International Journal of Acarology

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Life history, nesting behaviour, and social

behaviour of Stigmaeopsis spider mites (Acari:
Tetranychidae)

Muhammad Saad Waqas, Tian-Ci Yi, Muhammad Kamran, Liang-Yu Sun,

Rong Xiao & Dao-Chao Jin

To cite this article: Muhammad Saad Waqas, Tian-Ci Yi, Muhammad Kamran, Liang-Yu Sun,
Rong Xiao & Dao-Chao Jin (2021): Life history, nesting behaviour, and social behaviour of
Stigmaeopsis spider mites (Acari: Tetranychidae), International Journal of Acarology, DOI:
10.1080/01647954.2021.1968492

To link to this article: https://doi.org/10.1080/01647954.2021.1968492

Published online: 26 Aug 2021.

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INTERNATIONAL JOURNAL OF ACAROLOGY
https://doi.org/10.1080/01647954.2021.1968492

Life history, nesting behaviour, and social behaviour of Stigmaeopsis spider mites
(Acari: Tetranychidae)
Muhammad Saad Waqasa, Tian-Ci Yia, Muhammad Kamranb,c, Liang-Yu Suna, Rong Xiaoa and Dao-Chao Jina
a
Department of Plant Protection, Institute of Entomology, Guizhou University, Guiyang, P. R. China; Provincial Key Laboratory for Agricultural Pest
Management of Mountainous Regions, and Scientific Observing and Experimental Station of Crop Pest in Guiyang, Ministry of Agriculture and Rural
Affairs, Guiyang, P. R. China; bDepartment of Plant Protection, College of Food and Agriculture Sciences, King Saud University, Pakistan; cDepartment
of Plant Protection, Ghazi University, Dera Ghazi Khan, Punjab, Pakistan

ABSTRACT ARTICLE HISTORY

Stigmaeopsis mites are voracious feeders of bamboo plants, Pleioblastus spp., Miscanthus spp., and Sasa Received 28 October 2020
spp. Fifteen species of the genus Stigmaeopsis have been reported worldwide. Their life history is Accepted 2 August 2021
affected by temperature and they can have many generations overlap in their nests under favourable Published online 27 August
2021
conditions. Stigmaeopsis species overwinter as only adult stages in temperate zones. Males kill con­
specific males to mate with unmated females. Most Stigmaeopsis mites defaecate at pre-determined KEYWORDS
sites, which differ among species. Stigmaeopsis mites determine defaecation sites through volatile Stigmaeopsis mites; life
chemical cues or tactile stimuli. Herein, we discuss advances in the understanding of aspects of history; nest-weaving; nest
Stigmaeopsis mite biology and behaviour, such as life history, life types, reproduction, overwintering, sizing; waste management;
nest building and sizing, defaecation, mating, and defence against predators. We conclude that there is defence strategy;
a paucity in the literature concerning the biological traits of Stigmaeopsis. Furthermore, we suggest that overwintering
additional research be performed on the effects of temperature, geographic distribution, and biological
and chemical control on the species of this genus.

Introduction
Stigmaeopsis (Acari: Tetranychidae) is an important genus of social
and nest-weaving mites (Saito 1995a). To date, Stigmaeopsis contains
15 species (Saito et al. 2019), which live in groups and build tunnel-
like woven nests that consist of two entrances and a roof on the leaf
abaxial surface (Saito 1997; Saito et al. 2004). Stigmaeopsis celarius was
first collected from bamboo plants (Pleioblastus simonii) about
100 years ago (Banks 1917). Thirty-three years later, this species was
relocated to the genus Schizotetranychus by McGregor (1950). Saito
et al. (2004) reinstated the genus Stigmaeopsis Banks, described two
new species S. saharai and S. takahashii and moved more species that
were previously assigned to the Schizotetranychus celarius-species
group, to the genus Stigmaeopsis. These species include
S. tenuinidus Zhang et Zhang, S. longus Saito, S. nanjingensis Ma et
Yuan, S. miscanthi Saito et Mori, and S. celarius. Biology and morphol­
ogy of S. nanjingensis is closely related to those of S. celarius (Zhang
et al. 1999). S. malkovskii and S. meghalayensis were first reported by
Flechtmann (2012) in Brazil. Later in 2016 and 2018, S. tegmentalis
Saito et Lin, S. temporalis Saito et Ito (Saito et al. 2016a), and
S. continentalis Saito et Lin (Saito et al. 2018) were added to the
Stigmaeopsis genus. In 2019, two new Stigmaeopsis species,
S. inthanonsis Saito, Kongchuensin & Sahara and S. formosa Saito,
Chao & Sato, were described and collected from Miscanthus grasses
in Thailand and Taiwan, respectively (Saito et al. 2019).
Stigmaeopsis is globally widespread, including Australasia, Asia
(China, Japan, Saudi Arabia, and Thailand) (Saito et al. 2004, 2019),
Europe (e.g. Belgium, Britain, France, Italy, and the Netherlands)
(Pellizzari and Duso 2009; Alford 2012), and the United States of
America (Saito et al. 2004).
A behaviour shared by all Stigmaeopsis mites is that they live on
the abaxial side of the host plant. Stigmaeopsis damage the host
plant by sucking sap from the leaves (Yu and Shi 1991). The
infested parts generally develop yellowish-white or completely
whitish blotches on the upper side of leaves (Zhang et al. 1998).
Furthermore, the infested leaves become bleached, and flecked
with silvery-white spots where the contents of plant cells have
been removed (Alford 2012). S. nanjingensis reduces sugar and
chlorophyll levels in leaves, leading to a reduction in leaf area,
and an increase in culms thickness and length (Zhang and Zhang
2000). Consequently, photosynthetic activities are impaired, and
leaves may shed prematurely (Alford 2012).
Stigmaeopsis mites are significant threats to bamboo plants and
these are excellent species for the studies of evolutionary ecology
and genetic perspectives, because of their small size, less move­
ment, ease of rearing, short life history, social and sub-social life
types, and rapid development. The aim of this review is to provide
complete information and recent advances on the Stigmaeopsis
life history, life type, waste management, nesting behaviour, mat­
ing behaviour, diapause, male-to-male aggression, social beha­
viour, molecular phylogeny and defence against predators.
Phylogenetic relationship
Identification of Stigmaeopsis is very difficult morphologically, only
length of dorsal setae is a diagnostic characteristics to identify fresh
specimens. In addition, tarsus-I with 1 solenidion proximal to duplex
setae (Saito et al. 2016) and mediodorsal striae (Saito 2018) are also
important traits in identification. Sequencing of the cytochrome oxi­
dase subunit I (COI) gene of mitochondrial DNA and ribosomal RNA
genes (18S and 28S) are rapid identification methods (Sakamoto et al.
2017). Molecular phylogenetic relationship of Stigmaeopsis proved
that the genus is monophyletic. Schizotetranychus species formed
a different clade from Stigmaeopsis, although they show a very similar
life type (WN-c, Saito 1983) and morphology (Sakagami et al. 2009).
Phylogenetically, S. miscanthi and S. longus are derived from an
ancestor of S. takahashii and S. saharai and they can easily interbreed

CONTACT Dao-Chao Jin daochaojin@126.com; Tian-Ci Yi yitianci@msn.com Institute of Entomology, Guizhou University, Guiyang, P. R. China; Provincial
Key Laboratory for Agricultural Pest Management of Mountainous Regions, and Scientific Observing and Experimental Station of Crop Pest in Guiyang, Ministry of
Agriculture and Rural Affairs, Guiyang 550025, P. R. China
© 2021 Informa UK Limited, trading as Taylor & Francis Group

Published online 26 Aug 2021

2 M. S. WAQAS ET AL.

to form F1 (Mori and Saito 2013). Alignment of parasodium channel Adult males remain in their nests and mate with sisters or
gene and mtCOI, HG populations and LW populations formed a single disperse to other nests (Saito et al. 2016b). Reproduction of
clade and ML populations split into four clades: two populations from Stigmaeopsis spp. consists of the periods: preoviposition, oviposi­
Fujian and four populations from Taiwan of China, nine populations tion, and postoviposition. Preoviposition of S. nanjingensis and
from the Southern Ryukyus, and two populations from mountainous S. longus, and lasts 5 to 6 and 3 days, respectively, at 25°C (Saito
area in Taiwan (Sato et al. 2019). and Ueno 1979; Zhang et al. 1999; Saito et al. 2016b). Oviposition
starts after mating and each female lays one egg per day or some­
times 2 to 3 eggs on alternate days in S. nanjingensis (Yu and Shi
Life history 1991; Zhang et al. 1999; Alford 2012).
Eggs are spherical with an average length of 0.1 mm and adults
All Stigmaeopsis mites are phytophagous with five stages,
are whitish to translucent with an average length of 0.4–0.6 mm
namely, egg, larva, protonymph, deutonymph, and adult (Saito
(Alford 2012). At 25°C, each female of S. celarius produces an
et al. 2016b). However, the deutonymphal stage is absent in
average of 85 ± 4 eggs (Saito and Ueno 1979). Stigmaeopsis
S. nanjingensis males (Zhang et al. 1999). The life history of
nanjingensis produces an average of 30–35 eggs per female in
Stigmaeopsis mites varies among species (Saito 1979).
the Fujian province (Zhang et al. 1999), while this same species
Stigmaeopsis longus and S. celarius have a short developmental
produces an average of 17 eggs in the Zhejiang province, China
period (Saito and Ueno 1979; Saito 1986a, 2018). The develop­
(Yu and Shi 1991).
mental period from egg to adult of Stigmaeopsis mites at 25°C
takes approximately from 14 to 22 days. Longest generation
time reported by Saito (2018) for S. tenuinidusis 22.1 days; fol­ Overwintering
lowed by S. nanjingensis, 18.8 days (Zhang 2002); S. sabelisi,
Stigmaeopsis mites overwinter in the nest in temperate zones. In all
17.1 days; S. miscanthi, 14.6 days (Saito et al. 2013); S. Saharai,
spider mites, the term “overwinter” is used instead of “diapause”
14.5 days (Mori 2000); S. longus, 14.0 days (Saito and Ueno 1979);
because diapause is a physiological state and must be used only
S. celarius, 14.0 days; and S. takahashii, 13.9 days (Mori 2000). The
after it has been proved experimentally. The following overwinter­
average developmental period of most Stigmaeopsis mites is
ing stages have been observed in spider mites: egg (Lees 1953),
from 14 to 16 days at 25°C and 50–80% relative humidity
adult, reproductive diapause, and both egg and adult (Gotoh 1986).
(Saito et al. 2016b). Developmental time of S. longus is reduced
According to Ito and Yamanishi (2019), Panonychus and
significantly as the temperature increases from 15°C to 28°
Oligonychus species overwinter in egg stage, Stigmaeopsis,
C (Gotoh and Shida 2007). Development period of S. longus is
Tetranychus and Eotetranychus species overwinter in female adult
the longest at 15°C and shortest at 28°C (Gotoh and Shida 2007).
stage (Yanagida 1999) and Sasanychus akitanus (Ehara) overwinter
Egg hatchability of S. nanjingensis is 81–100% at 22–30°C, 33% at
in both eggs and adult females stage (Gotoh 1986aa). The over­
35°C, and the developmental period of immature stages
wintering stage of each genus is associated with host phenology
decreases as temperature increases from 15 to 29°C. Total devel­
(Gutierrez and Helle 1985). Stigmaeopsis akitanus (Ehara) overwinter
opmental period from egg to adult is completed within 2 weeks
on evergreen Sasa bamboos (Gotoh 1986aa, b). In S. miscanthi,
at 29°C (Zhang et al. 1999). Stigmaeopsis miscanthi immatures
S. saharai, and S. takahashii, only female adults are capable of
could survive on Sasa bamboo for a long period and S. longus
overwintering (Saito et al. 2005). Some females overwintered in
immatures could not survive on Miscanthus grass (Sakagami
the fertilized (mating with male) state and some females overwin­
et al. 2009).
tered in unfertilized (without mating) state. For example, 8.7–21.1%
of S. longus and 36% of S. miscanthi females overwintered in the
unfertilized state and remaining females overwintered in fertilized
Life type diversity state (Saito 1987).
Different web-weaving patterns were first described by Gutierrez
et al. (1970). Later, the concept of life type was defined as Overwintering in females
“different strategies by which different mite species develop
Overwintering females find nests solitarily during spring, and
silken threads on the host plant leaves” (Saito 1983; Saito and
unfertilized females produce haploid males, and in cold areas
Mori 2005). Life types of Stigmaeopsis mites are characterized by
where a small population of males occurs during the spring
different nest architecture (Saito et al. 2016b) and other charac­
season, mother–son inbreeding is observed (Saito et al. 2000).
teristics, such as feeding preferences, quiescent stage, egg cover­
Under laboratory conditions, low temperature (18°C) and
ing, nest construction, as well as preferred sites for faeces
short day length induce reproductive diapause in female adults
deposition, oviposition, and walking sites (Yu and Shi 1991;
of S. longus, S. miscanthi LW and T. kanzawai (Saito et al. 2005).
Zhang et al. 1999; Sato and Saito 2006; Kanazawa et al. 2011;
Under field conditions, S. longus egg density started to diminish
Sato et al. 2013). Based on these characteristics, the life type of
during the last week of September and declined during October
spider mites can be broadly divided into three small groups:
and after November, adult females congregate with a few males
complicated web (CW), little web (LW), and woven nest (WN);
and nymphs in the nests to diapause. Overwintering females lay
these three groups are divided into other small subgroups based
eggs after the completion and breakdown of diapause condi­
on species association with host plants (Table 1) (Saito 1979,
tion during the early spring by initiating a small colony in
1983, 1985). All Stigmaeopsis species belong to the WN life
a densely woven silken nest on the abaxial side of young leaves
type. Differences in body structure, type locality, host plant, life
(Alford 2012). The diapausing females lay orange eggs, and the
type, and morphology among Stigmaeopsis mite species, except
immature mites and adult males leave the nests. Egg hatching
S. malkovskii and S. meghalayensis, are illustrated in Table 1.
occurs in March and early development starts in April.
Overwintering S. nanjingensis females produce eggs in late
February, which hatch in April (Zhang et al. 2001).
Reproduction
Sex ratio varies among Stigmaeopsis species and all species exhibit
Male performance
female-biased sex ratio (Saito and Ueno 1979; Mori 2000; Zhang
2002; Sato et al. 2013). The male sex ratio of S. celarius is 0.28; There are three forms of male–male aggressiveness: low aggressive­
S. saharai, 0.21 (Mori 2000); S. longus, 0.14; S. takahashii, 0.16 (Saito ness, mild aggressiveness and high aggressiveness (Sato et al. 2019).
and Takahashi 1982); and S. miscanthi 0.15 (Sato et al. 2000). Populations of low, mild and high aggressiveness are discriminated
Table 1. Body structure, type locality, host plant, life type and diagnostic characteristics of Stigmaeopsis species.
Specie name and year of
discover Body structure Host plant Life type Diagnostic characters Reference
Stigmaeopsis celarius Banks 1917 Body flattened. Strawy to greenish yellow Pseudosasa japonica WN-c This specie is distinguished from other genera (Banks 1917)
with small blackish green spots. of Tetranychidae by the chaetotaxy of the distal segment
of palp.
Stigmaeopsis longus Saito (1990) Body flattened. Strawy to greenish yellow Sasa senanensi, WN-c S. longus is differentiated from other species by the second propodosomal (Saito et al. 2004)
with small blackish green spots. S. kurilensis and Sasa setae, second and third dorsocentral hysterosomal setae (C2 and C3)
species
Stigmaeopsis nanjingensis　 * Phyllostachys pubescens, WN-c This species is distinguished from other species by the arrangement of dorsocentral (Saito et al. 2004)
Ma & Yuan (1980) P. praecox and P. vivax setae
Stigmaeopsis tenuinidus　Zhang * Phyllostachys pubescens WN-p * (Saito et al. 2004;
and Zhang 2000 Saito et al.
2016a)
Stigmaeopsis saharai Sasa senanensis, sasa WN-c S. saharai (Saito et al. 2004)
Saito & Mori (2004) kurilensis, and other distinguished from other Stigmaeopsis species by the lengths of P2 and
Sasa C2 setae.
spp.
Stigmaeopsis takahashii Body flattened, strawy to greenish yellow Sasa senanensis, sasa WN-c S. takahashii i (Saito et al. 2004)
Saito & Mori (2004) with small blackish green spots. kurilensis, and other distinguished from other species by the length of second dorsocentral hysterosomal setae (C2).
Sasa
spp.
Stigmaeopsis miscanthi　 Body flattened, strawy to greenish yellow Miscanthus sinensis, WN-c S. miscanthi　 (Saito et al. 2004)
Saito & Mori (1990) with small blackish green spots. Miscanthus sp differentiated from other species by the proportions of male leg I and the reproductive isolation
behaviour
Stigmaeopsis temporalis Body flattened and wide, straw to greenish Pleioblastus argentostria- WN-p S. temporalis (Saito et al. 2016a)
Saito & Ito (2016) yellow with tus distinguished from other species by the numbers of setae on leg segments
small blackish green spots
Stigmaeopsis tegmentalis Body cylindrical and slender, straw yellow Phyllostach-ys makinoi WN-h S. tegmentalis (Saito et al. 2016a)
Saito & Lin (2016) with small blackish spots distinguished other species by dorsal setae c1 and c2.
Stigmaeopsis sabelisi Body flattened and wide, straw to greenish Miscanthus sinensis and WN-c S. sabelisi (Saito et al. 2018)
Saito & Sato (2018) yellow with small blackish green Miscanthus species distinguished from other species by
spots. the length of dorsal propodosomal setae
Stigmaeopsis continentalis Body flattened and wide, straw to greenish Miscanthus WN-c S. continentalis (Saito et al. 2018)
Saito & Lin (2018) yellow with small blackish green Sinensis and differentiated from other species by the distance between bases of dorsolateral hysterosomal setae
spots Miscanthus species.
Stigmaeopsis inthanonsis Saito, Body flattened and wide, straw to greenish Miscanthus species WN-c S. inthanonsis distinguished from other species by the length of dorsal propodosomal setae sc1 and (Saito et al. 2019)
Kongchuensin & Sahara yellow dorsocentral hysterosomal setae c1.
(2019)
Stigmaeopsis formosa Saito, Body flattened and wide, straw to greenish Miscanthus species WN-c S. formosa distinguished from other species by the distance between the bases of dorsolateral (Saito et al. 2019)
Chao & Sato (2019) yellow with small blackish green spots. hysterosomal setae c2, dorsocentral hysterosomal setae d1 and f1, and the length of c1 and c3 setae.
*Literature available in Chinese with English abstract.
S. malkovskii and S. meghalayensis are not included due to lack of information on body structure, host plant, life type and diagnostic characters.
INTERNATIONAL JOURNAL OF ACAROLOGY
3
4 M. S. WAQAS ET AL.
against as LW, ML and HG forms, because of their behavioural,
ecological, and morphological differences (Sakagami et al. 2009).
LW populations and HG populations both clustered into single
clade (Saito 2018) and ML populations split into four clades (Sato
et al. 2019). LW male overwintering is very difficult in cooler region
(northern or high altitude region of Japan), and nests only contain
deep diapause fertilized females, and reproduce in spring.
Consequently, nest mates in the later seasons consist of parents,
offspring and grand offspring. In contrast, HG males generally can
survive and overwinter successfully in warmer regions (low altitude
southern region), hibernated males from different nests visit the nest
and mates with females and females reproduce during spring (Sato
et al. 2013). LW S. miscanthi males have deeper diapause than that of
HG males (Saito et al. 2002).
Survival rates of adult females, adult males, and deutonymphs
of S. Longus are very high, 99.5%, 84.2–98.7%, and 89.7–89.8%,
respectively, whereas the survival rate of the eggs and quiescent
stages are extremely low, 16–23% and 0–20%, respectively, after
rearing under diapause-inducing short-day and low temperature
conditions (Ito and Chae 2019). High temperature during the
summer could reduce the S. nanjingensis population, leading to
aestivation and a new population is established from late summer
to November (Zhang et al. 1999).
Nest building
Stigmaeopsis mites build nest-like structures as living quarters
(Saito 1985; Zhang et al. 1999; Chae et al. 2015). They are consid­
ered a very unique group of mites, as its members exhibit social
behaviour. Deutonymphs and adult females play an important role
in nest building (Zhang et al. 1999). Adult females clean the target
spot on the leaf surface during nest construction (Kanazawa et al.
2011). Through silk-weaving, these individuals produce silk
threads onto the clean leaf surface by pressing their mouthparts,
the silk production organ, on one side of the leaf depression (Saito
1977), and by walking straight to the other side of the leaf depres­
sion, threads are extruded, enabling silk production (Kanazawa
et al. 2011). This process is repeated many times until the nest
building is complete. This behaviour corresponds to a single nest-
building trial (Mori and Saito 2006), and the whole process is called
silk-weaving behaviour (Jeppson et al. 1975). Kanazawa et al.
(2011) observed that S. longus females spent up to 3 hours build­
ing a new nest.
S. longus females exhibit social behaviour during nest building.
Indeed, females cooperate with each other and built large nests on
Sasa leaves, and adult males and females defend their nests and
nest members from the attack of predators (Saito 1986a, 1986b).
Therefore, this species is described as a cooperative social mite.
Nest sizing
Nest sizing varies among Stigmaeopsis species. Stigmaeopsis longus,
S. nanjingensis, and S. miscanthi build large woven nests, while
S. celarius and S. takahashii build mid-sized nests on the abaxial side
of Pleioblastus leaves and Sasa leaves, respectively, and S. saharai builds
smaller nests on Sasa leaves. Nest size is a strategy against the invasion
of other small animals, including mites, for example, the small size of
nests of S. saharai prevents invasion of large-sized S. longus (Saito 1990;
Mori 2000; Mori and Saito 2004).
The nest size is related to the morphology of Stigmaeopsis species.
Indeed, Stigmaeopsis species with longer setae sc1 construct larger
nests, whereas those species with smaller setae sc1 construct smaller
nests, which supports the idea that the length of dorsal setae sc1
determines the nest size (Mori and Saito 2013). Nest area (maximum
length × maximum width) constructed by S. longus, S. celarius,
S. takahashii and S. saharai within 48 h are 12.6 mm2, 6.9 mm2,
5.9 mm2, and 4.2 mm2, respectively (Mori and Saito 2005) .
Social behaviour is another factor affecting nest size.
Stigmaeopsis longus and S. celarius extend their nests continuously
as colonies increase, which allows them to remain in the nest
throughout their life span. More than 100 juvenile and adult
individuals of S. longus were observed cohabiting large nests for
long periods (Mori and Saito 2004). In contrast, Zhang et al. (1999)
observed that each S. nanjingensis nest generally contained three
females, 14 eggs, and five larvae. Stigmaeopsis miscanthi and
S. saharai also reside in small colonies. Females of S. takahashii
and S. saharai leave their parental nest before oviposition and
construct new nests for egg deposition and offspring develop­
ment (Saito and Takahashi 1982). Therefore, these observations
show that nest size is proportional to the number of colony
members (Saito 1990a). Furthermore, the number of males is
lower than the number of females in all colonies of Stigmaeopsis
(Mori and Saito 2005).
Waste management
All Stigmaeopsis have a well-developed nest waste management
system. Defaecation at a pre-determined site of the nest is one of
the main waste management behaviours of Stigmaeopsis mites
(Saito 1986a). However, the spot where the “toilet” is established
is defined differently among species (Sato and Saito 2006).
Stigmaeopsis nanjingensis and S. miscanthi defaecate inside the
nest, while S. longus, S. takahashii, S. saharai, S. celarius,
S. tenuinidus, S. temporalis, and S. tegmentalis defaecate outside
their nests (Saito et al. 2016b). Nest waste management by chemi­
cal and tactile cues is a newly evolved behaviour (Sato and Saito
2006). Different Stigmaeopsis species use volatile chemical cues or
tactile stimuli to manage waste (Sato and Saito 2008). Stigmaeopsis
takahashii and S. saharai recognize excrements by tactile cues
(Sato and Saito 2006), since their eyes cannot form fast images
(McEnroe and Dronka 1969). Stigmaeopsis miscanthi and S. longus
manage waste in two ways: they use tactile cues to defaecate at
the nest entrance when there is no excrement deposit, and they
use volatile chemical cues to defaecate at a site where excrements
have been deposited previously (Sato et al. 2003). Using volatile
chemical cues to confirm that excrements have been deposited at
a determined site of the nest is an adaptive behaviour of
S. miscanthi and S. longus, whose nests are extended continuously.
If these mites defaecated at every nest entrance, they would have
to establish a new faeces deposit site whenever they extended
nests, and this would result in excessive faecal deposits in the nests
(Sato and Saito 2006). The large-size nest builder, S. longus, defae­
cates in a single site of the nest by detecting leaf chemical cues,
whereas middle-size nest builders use tactile cues to defaecate
outside of nest entrances (Sato and Saito 2008).
Saito et al. (2016b) reported five faeces deposition combinations.
Stigmaeopsis longus and S. nanjingensis deposit massive faecal pile sat
a single site near the outside of the nest entrance. Stigmaeopsis longus
defaecate inside the nest. Stigmaeopsis nanjingensis, S. takahashii,
S. saharai, and S. celarius deposit massive faecal piles at two deter­
mined sites outside the nest. Stigmaeopsis tenuinidus, S. tegmentalis,
and S. temporalis defaecate in multiple sites of the nest.
Mating behaviour
Phytophagous male mites may kill the conspecific male to mate with
immature females immediately after its moulting (Boudreaux 1963).
The aggressive mating behaviour of male spider mites Tetranychus
urticae was first reported by Potter et al. (1976) and by Saito
et al. (2002) in Stigmaeopsis species. Large groups of Stigmaeopsis
males live in the same nest for long periods (Saito 1990b). Moreover,
multiple males simultaneously protect one static female. Males of
S. miscanthi often kill the opponent male and sucks its body fluid
(Saito 1995a).

Male–male aggression behaviour is correlated with leg I length,
which is a measuring tool of defence ability in males (Saito 1995b;
Sato et al. 2013). The frequency at which males kill conspecific males
varies from region to region, being low in colder regions, mild in
subtropical regions, and high in warmer regions (Sato et al. 2013).
Male aggressive behaviour is also species-specific. Stigmaeopsis
miscanthi are sibling species in the Tetranychidae and both have
well-developed sociality (Sakagami 2002; Saito et al. 2004; Mori
and Saito 2005). Male aggression behaviour of S. longus,
S. miscanthi, and S. sabelisis is significantly different owing to
environmental conditions (Saito 1990b, 1995a; Sato et al. 2013,
2018). Generally, male–male aggressive behaviour is lethal and
stronger in S. miscanthi compared with S. sabelisi and S. longus
(Saito and Sahara 1999). Additionally, S. miscanthi HG males fre­
quently kill conspecific males, whereas their LW males rarely kill
conspecific males (Saito and Sahara 1999). Stigmaeopsis miscanthi
ML form, S. formosa, and S. continentalis have been associated with
male fighting (Sato et al. 2013, 2019).
Defence against predators
Stigmaeopsis mites have strong defence mechanisms against
immature stages of phytoseiid predatory mites (Saito 1986a, b;
Mori and Saito 2004; Yano et al. 2011). Adults of S. longus strongly
counterattack the predator Typhlodromus bambusae Ehara (Saito
1986b) and adult males frequently kill the larval and protonymphal
stages of this predator (Saito 1986a). LG and HG parental males
and females of S. miscanthi strongly counterattack the predator
T. Bambusae, and compared with LG, the HG individuals are more
aggressive to kill T. bambusae (Yano et al. 2011). Stigmaeopsis
nanjingensis males and females have strong but different counter­
attacking behaviour: females trap T. bambusae immature stages in
the nest silk web, which starves them to death, while males directly
kill immature individuals of T. bambusae during predator invasion
(Saito and Zhang 2017). Stigmaeopsis takahashii and S. saharai
build small and separate nests to protect colonies from predator
intrusion and attack. S. miscanthi and S. longus build large nests by
continuously extending their original ones to counterattack
intruding predators (Mori and Saito 2004, 2005).
Nest size plays an important role in anti-predatory intrusion.
There are two anti-predation strategies in Stigmaeopsis. First, pro­
tection by smaller nests; second, long-term defence by multiple
individuals in larger nests (Mori and Saito 2005). Stigmaeopsis
mites living in large continuous nests have stronger anti-
predation behaviour than those living in smaller nests (Saito
et al. 2016b). Living in small nests protects the eggs of
Stigmaeopsis from Phytoseius tenuiformis, Agistemus iburiensis,
and Agistemus summersi, whereas living in larger nests protect
their eggs from Amblyseius spp. (Mori and Saito 2004).
Conclusions and prospects
Life history of Stigmaeopsis differ among species. The most dis­
tinctive biological trait in the genus Stigmaeopsis is that deuto­
nymphs and females build their nests. Additionally, these mites
use volatile chemical cues or tactile stimuli to establish faeces
deposit sites either inside or outside nests. To build nests,
Stigmaeopsis females clean the target site on the leaf surface
through the silk-weaving mechanism. Furthermore, males kill con­
specific males to mate with immature females. The killing fre­
quency of the same species varies among the geographical
regions due to climatic changes.
There is still a need for more studies on the biology of
Stigmaeopsis that investigate the effects of constant and fluctuat­
ing temperature, geographic distribution, biological control, and
chemical control on species of this genus. There is no published
literature available on the management of most species.
Appropriate funding for basic research and training of specialists
to manage Stigmaeopsis spp. are important aspects to develop and
INTERNATIONAL JOURNAL OF ACAROLOGY 5
implement economically and ecologically sustainable manage­
ment programmes against Stigmaeopsis mites.
Acknowledgments
This study was supported by the National Natural Science
Foundation of China (31750002 and 31472034) and Annual
Program of the Ministry of Agriculture and Rural Affairs of PRC
for Integrated Control of Pests, Disease and Rats (19150061).
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
This work was supported by the Annual Program of the Ministry of
Agriculture and Rural Affairs of PRC for Integrated Control of Pests,
Disease and Rats [19150061]; National Natural Science Foundation
of China [31472034,31750002].
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