characteristics.

Results and discussion

Subfamily Tetranychinae Berlese

Tribe Tetranychini Reck

Schizotetranychus Trägårdh

Type species

Tetranychus schizopus Zacher, 1913.

Diagnosis

Dorsal hysterosoma with ten pairs of setae (c1-3, d1-2, e1-2, f1-2, and h1), setae h2 and h3 present on ventral opisthosoma, empodial claw divided deeply into two claw-like structures, palp tarsus with seven structures/ setae (one spinneret, two eupathidia, one solenidion, three setiform setae); dorsal hysterosoma medially usually with transverse striations, but may be longitudinal or irregular between d1 and e1; two sets of duplex setae on tarsus I present distally, nearly adjacent to each other.

Background and taxonomic review of the genus Schizotetranychus

The genus Schizotetranychus was erected by Trägårdh (1915) based on Tetranychus schizopus Zacher, 1913 and distinguished from the genus Tetranychus by having the leg empodia divided deeply into two claw-like structures. Two years later, Banks (1917) erected the genus Stigmaeopsis and designated St. celarius Banks its type species. Banks (1917) described the genus Stigmaeopsis very briefly and did not provide any diagnostic characters which could separate it from the closely related genus Schizotetranychus. Later, McGregor (1950) synonymized the monospecific genus Stigmaeopsis with Schizotetranychus. This synonymy was accepted and followed by Pritchard and Baker (1955), Baker and Pritchard (1960), Tuttle and Baker (1968), Gutierrez (1968), Meyer (1974, 1987), Tuttle et al. (1976), Bolland et al. (1998), and Ehara (1999).

For the first time, Ehara (1999) introduced species groups in the genus Schizotetranychus by dividing the nine Schizotetranychus species reported from Japan into two species groups: schizopus (six species) with transverse striations in the anterior portion of dorsocentral area on dorsal opisthosoma and celarius (three species) with longitudinal striations in the anterior portion of dorsocentral area on dorsal opisthosoma.

Later, Saito et al. (2004) reinstated the genus Stigmaeopsis and distinguished it from Schizotetranychus and other genera of Tetranychidae by having six setae/ structures on the palp tarsus in the female (instead of seven); dorsal striations between c1 and d1, clearly longitudinal, forming a trapezoidal shape instead of having mostly transverse or longitudinal irregular without forming a trapezoidal shape in Schizotetranychus. Also, the bases of setae e1, d1, and c1 gradually become further apart than the bases of f1 setae; if hypothetical lines connecting their bases are drawn, they form a V-shaped pattern vs these lines being almost parallel in Schizotetranychus and related genera, as described by Saito et al. (2004). Based on these characteristics, Flechtmann (2012) transferred two Schizotetranychus species, S. malkovskii Wainstein, 1956 and S. meghalensis (Gupta & Gupta, 1994) to Stigmaeopsis. Although S. meghalensis has transverse striations between setae c1, d1, and e1, and does not satisfy several characters of Stigmaeopsis, despite the fact that Saito et al. (2016, 2018, 2019) provisionally included this species in Stigmaeopsis because of its six setae/ structures on the palp tarsus.

Morphological diagnostic features previously used for grouping Schizotetranychus species

Flechtmann (2012) categorized world 106 Schizotetranychus species into 17 groups based on following female morphological characters: body length: width ratio, dorsal setal length, shape of peritremes, number of tactile setae on tibia I. As a result, numerous species groups in the genus Schizotetranychus based on variable morphological characters are causing confusion and misunderstanding in species identification.

  1. Peritremes distally are variously developed in Schizotetranychus species, straight in most species, and either making a U-shape, ring, or looped distally others. Peritremes distally are anastomosing in two species S. cajani Gupta, 1996, S. prosopis Tuttle, Baker & Abbatiello, 1976. Flechtmann (2012) used this character to arbitrarily propose different groups for Schizotetranychus species. We consider the shape of the peritreme at species level a misleading character because it is variously developed distally even in different specimens of the same species, and described and illustrated differently for one species by various authors. Also, this character was already causing confusion while attempting to separate the species groups of Schizotetranychus created by Flechtmann (2012).
  2. Mite body shape is either oval (longer than wide in most of species), or orbicular (as long as wide in few species), or elongate (more than 2× longer than width of body in few species). This character was used by Flechtmann (2012) to develop groups in Schizotetranychus. However, it caused confusion in the identification of those groups because some species lie on the borderline in length and width ratios. Also, the length and width ratios could be affected by the mounting of specimens on glass slides.
  3. Dorsal and ventral idiosoma is entirely striated in almost all Schizotetranychus species, either widely or closely spaced, except S. reticulatus Baker & Pritchard, 1960 with reticulations on the propodosoma medially, and the hysterosoma is entirely striated and rugose. The dorsal hysterosoma between setae c 1 , d 1 , e 1 , and f 1 with transverse striations entirely in all Schizotetranychus species except six, namely S. hidayahae Yusof & Zhang, 2003, S. baltazarae Rimando, 1962, S. spiculus Baker & Pritchard, 1960, S. brevisetosus Ehara, 1989, S. rhodanus Baker & Pritchard, 1960, and S. colocasiae Ehara, 1988 (as in Ehara & Tho, 1988), in which the striations between setae e 1 and d 1 form a V-shaped pattern or is irregular.

Dorsal body setae are usually setiform in Schizotetranychus species. However, few species have awl-shaped dorsal setae with slightly expanded bases. Flechtmann (2012) used this character to develop species groups in Schizotetranychus. However, dorsal setae were not properly illustrated or described in detail for many Schizotetranychus species, so for those species it is very difficult to discern the exact shape (awl or setiform) of the dorsal setae. This character can be considered as supporting species level character.

Taxonomic notes about two Schizotetranychus species having six setae/ structures on the palp tarsus as in the genus Stigmaeopsis

As mentioned earlier, the genus Stigmaeopsis differs from Schizotetranychus by having six setae/ structures on the palp tarsus in females instead of seven; dorsal striations lie between c1 and d1 are clearly longitudinal and forming a trapezoidal shape instead of being mostly transverse or irregularly longitudinal between setae d1 and e1 in six Schizotetranychus species, namely, S. hidayahae, S. baltazarae, S. spiculus, S. brevisetosus, S. rhodanus and S. colocasiae without forming a trapezoidal shape. Also, the bases of setae e1, d1, and c1 gradually widen further apart than the bases of f1 setae if hypothetical lines connecting their bases are drawn. They form a V-shaped pattern vs almost parallel lines are in Schizotetranychus and related genera, as described by Saito et al. (2004).

In the present study, it was found that two Schizotetranychus species, S. gausus Baker & Pritchard, 1960 and S. luculentus (Tseng, 1990) have six setae/ structures including spinneret and solenidion on palp tarsus. The original description of these species lacking information of palp setae. So, relying on the original illustrations, these two Schizotetranychus species having six setae on palp tarsus are provisionally transferred to Stigmaeopsis. Also, dorsum is entirely reticulated in S. luculentus (Tseng, 1990). However, dorsum with transverse striations between setae c1, d1 and irregular longitudinal between setae e1 and d1 in S. gausus. Moreover, 16 known species of Stigmaeopsis have five tactile setae on tibia II except S. gausus having seven setae on tibia II.

Furthermore, bases of length dorsal setae c1, d1 which is ~ 2× more widely spaced to the bases of e1 and f1 (bases of c1, d1, e1, f1 forming a V-shaped pattern) in S. Attiah, 1967 as in all known 16 Stigmaeopsis species. Few other Schizotetranychus species have a similar pattern of dorsal setal bases. So, the supporting diagnostic character that hypothetical lines connecting the bases of setae c1, d1, e1, and f1 forming a V-shaped taken by Saito et al. (2004) for Stigmaeopsis while reinstating this genus to separate it from Schizotetranychus becomes impractical.

Hence it is understood from the above discussion that genus Stigmaeopsis is different from Schizotetranychus by only one character, the presence of six setae/ structures on palp tarsus vs seven in Schizotetranychus. All other supporting characters (longitudinal striations between setae c1 and d1, bases of setae c1, d1, e1, and f1 forming a V-shaped pattern) of Stigmaeopsis taken by Saito et al. (2004, 2018) as a generic character could be considered as species level characters.

Species groups and subgroups of Schizotetranychus developed in the current study

In the present research, after comprehensive taxonomic assessment of descriptions and illustrations of all known (116) species of the genus Schizotetranychus, species grouping in this genus is reconsidered based on females using only the number of tactile setae on tibia II and species subgroups based on only the number of tactile setae on tibia I. The number of tactile setae on tibia II is found to be a consistent diagnostic character in Schizotetranychus species and described in 110 Schizotetranychus species, even those which were very briefly described. Flechtmann (2012) used tactile setae on tibia I to separate some Schizotetranychus groups. Pritchard and Baker (1955) and Mushtaq et al. (2021) also used tactile setae on tibia I to develop species groups in the genus Oligonychus. Based on tibial setal counts, species groups of Schizotetranychus can easily be recognized.

In the present study, the genus Schizotetranychus can be divided into five species groups based on the number of tactile setae on tibia II in the female: schizopus group (52 spp.) with five setae, asparagi group with seven setae (20 spp.), bambusae group with eight setae (22 spp.), spireafolia group with six setae (10 spp.) and vermiculatus group with four setae on tibia II (four spp.). Also, keys to the world Schizotetranychus species, species groups, and subgroups are developed for the first time. Eight Schizotetranychus species were not assigned any species group because these have been described and illustrated very briefly without information on the number of setae on tibia I and II.

1. Species group schizopus

Diagnosis. Female: Tibia II with five setae (52 species).

Exemplar species. Schizotetranychus schizopus (Zacher, 1913)

Species group schizopus is further divided into three species subgroups based on number of tactile setae excluding solenidion on tibia I.

i) Species subgroup schizopus

Diagnosis. Female. Tibia I with eight/ nine setae (21 species).

Exemplar species. Schizotetranychus schizopus

ii) Species subgroup andropogoni

Diagnosis. Female. Tibia I with seven setae (26 species).

Exemplar species. Schizotetranychus andropogoni (Hirst, 1926)

iii) Species subgroup taquarae

Diagnosis. Female. Tibia I with six setae (5 species).

Exemplar species. Schizotetranychus taquarae Paschoal, 1971

2. Species group asparagi

Diagnosis. Female. Tibia II with seven setae (20 spp.)

Exemplar species. Schizotetranychus asparagi (Oudemans, 1928)

3. Species group bambusae

Diagnosis. Female. Tibia II with eight setae (22 spp.).

Exemplar species. Schizotetranychus bambusae Reck, 1941

4. Species group spireafolia

Diagnosis. Female. Tibia II with six setae (10 spp.).

Exemplar species. Schizotetranychus spireafolia Garman, 1940

5. Species group vermiculatus

Diagnosis. Female. Tibia II with four setae (04 spp.).

Exemplar species. Schizotetranychus vermiculatus Ehara & Wongsiri, 1975

Ungrouped species

The following eight species were not assigned any species group because these have been described and illustrated very briefly without information of number of tactile setae on tibiae I and II. Schizotetranychus setariae Meyer, 1987 was not assigned to any species group/ subgroup because it was only described/ known from the male.

  1. S. graminicola Goux, 1949
  2. S. glabrisetus (Ugarov & Nikolskii, 1937)
  3. S. tuberculatus (Ugarov & Nikolski, 1937)
  4. S. guatemalae-novae (Stoll, 1886)
  5. S. hindustanicus (Hirst, 1924)
  6. S. mustafaii Mustafa & Chaudri, 1972 (as in Mustafa et al. 1972)
  7. S. oudemansi Reck, 1948
  8. S. setariae Meyer, 1987

Moreover, two Schizotetranychus species, S. gausus and S. luculentus, that have six setae/ structures including spinneret and a solenidion on the palp tarsus based on original illustrations, are provisionally transferred to Stigmaeopsis. Further studies are required to confirm the clear taxonomic status of these two species.

1. Schizopus species group (52 species)

I- Schizopus species subgroup (21 species)

(Female: tibia II with 5 setae, tibia I with 8 or 9 tactile setae excluding solenidion)

Key to species of schizopus species subgroup of schizopus species group

1 Propodosoma dorsomedially with reticulate pattern S. reticulatus Baker & Pritchard, 1960
– Propodosoma dorsomedially striated 2
2 Peritremes distally curved U- or ring-shaped/ looped 3
– Peritremes distally straight or slightly hooked 7
3 Tibia I with 9 tactile setae and 1 sensory seta S. australis Gutierrez, 1968
– Tibia I with 8 tactile setae and 1 sensory seta 4
4 Dorso-opisthosomal setae c1, d1, e1 at least reaching 1/2 to 2/3 of the setae next in line S. pennamontanus Meyer, 1987
– Dorso-opisthosomal setae c1, d1, e1 almost reaching or crossing the setae next in line 5
5 Male aedeagus upturned part not sigmoid, distal part not projecting posteriorly S. russeus Davis, 1969
– Male aedeagus upturned part sigmoid, distal part projecting posteriorly 6
6 In male: tarsus I with 3 and tibia I with 1 spindle-shaped setae; in female: setae c1, d1, e1 at least reaching the setae next in line S. eremophilus McGregor, 1950
– In male: tarsus I and tibia I without spindle-shaped setae; in female: setae c1, d1, e1 crossing the setae next in line S. elymus McGregor, 1950
7 Tibia I with 9 tactile setae and 1 solenidion 8
– Tibia I with 8 tactile setae and 1 solenidion 10
8 Male: aedeagus upturned part almost sigmoid, without anterior projection, tibia I with 7 tactile setae and 2 solenidia S. mansoni Gupta, 1980
– Male: aedeagus upturned part, not sigmoid, with anterior projection, tibia I with 9 tactile setae and 3 or 4 solenidia 9
9 Male: tibia I with 9 tactile setae and 3 solenidia, tibia II with 5 tactile setae S. schizopus (Zacher, 1913)
– Male: tibia I with 9 tactile setae and 4 solenidia, tibia II with 6 tactile setae S. lechrius Rimando, 1962
10 Female: genu III with 4 tactile setae 11
– Female: genu III with 3 tactile setae 14
11 Female: setae c1, d1, e1 far behind the bases of setae next in line S. agropyron Tuttle & Baker, 1976
– Female: setae c1, d1, e1 crossing the bases of setae next in line 12
12 Male: aedeagus upturned part at right or acute angle to the shaft, does not project posteriorly S. nesbitti Meyer, 1965
– Male: aedeagus upturned part making obtuse angle to the shaft, projecting posteriorly 13
13 Male: aedeagus not sigmoid, upturned part gradually narrowing toward distal end, 1.5× longer than max. width of shaft S. cynodonis McGregor, 1950
– Male: aedeagus almost sigmoid, upturned part abruptly narrowing toward distal end, less than max. width of shaft S. paezi Alvarado & Freitez, 1976
14 Female: femur I with 7 or 8 setae 15
– Female: femur I with 9 or 10 setae 17
15 Female: genu IV with 2 setae S. echinulatus Mitrofanov, 1978
– Female: genu IV with 3 setae 16
16 Dorso-central hysterosomal setae shorter than distance/interval to base of seta immediately behind S. saba-sulchani Reck, 1956
– Dorso-central hysterosomal setae as long as or longer than distance/interval to base of seta immediately behind S. yoshimekii Ehara & Wongsiri, 1975
17 Femur I with 9 setae 18
– Femur I with 10 setae 19
18 Female: tibia III with 5 setae. Male: aedeagus upturned part with knob, prominent neck, and anterior projection S. hilariae Tuttle & Baker, 1968
– Female: tibia III with 5 setae. Male: aedeagus upturned part without knob, neck, or anterior projection S. tuttleii Zaher, Gomaa & El-Enany, 1982
19 Female: femur III with 2 setae, peritremes L-shaped distally. Male: aedeagus upturned part length of shaft less than max. width of shaft S. kochummeni Ehara, 1988 (as in Ehara and Tho 1988)
– Female: femur III with 3 setae, peritremes straight distally. Male: aedeagus upturned part of shaft length > 2× the max. width of shaft 20
20 Female: femur IV with 2 setae. Male: aedeagus upturned part almost at right angle to the shaft, not projecting posteriorly S. tbilisiensis Reck, 1959
– Female: femur IV with 3 setae. Male: aedeagus upturned part making obtuse angle (120°) to the shaft, projecting posteriorly S. tumidus Wang, 1981

II- andropogoni species subgroup (26 species)

(Female: tibia II with 5 setae, tibia I with 7 tactile setae excluding solenidion)

Key to species of andropogoni species subgroup

1 Peritremes distally hooked, branched, or looped 10
– Peritremes distally simple (bulb-like) 2
– Peritremes distally anastomosing S. cajani Gupta, 1996
2 Dorsal setae comparatively shorter in length, setae c1 reaching at least 2/3 the distance c1-d1 3
– Dorsal setae comparatively long, setae c1 reaching/ crossing the bases of d1, almost equal to/ longer than the distance c1-d1 7
3 Idiosoma elongate, ratio of body length (not including rostrum): width > 2 4
– Idiosoma oval/ orbicular, ratio of body length (not including rostrum): width < 2 5
4 Tarsus I with 3 tactile setae and 1 solenidion proximal to proximal duplex setae S. lycurus Tuttle & Baker, 1964
– Tarsus I with 1 tactile seta and 1 solenidion proximal to proximal duplex setae S. boutelouae Tuttle & Baker, 1968
5 Femur I with 7 setae, area anterior to genital flap with transverse striations, tarsus I with 2 tactile setae and 7 solenidion proximal to proximal duplex setae S. celtidis Tuttle & Baker, 1968
– Femur I with 9 setae, area anterior to genital flap with longitudinal striations, tarsus I with 3 tactile setae and 1 solenidion proximal to proximal duplex setae 6
6 Genu III with 2 setae, genu IV with 1 seta, area on hysterosoma between d1 and e1 with longitudinal striations S. hidayahae Yusof & Zhang, 2003
– Genua III and IV each with 3 setae, hysterosoma dorsomedially with transverse striations entirely S. montanae Tuttle & Baker, 1968
7 Setae c1 very long, crossing the bases of d1, reaching to the bases of e1 S. longirostris Feres & Flechtmann, 1995
– Setae c1 reaching at least to the bases of d1, almost equal to the distance c1-d1 8
8 Area anterior to genital flap with longitudinal irregular striations S. paraelymus Feres & Flechtmann, 1995
– Area anterior to genital flap with transverse striations 9
9 In female: seta sc1 much longer than sc2, setae d1 and e1 not reaching to the bases of setae next in line. Male: tarsus I with 3 solenidia and 1 tactile seta proximal to proximal duplex setae, aedeagus upturned part sigmoid without anterior projection S. camur Pritchard & Baker, 1955
– In female: setae sc1 and sc2 almost subequal, setae d1 and e1 reaching to the bases of setae next in line. Male: tarsus I with 1 solenidion and 1 tactile seta proximal to proximal duplex setae, aedeagus upturned part with anterior projection S. andropogoni (Hirst, 1926)
10 Dorso-central setae c1, d1, and e1 reaching or crossing the bases of next setae in line 11
– Dorso-central setae c1, d1, and e1 not reaching behind the bases of next setae in line 16
11 Female: tarsus I with 2 or 4 setae and 1 solenidion proximal to proximal duplex setae 12
– Female: tarsus I with 1 seta and 1 solenidion proximal to proximal duplex setae 14
12 Female: tarsus I with 4 setae and 1 solenidion proximal to proximal duplex setae S. filifolius Meyer, 1974
– Female: tarsus I with 2 setae and 1 solenidion proximal to proximal duplex setae 13
13 Female: All hysterosomal setae longer than longitudinal interval between their bases. Male: tarsus I with 2 setae and 2 solenidia proximal to proximal duplex setae, upturned part of aedeagus making almost right angle with shaft S. sacharum Flechtmann & Baker, 1975
– Female: most of hysterosomal setae approximately as long as the longitudinal interval between their bases. Male: tarsus I with 3 setae and 2 solenidia proximal to proximal duplex setae, upturned part of aedeagus making acute angle with shaft S. youngi Tseng, 1975
14 Female: hysterosomal setae especially c1, d1, e1 barely reaching the bases of next setae in line. Male: tarsus I with 1 solenidion and 1 tactile seta proximal to proximal duplex setae S. krungthepensis Auger & Naing, 2014 (as in Naing et al. 2014)
– Female: hysterosomal setae especially c1, d1, e1 longer than distance to bases of next setae in line. Male: tarsus I with 2 or 3 solenidia and 1 tactile seta proximal to proximal duplex setae 15
15 Male: tibia I with 7 tactile setae and 3 solenidia, tibia II with 5 tactile setae and 1 solenidion, tarsus I with 1 tactile seta and 3 solenidia proximal to proximal duplex setae S. arcuatus Meyer, 1974
– Male: tibia I with 7 tactile setae and 4 solenidia, tibia II with 5 tactile setae only without solenidion, tarsus I with 1 tactile seta and 2 solenidia proximal to proximal duplex setae S. rhynosperus Flechtmann & Baker, 1970
16 Female: Dorsal hysterosoma medially with transverse striations except area between setae c1, d1, and e1 forming V-shaped or longitudinal pattern 17
– Female: Dorsal hysterosoma medially with transverse striations entirely 19
17 Female: Stylophore anteriorly emarginate, with notch 18
– Female: Stylophore anteriorly without notch S. sacrales Baker & Pritchard, 1960
18 Female: Tarsus I with 2 solenidia and two tactile setae proximal to proximal duplex setae; striations in between setae c1 to d1 longitudinal S. baltazarae Rimando, 1962
– Female: Tarsus I with 1 solenidion and 2 tactile setae proximal to proximal duplex setae; striations in between setae c1 to d1 transverse S. spiculus Baker & Pritchard, 1960
19 Peritremes looped (making a loop) distally S. nugax Pritchard & Baker, 1955
– Peritremes hooked or making L-shape distally 20
20 Idiosoma elongate, ratio of body length (not including rostrum): width > 2 23
– Idiosoma oval/ orbicular, ratio of body length (not including rostrum): width < 2 21
21 Female: Dorsal striations smooth without lobes. Male: aedeagus upturned part making right angle with the shaft S. sagatus Davis, 1969
– Female: Dorsal striations with lobes. Male: aedeagus upturned part making obtuse angle with the shaft 22
22 Femur I with 7 setae, stylophore notched anteriorly, striations in pregenital area making strongly arched S. denmarki Baker & Tuttle, 1994
– Femur I with 9 setae, stylophore rounded anteriorly, striations in pregenital area transverse S. pseudolycurus Ochoa, Gray & von Lindeman, 1990
23 Tarsus I with 4 tactile setae proximal to proximal duplex S. fluvialis McGregor, 1928
– Tarsus I with 3 tactile setae proximal to proximal duplex 24
24 Striations in pregenital area almost transverse slightly curved S. freitezi Ochoa, Gray & von Lindeman, 1990
– Striations in pregenital area longitudinal irregular S. oryzae Rossi de Simons, 1966

III- taquarae species subgroup (05 species)

(Female: tibia II with 5 setae, tibia I with 6 tactile setae excluding solenidion)

Key to species of taquarae species subgroup

1 Dorsal setae very short, far behind the bases of next setae in line 2
– Dorsal setae long, at least reaching or crossing the bases of next setae in line 3
2 Female: peritremes hooked distally, dorsal striations without lobes, striations on pregenital area transverse; femur IV with 3 setae. Male: aedeagus upturned part sigmoid without anterior projection S. tegophallos Flechtmann & Peralta-Alba, 2012
– Female: peritremes simple, without hook distally, dorsal striations with lobes, striations on pregenital area longitudinal; femur IV with 2 setae. Male: aedeagus upturned part not sigmoid with anterior projection S. umtaliensis Meyer, 1974
3 Female: tarsus I with 4 tactile setae and a solenidion proximal to proximal duplex setae. Male: aedeagus not sigmoid, shaft almost straight, narrowing toward distal end S. triquetrus Meyer, 1987
– Female: tarsus I with 2 or 3 tactile setae and a solenidion proximal to proximal duplex setae. Male: aedeagus almost sigmoid, upturned 4
4 Dorsal striations with lobes, tarsus I with 2 tactile setae and a solenidion proximal to proximal duplex setae S. taquarae Paschoal, 1971
– Dorsal striations without lobes, tarsus I with 3 tactile setae proximal to proximal duplex setae S. papillatus Flechtmann, 1995

2. asparagi species group (20 species)

(Female: tibia II with 7 setae)

Key to species (20) of asparagi species group

1 Tibia I with 12 setae including solenidia 2
– Tibia I with 8–10 setae and 1 solenidion 3
2 Female: femur IV with 4 setae, genua III and IV with 3 setae. Male: aedeagus downturned, with only posterior projection S. emeiensis Wang, 1983
– Female: femur IV with 4 setae, genua III and IV with 3 setae. Male: aedeagus upturned, with anterior and posterior projections S. kreiteri Flechtmann, 1999 (as in Flechtmann et al. 1999)
3 Tibia I with 7 or 8 setae and a solenidion 4
– Tibia I with 9 setae and a solenidion 7
4 Tibia I with 7 tactile setae and a solenidion S. lanyuensis Tseng, 1975
– Tibia I with 8 tactile setae and a solenidion 5
5 Dorsal hysterosoma medially with transverse striations entirely, dorsal setae, especially c1, d1, e1, longer than interval between their bases S. miyatahus (Meyer, 1974)
– Dorsal hysterosoma medially between setae d1 and e1 with longitudinal irregular striations, dorsal setae shorter than interval between their bases 6
6 Female: Dorsal setae serrated, tarsus I with 5 tactile setae and 1 solenidion proximal to proximal duplex setae. Male: aedeagus upturned with small anterior projection and long posterior projection S. brevisetosus Ehara, 1989
– Female: Dorsal setae nude, tarsus I with four tactile setae and a solenidion proximal to proximal duplex setae. Male: aedeagus almost straight without anterior projection S. rhodanus Baker & Pritchard, 1960
7 Femur I with 10 setae 8
– Femur I with 8 or 9 setae 12
8 Tibia IV with 6 setae 9
– Tibia IV with 7 setae 10
9 Peritremes hooked distally S. lushanensis Wang, 1994
– Peritremes simple/ straight distally S. zhangi Wang & Cui, 1992
10 Femur II with 6 setae, femur IV with 4 setae S. kaspari Manson, 1967
– Femur II with 7 setae, femur IV with 2 or 3 setae 11
11 Femur III with 4 setae, femur IV with 3 setae S. tuminicus Ma & Yuan, 1982
– Femur III with 3 setae, femur IV with 2 setae S. halimodendri Wainstein, 1958
12 Femur I with 8 setae S. sayedi Attiah, 1967
– Femur I with 9 setae 13
13 Tibia III with 5 setae 14
– Tibia III with 6 setae 15
14 Female: Tibia IV with 5 setae, femur IV with 4 setae. Male: aedeagus distal part downturned, with small posterior projection S. asparagi (Oudemans, 1928)
– Female: Tibia IV with 6 setae, femur IV with 3 setae. Male: aedeagus distal part upturned, with large posterior projection S. tephrosiae Gutierrez, 1968
15 Genu IV with 3 setae S. lespedezae Beglyarov & Mitrofanov, 1973
– Genu IV with 4 setae 16
16 Striations on dorsal hysterosoma medially between setae e1 forming V-shaped or irregular longitudinal patterns 17
– Dorsal hysterosoma medially with entirely transverse striations 18
17 Female. Dorsal setae c1, d1, and e1 just reaching the bases of next consecutive setae, peritremes slightly hooked distally. Male. Aedeagus with very minute anterior projection, aedeagal knob making acute angle with shaft S. colocasiae Ehara, 1988 (as in Ehara and Tho 1988)
– Female. Dorsal setae c1, d1 and e1 crossing the bases of next consecutive setae, peritremes almost straight distally. Male. Aedeagus with very prominent anterior projection, aedeagal knob making obtuse angle with shaft S. malodhensis Sadana et al., 1985
18 Setae c1 and d1 reaching maximum up to 2/3 distance to setae next in line S. protectus Meyer, 1965
– Setae c1 and d1 as long as or crossing the bases of setae next in line 19
19 Male: Eupathidium on palp tarsus absent, aedeagus knob of upturned part parallel with the shaft S. malayanus Ehara, 1988 (as in Ehara and Tho 1988)
– Male: Eupathidium on palp tarsus present, aedeagus knob of upturned part making obtuse angle with the shaft S. bhandhufalcki Ehara & Wongsiri, 1975

3. bambusae species group (22 species)

(Female: Tibia II with 8 setae)

Key to species of bambusae species group

1 Tibia I with 10 or 11 setae including solenidion 2
– Tibia I with 9 setae including solenidion S. indicus Gupta & Gupta, 1994
2 Peritremes distally hooked, U/L-shaped 3
– Preitremes almost straight, slightly expanded distally, not hooked/ U/L-shaped distally 11
3 Genua III and IV each with 3 setae 4
– Genua III and IV each with 4 setae 7
4 Female: Dorsal setae comparatively short, far behind the bases of next setae in line. Male: aedeagus distal part upturned with anterior projection S. gilvus Ehara & Ohashi, 2005
– Female: Dorsal setae long, crossing the bases of next setae in line. Male: aedeagus distal part straight undulating or downturned without anterior projection 5
5 Female: setae c1 just crossing setae d1, far behind the bases of setae e1, tarsus I with 2 or 3 tactile setae and a solenidion proximal to proximal duplex setae S. minutus Wang, 1985 (as mentioned in Wang et al. 1985)
– Female: setae c1 reaching the bases of setae e1, tarsus I with 4 or 5 tactile setae and 1 solenidion proximal to proximal duplex setae 6
6 Female: pregenital area with longitudinal striations, tarsus I with 4 tactile setae and 1 solenidion proximal to proximal duplex setae. Male: aedeagus distal part almost straight, undulating, slightly turning up S. gahniae Davis, 1969
– Female: pregenital area with transverse striations, tarsus I with 5 tactile setae and 1 solenidion proximal to proximal duplex setae. Male: aedeagus distal part almost straight, downturned S. cercidiphylli Ehara, 1973
7 Femur IV with 3 setae 8
– Femur IV with 4 setae 9
8 Femur II with 6 setae, peritremes V-shaped distally S. imperatae Wang, 1983
– Femur II with 7 setae, peritremes L-shaped distally, slightly hooked S. textor Wainstein, 1954
9 Female: dorsal setae especially c1, d1, e1 almost reaching to the bases of setae next in line. Male: aedeagus distal part downturned and sigmoid S. fauveli Gutierrez, 1978
– Female: dorsal setae especially c1, d1, e1 well crossing to the bases of setae next in line. Male: aedeagus distal part not sigmoid, almost straight/ undulating 10
10 Male: aedeagus distal part very long needle-like undulating S. alni Beglyarov & Mitrofanov, 1973
– Male: aedeagus distal part downturned slightly gradually narrowing toward distal end, not needle-like S. zhongdianensis Wang & Cui, 1992
11 Tibia I with 10 tactile setae and 1 solenidion 12
– Tibia I with 9 tactile setae and 1 solenidion 13
12 Female: femur I with 9 setae, femur II with 6 setae genu III with 3 setae. Male: aedeagus distal part straight undulating S. garmani Pritchard & Baker, 1955
– Female: femur I with 10 setae, femur II with 7 setae, genu III with 4 setae. Male: aedeagus distal part slightly upturned, dorsally making slight knob, then bent down distally S. levinensis Manson, 1967
13 Male: aedeagus distal part upturned 14
– Male: aedeagus distal part, straight or downturned 18
14 Male: aedeagus distal upturned part with prominent anterior projection and long posterior projection 15
– Male: aedeagus distal upturned part without anterior projection 16
15 Eupathidium on male palp tarsus long, almost as long as eupathidia aedeagus posterior projection of upturned part is 4× longer than width of aedeagus neck and making prominent angle with neck S. beckeri Wainstein, 1958
– Eupathidium on male palp tarsus minute, eupathidia 3× longer than, aedeagus posterior projection of upturned part is 2–3× longer than width of neck and not making angle with neck S. brachypodii Livshits & Mitrofanov, 1968
16 Male: aedeagus distal part turn dorso-caudally, almost sigmoid in shape S. ibericus Reck, 1947
– Male: aedeagus distal part turn dorsally, not sigmoid in shape 17
17 Female: pregenital area with transverse striations, tarsus I with 4 tactile setae and a solenidion, tarsus II with 4 setae and a solenidion proximal to proximal duplex setae. Male: aedeagus upturned part greatly narrowing, needle-like S. floresi Rimando, 1962
– Female: pregenital area with longitudinal striations, tarsus I with 5 tactile setae and 1 solenidion, tarsus II with 3 setae and 1 solenidion proximal to proximal duplex setae. Male: aedeagus upturned part blunt distally, not narrowing S. bambusae Reck, 1941
18 Male: aedeagus distal part straight, undulating S. jachontovi Reck, 1953
– Male: aedeagus distal part down turned 19
19. Female: femur IV with 3 setae. Male: aedeagus distal part slightly downturned without anterior projection 20
– Female: femur IV with 4 setae. Male: aedeagus distal downturned part with distal knob, neck, and anterior projection 21
20 Female: tarsus I with 19 and tarsus II with 16 setae. Male: eupathidium su on palp tarsus long, longer than eupathidia S. smirnovi Wainstein, 1954
– Female: tarsus I with 18 and tarsus II with 15 setae. Male: eupathidium su on palp tarsus almost half in length than eupathidia S. iraniensis Mahdavi & Asadi, 2015
21 Male: aedeagus anterior and posterior projections almost equal, knob forming obtuse angle with the shaft S. chiangmaiensis Ehara & Wongsiri, 1975
– Male: aedeagus posterior projections much longer (2–3×) than anterior projection, knob forming acute angle with the shaft S. euphorbiae Livshitz & Mitrofanov, 1968

4. vermiculatus species group (4 species)

(Female: Tibia II with 4 setae)

Key to species of vermiculatus species group

1 Tibia I with 7 setae and a solenidion, tibiae III and IV each with 4 setae S. vermiculatus Ehara & Wongsiri, 1975
– Tibia I with 6 setae and a solenidion, tibiae III and IV each with 3 setae 2
2 Genua III and IV each with 3 setae S. approximatus Ehara, 1988
– Genua III and IV each with 2 setae 3
3 Dorsocentral area between setae c1, d1, e1, and f1 smooth, without striations S. laevidorsatus Ehara, 1988
– Dorsocentral area between setae c1, d1, e1, and f1 with transverse striations S. saitoi Ehara, 1988

5. spireafolia species group (10 species)

(Female: Tibia II with 6 setae)

Key to species of spireafolia species group

1 Tibia I with 6 tactile setae and a solenidion S. prosopis Tuttle, Baker & Abbatiello, 1976
– Tibia I with 7 tactile setae and a solenidion 2
– Tibia I with 8 or 9 tactile setae and a solenidion 4
2 Dorsal setae very long, c1 crossing the bases of d1 reaching up to the bases of e1, setae d1 reaching up to the bases of f1 S. parasemus Pritchard & Baker, 1955
– Dorsal setae, short, setae c1 and d1 and e1 almost reaching up the bases of setae next in line or just crossing the bases of setae next in line 3
3 Female: Tarsus I with 2 setae and 1 solenidion proximal to proximal duplex setae, tarsus II with 1 tactile seta and 1 solenidion proximal to duplex setae. Male aedeagus upturned distal part shorter (less than half) the length of shaft S. recki Ehara, 1957
– Female: Tarsus I with 5 tactile setae proximal to duplex setae, Tarsus II with 4 tactile setae and 1 solenidion proximal to duplex setae. Male aedeagus upturned distal part as long as the length of shaft S. undulatus (Beer & Lang, 1958)
4 Tibia I with 9 tactile setae and a solenidion S. ugarovi Wainstein, 1960
– Tibia I with 8 tactile setae and a solenidion 5
5 Peritremes hooked distally. Male aedeagus upturned part with neck and anterior projection S. shii Ehara, 1965
– Peritremes straight distally. Male aedeagus without anterior projection 6
6 Dorsal hysterosomal setae (most of them) awl-shaped, acutely tapering from the widened proximal (basal) portion S. spireafolia Garman, 1940
– Dorsal hysterosomal setae setose 7
7 Tibiae III and IV each with 6 setae S. dalbergiae Meyer, 1974
– Tibiae III and IV each with 5 setae 8
8 Femur II, III, and IV with 8, 4, and 2 setae, respectively. Male aedeagus upturned part as long as the length of shaft S. elongatus Wang & Cui, 1991
– Femur II, III, and IV with 6, 3, and 3 setae, respectively. Male aedeagus upturned part very minute as compared to the length of shaft S. avetjanae Bagdasarian, 1954

Acknowledgments

The authors would like to extend their sincere appreciation to Yutaka Saito (Research Faculty of Agriculture, Hokkaido University, Sapporo, Japan) for sharing the published descriptions, redescriptions, and illustrations of all the species of the genus Stigmaeopsis.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The authors would like to extend their sincere appreciation to the Researchers Supporting Project number [RSPD2024R807], King Saud University, Riyadh, Saudi Arabia for funding this research.

Author contributions

All authors have contributed equally.

Author ORCIDs

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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