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Waking beauties: Mating quiescence in arachnid

females
Article · April 2023

DOI: 10.1016/bs.asb.2023.03.001
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Waking beauties: Mating

quiescence in arachnid females
Franco Cargneluttia,b,†, Fedra Bollattia,b,†, Matías A. Izquierdoa,b,†,
Debora Abregúb, Mariela Oviedo-Diegoa,b, David Vrecha,b,
Paola Oliveroa,c, Lucía Calbacho-Rosaa,b, Catalina Simiana,b,
Rocío Palen-Pietria,b, Camilo Mattonia,b, and Alfredo V. Perettia,b,⁎
a
Consejo Nacional de Investigaciones Cientı́ficas y Tecnicas (CONICET), Laboratorio de Biologı́a
Reproductiva y Evolución, Instituto de Diversidad y Ecologı́a Animal (IDEA), Córdoba, Argentina
b
Facultad de Ciencias Exactas, Fı́sicas y Naturales, Departamento de Diversidad Biológica y Ecologı́a,
Universidad Nacional de Córdoba, Córdoba, Argentina
c
Facultad de Ciencias Exactas, Fı́sicas y Naturales, Departamento de Fisiologı́a, Universidad Nacional de
Córdoba, Córdoba, Argentina
⁎
Corresponding author: e-mail address: aperetti@unc.edu.ar
Contents
1. Introduction 2
2. Literature search 50
3. Terminologies, history, and other concerns 51
4. General results of the survey 54
5. Quiescence and mating phase 55
6. Arachnids and quiescence levels 56
7. Behavioral and morphological traits associated with quiescence 57
7.1 Quiescence and female body position 58
7.2 Quiescence and female movements 60
7.3 Quiescence and male behaviors 61
7.4 Quiescence, aggression, and cannibalism 64
7.5 Quiescence and sexual size dimorphism 66
8. Quiescence and other biological traits in spiders 67
8.1 Quiescence and hunting strategies 67
8.2 Quiescence and copulation position 67
8.3 Quiescence and genitalia 69
9. Contexts that may have favored the evolution of female quiescence 72
9.1 Female sexual aggressiveness and cannibalism 72
9.2 Female and male genital complexity 76
9.3 Sperm transfer 79
†
The authors contributed equally to the work and should therefore be considered as first authors.
Advances in the Study of Behavior Copyright # 2023 Elsevier Inc. 1

ISSN 0065-3454 All rights reserved.
https://doi.org/10.1016/bs.asb.2023.03.001
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2 Franco Cargnelutti et al.
9.4 Copulation substrate constraints 81

9.5 Intersexual communication and mate choice 81
9.6 Postcopulatory sexual selection 86
10. Limitations of the present study 87
11. Conclusions and prospects 89
Acknowledgments 91
References 91
1. Introduction
Arachnids are a highly diverse group of arthropods with 13 living
orders (Ballesteros et al., 2022; Fig. 1). They have a great diversity of repro-
ductive modes and sperm transfer strategies. Some of these strategies can be
broadly classified as direct, mediated by a penis or modified body appendages,
as occurs in spiders (Araneae), harvestmen (Opiliones), solifuges (Solifugae),
some mites (Acariformes), and hooded tickspiders (Ricinulei). Other modes
or strategies can be considered indirect, mediated by spermatophores,
Fig. 1 The class Arachnida is composed of 13 orders and is sister group to the class
Pycnogonida, which includes the sea-spiders. The phylogeny is based on Ballesteros
et al. (2022) and unresolved clades are presented as a polytomy. The orders where
female mating quiescence occurs are marked with red dots. The forms of sperm transfer
in arachnids can be direct (D), by means of intromittent structures such as a penis, mod-
ified legs, or modified pedipalps, or indirect (I), by means of a spermatophore.
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Waking beauties: Mating quiescence in arachnid females 3
as occurs in scorpions (Scorpiones), pseudoscorpions (Pseudoscorpiones),

whip spiders (Amblypygi), whip scorpions (Uropygi), ticks (Parasitiformes),
most mites, and micro-whipscorpions (Schizomida) (Proctor, 1998; Fig. 1).
In spiders (Fig. 2A), for instance, the pedipalps in males are modified for direct
sperm transfer (Foelix, 2011). The sperm is deposited in a sperm web by the
recently molted male, collected by the pedipalp bulbs’ embolus, and stored
until copulation. A different strategy of direct sperm transfer occurs in solifuges
(Fig. 2B), which use their chelicerae to introduce a spermatophore or a ball of
sperm into the female genitalia. In harvestmen (Fig. 2C), direct sperm transfer
requires contact between genital structures (penis or spermatopositor in males,
ovipositor in females, Macı́as-Ordóñez, Machado, Perez-González, & Shultz,
2010). In some pseudoscorpions and mites, the male deposits spermatophores
on the substrate without encountering the female (Weygoldt, 1969, 1990). In
scorpions and some pseudoscorpions, indirect sperm transfer is preceded by a
mating dance (Olivero, Mattoni, & Peretti, 2017; Proctor, 1998; Weygoldt,
1969). During the dance, males search for an appropriate substrate to deposit
the spermatophore (Fig. 2D), which in some species can be morphologically
complex (Francke, 1979; Peretti, 2003; Weygoldt, 1969). After sperm trans-
fer, the mating pair separates (Olivero et al., 2017). In whip spiders, males
deposit spermatophores on the substrate for females to collect, which they
do using sclerotized claspers from their genitals (Weygoldt, 2000). In this
order, pair formation between sexes is a prerequisite for indirect sperm transfer
(Chapin & Hebets, 2016).
In animal reproduction, events that follow sperm transfer yet precede
fertilization can be shaped by postcopulatory sexual selection. These hidden
but strong selective mechanisms establish a field rich for quantitative surveys
and theoretical assays (Eberhard, 2009). They offer the chance to explore
issues related to sperm competition and cryptic female choice in an evolu-
tionary framework covering aspects of behavior, physiology, and genetics
(Andersson, 1994; Birkhead & Pizzari, 2002; Eberhard, 1996; Parker,
1970, 1984, 1998). In the case of arachnids, however, many aspects of their
sexual biology remain unknown or poorly explored. An intriguing behavior
reported for certain spiders (and also other arachnids) is that females adopt a
“passive” behavior and remain immobile in an apparent state of catalepsy
before, during, and after mating (e.g., Aisenberg & Costa, 2005; Just,
Opatova, & Dolejš, 2018; Xiao et al., 2015) (Fig. 2E). This behavior,
referred to most generally as “quiescence,” has also been reported in other
animals, including vertebrates and insects. Among vertebrates, for example,
females of the North American-tailed frog Ascaphus truei adopt a state of
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Fig. 2 Arachnids and their reproductive mode. (A) A mating pair of a wolf spider from
the Thorelli group (Lycosidae). The male is on top of the female, which remains quies-
cent (level III). Note the female’s extended legs and the position of her abdomen
during swiveling behavior. (B) A mating pair of a solifuge of the genus Eremobates
(Eremobatidae). Note how the female (left) bends her abdomen forward while the
male transfers sperm using his chelicerae. (C) The harvestman Iporangaia pustulosa
(Gonyleptidae), a species with exclusive paternal care. Both male (left) and female
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so-called “tonic immobility” during first episodes of amplexus (Noble &

Putnam, 1931). In this species, fertilization is internal and is accomplished
by an intromittent organ, the tail of the males (Stephenson & Verrell,
2003). The reason for the female’s immobility is uncertain, and authors have
proposed that it may facilitate intromission (Noble & Putnam, 1931), or per-
haps prevent it (Metter, 1964). Shapiro (1936) suggested that the female’s
role in copulation is passive, important for the successful coupling reflex,
whereas the male’s role is active. The author also speculates that the male
is unable to grip the female unless she exhibits a certain degree of passivity.
Interestingly, female immobility may last for several minutes after she has been
released by the male, being the strong grip during the amplexus necessary to
accomplish the immobility (Noble & Putnam, 1931; Stephenson & Verrell,
2003; Wernz & Storm, 1969). Another example of female immobility in ver-
tebrates occurs in rats. Here, however, the pair (not just the female) remains
immobile for a few seconds, during which time the male deposits sperm and a
coagulated copulatory plug into the vagina. Immobility might facilitate the
formation of tighter plugs and hence the transfer of more sperm (Adler &
Toner, 1986; Matthews & Adler, 1977, 1978, 1979).
In insects, a state of female quiescence during mating has been reported
in species belonging to different orders. In the flour beetle Tribolium
castaneum, females stop moving during copulation, although they will per-
form postural changes including foreleg extension, head lifting, and anten-
nae elevation (Bloch-Qazi, 2003). Longer female quiescence seems to reflect
longer relaxation of female bursal muscles, allowing males more time to
inject sperm into his spermatophore (Fedina, 2007). Relatedly, longer
are quiescent after copulation. (D) A scorpion of the species Timogenes dorbignyi
(Bothriuridae) performing a mating dance (or promenade à deux). During this phase
of the courtship, the male (left) and female interlock their pedipalps and walk together
until the male deposits the spermatophore on the substrate. (E) After copulation, in the
spider Agelenopsis pennsylvanica (Agelenidae), the female (below) remains quiescent
(level III), lying on her side. (F) A mating pair of the scorpion Timogenes dorbignyi
(Bothriuridae), showing the moment of sexual stinging. The male (right) stings the
female pedipalp with the stinger in telson during copulatory courtship. (G) A mating
pair of the tarantula Sickius longibulbi (Theraphosidae) with the male in a vertical posi-
tion and the female lying on her back. (H) A mating pair of the tarantula Acanthogonatus
centralis (Pycnothelidae) after copulation. Note how the female stays in the copulatory
position after male separation. Photo credits: (A) Matías A. Izquierdo; (B) Marco Zozaya;
(C) Gustavo S. Requena; (D) Camilo I. Mattoni; (E) Brandi Pessman; (F) Camilo I. Mattoni;
(G) Rogerio Bertani; (H) Sofía Copperi.
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quiescence duration has been associated with increased numbers of sperm

transferred to females, as well as higher second male sperm precedence
(Bloch-Qazi, 2003). In the earwig Vostox apicedentatus, female quiescence
occurs during courtship as the male taps the female’s abdomen and grabs
her with his forceps (Moore & Wilson, 1993). Finally, in the moth
Utetheisa ornatrix, quiescent females tend to expose their abdomens when
stimulated with an excised coremata, a male extrusive brush organ that
produces a pheromone (Eisner & Meinwald, 1995).
Although the reasons for female immobility varied in the cases men-
tioned above, we can discern a common factor in all: females’ lack of move-
ment was not prompted by males. Rather, we can infer that the females’
immobility was started voluntarily. Part of the difficulty in reaching this
inference is that some prior terminologies seem to automatically cancel
out the possibility that females are making active decisions. For example,
the term “passivity” can might imply in females a lack of interest, or lack
of active response, or both. A lack of attention to female behaviors during
copulation predates Darwin’s (1871) ideas of sexual choices in evolution,
and continues to the present day. Often, females are considered passive or
even just mere receptors of sperm who did not participate actively as mates.
Consequently, Darwin’s emphasis on the evolutionary role of females has
stimulated rich reflections on other current disciplines such as gender studies
(Cohen, 2010). Although his theory of natural selection continues to be
broadly accepted and supported by data, Darwin’s theory of sexual selection
was not immediately embraced, and debates still persist (Carranza, 2010).
Much debate focuses on female choice and the anthropocentric biases that
have colored some conceptions (Ah-King, 2011). There is good reason to
believe that some presuppositions in animal behavior research reflect biases
in the ideological values of society at large (Ah-King, 2007).
The undervalued role that females can play during copulation is also evi-
dent in arachnids, in which a plethora of terms have been proposed to
explain female mating immobility (Tables 1–5; Fig. 3A). In certain spiders,
for example, catalepsy (or cataleptic posture) has been defined as the lack of
response to external stimuli, and is usually accompanied by leg curving as
the mating pair makes contact (Becker et al., 2005). Such a state is apparently
triggered by chemicals that males release during copulatory behavior
(Becker et al., 2005). The body postural change of some spiders is reminis-
cent of that already mentioned in the flour beetle Tribolium castaneum.
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Table 1 Spiders for which female mating quiescence has been reported. The
table shows some aspects of the biology of the group that may be related to the
presence of female mating quiescence.
Sexual Mating Web Genital
Species cannibalism SSD position Lifestyle type type
Araneomorphae: Agelenidae (RTA)

Agelena labyrinthica No No III Web builder SF Entelegyne
Agelenopsis aperta Yes No III Web builder SF Entelegyne
Agelenopsis Yes No III Web builder SF Entelegyne
pennsylvanica
Cabolena huiztocatl No No III Web builder SF Entelegyne
Coelotes atropos No No III Web builder SF Entelegyne
Histopona torpida No No III Web builder SF Entelegyne
Hololena adnexa Yes No III Web builder SF Entelegyne
Hololena curta No No III Web builder SF Entelegyne
Wadotes bimucronatus No No III Web builder SF Entelegyne
Araneomorphae: Cybaeidae (RTA)
Cedicus israeliensis No No III Web builder SF Entelegyne
Cybaeus angustiarum No No III Web builder SF Entelegyne
Araneomorphae: Lycosidae (RTA)
Aglaoctenus oblongus Yes No III Web builder SF Entelegyne
Alopecosa albofasciata No No III Wandering – Entelegyne
Alopecosa cuneata No No III Wandering – Entelegyne
Alopecosa farinosa No No III Wandering – Entelegyne
(Czech population)
Alopecosa farinosa No No III Wandering – Entelegyne
(Italian population)
Alopecosa inquilina No No III Wandering – Entelegyne
Alopecosa mariae No No III Wandering – Entelegyne
Alopecosa pinetorum No No III Wandering – Entelegyne
Alopecosa psammophila No No III Wandering – Entelegyne
Alopecosa pulverulenta No No III Wandering – Entelegyne
Alopecosa schmidti No No III Wandering – Entelegyne
Continued
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presence of female mating quiescence.—cont’d
Alopecosa solitaria Yes No III Wandering – Entelegyne

Alopecosa striatipes No No III Wandering – Entelegyne
Alopecosa sulzeri No No III Wandering – Entelegyne
Alopecosa taeniata No No III Wandering – Entelegyne
Alopecosa trabalis No No III Wandering – Entelegyne
Arctosa alpigena No F > M III Wandering – Entelegyne
lamperti
Arctosa littoralis Yes No III Wandering – Entelegyne
Arctosa lutetiana No No III Wandering – Entelegyne
Diapontia uruguayensis No No III Web builder SF Entelegyne
Geolycosa turricola Yes No III Sedentary OB Entelegyne
Hogna gumia No No III Wandering – Entelegyne
Hogna sp. No No III Wandering – Entelegyne
Lycosa carbonelli No No III Wandering – Entelegyne
Lycosa chaperi No No III Wandering – Entelegyne
Lycosa inornata Yes No III Wandering – Entelegyne
Lycosa poliostoma No No III Wandering – Entelegyne
(Argentina
population)
Lycosa poliostoma No No III Wandering – Entelegyne
(Uruguay
population)
Lycosa thorelli No No III Wandering – Entelegyne
Pardosa lapidicina No No III Wandering – Entelegyne
Rabidosa rabida Yes No III Wandering – Entelegyne
Rabidosa santrita No No III Wandering – Entelegyne
Schizocosa malitiosa Yes No III Wandering – Entelegyne
Schizocosa ocreata Yes No III Wandering – Entelegyne
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Tigrosa helluo Yes No III Wandering – Entelegyne

Trochosa robusta No No III Wandering – Entelegyne
Trochosa ruricola No No III Wandering – Entelegyne
Trochosa spinipalpis No No III Wandering – Entelegyne
Trochosa terricola Yes No III Wandering – Entelegyne
Araneomorphae: Homalonychidae (RTA)
Homalonychus Yes No III Wandering – Entelegyne
selenopoides
Homalonychus Yes No III1 Wandering – Entelegyne
theologus
Araneomorphae: Philodromidae (RTA-D)
Thanatus fabricii Yes No III Wandering – Entelegyne
Araneomorphae: Desidae (RTA-M)
Metaltella iheringi Yes No II Web builder SCr Entelegyne
Metaltella simoni No No II Web builder SCr Entelegyne
Araneomorphae: Oxyopidae (RTA-OC)
Oxyopes schenkeli Yes No III Wandering – Entelegyne
Araneomorphae: Pisauridae (RTA-OC)
Pisaurina mira Yes No III Wandering – Entelegyne
Araneomorphae: Thomiside (RTA-OC)
Xysticus ferox Yes F > M III Wandering – Entelegyne
Araneomorphae: Hypochilidae (NE)
Hypochilus pococki Yes No I Web builder LS Haplogyne
Araneomorphae: Theridiidae (ARA)
Argyrodes antipodianus Yes No II Web builder Kp Entelegyne
Echinotheridion Yes F > M II Web builder TS Entelegyne
gibberosum
Latrodectus bishopi No F > M II Web builder TS Entelegyne
Continued
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Latrodectus geometricus No F > M II Web builder TS Entelegyne

Latrodectus hasselti Yes F > M II Web builder TS Entelegyne
Latrodectus hesperus Yes F > M II Web builder TS Entelegyne
Latrodectus mactans Yes F > M II Web builder TS Entelegyne
Latrodectus pallidus Yes F > M II Web builder TS Entelegyne
Latrodectus revivensis Yes F > M II Web builder TS Entelegyne
Latrodectus No F > M II Web builder TS Entelegyne
tredecimguttatus
Tidarren sisyphoides No F > M II Web builder TS Entelegyne
Mygalomorphae: Atracidae (AVV)
Atrax robustus No No I Web builder SLB Haplogyne
Mygalomorphae: Euagridae (AVI)
Euagrus sp. No No I Sedentary MFS Haplogyne
Mygalomorphae: Ischnothelidae (AVI)
Thelechoris striatipes Yes No I Web builder PC Haplogyne
Mygalomorphae: Mecicobothriidae (ATY)
Mecicobothrium thorelli Yes No I Web builder DSF Haplogyne
Mygalomorphae: Microhexuridae (AVI)
Microhexura montivaga No No I Web builder TSW Haplogyne
Mygalomorphae: Pycnothelidae (AVN)
Acanthogonatus No No I Web builder T Haplogyne
centralis
Acanthogonatus No No I Web builder T Haplogyne
tacuariensis
Xenonemesia platensis No No I Sendentary SC Haplogyne
Mygalomorphae: Theraphosidae (AVC)
Aphonopelma hentzi No No I Sendentary B Haplogyne
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Eupalaestrus No No I Sendentary B Haplogyne

weijenberghi
Grammostola vachoni Yes No I Sedentary B Haplogyne
Plesiopelma No No I Sedentary B Haplogyne
longisternale
Sickius longibulbi No No I2 Sedentary B Haplogyne
Vitalius roseus Yes No I Sedentary B Haplogyne
Except for the “Non-Entelegyne” (NE), families are organized into major clades according to Hedin et al.
(2019), Wheeler et al. (2016), and Opatova et al. (2020) as follows: ARA ¼ Araneoidea, ATY ¼ Atypoidea,
AVI ¼ Avicularioidea, AVC ¼ Avicularioidea, Crassitarsae, AVN ¼ Avicularioidea, “Nemesioidina,” AVV ¼
Avicularioidea, venom clade, RTA ¼ Retrolateral Tibial Apophysis clade, RTA-D ¼ RTA, Dionycha,
RTA-OC ¼ RTA, Oval Calamistrum, RTA-M ¼ RTA, Marronoid. SSD ¼ Sexual size dimorphism
(F ¼ female; M ¼ Male). For a description of the mating positions, see Section 8.2. Web types: B ¼ Burrow-
er, DSF ¼ Dense-sheet funnel web, Kp ¼ Kleptoparasitic of orb-web spiders, LS ¼ Lampshade shaped
web, MFS ¼ Mixed funnels and sheets with retreat, OB ¼ Obligate burrower, PC ¼ Perennial capture web,
SC ¼ Shallow cave, SF ¼ sheet web with funnel, Scr ¼ Sheet cribellate, SLB ¼ Silk-lined tubular retreats
with funnel, T ¼ Tunnel web, TS ¼ Tangle space web-gumfoot-web, TSW ¼ Tube-shaped web. Sources of
information are presented in Table 2.
However, as we will show, female immobility may not necessarily include

any visible postural change. In this sense, there may be different levels of
female quiescence, and there is no consensus on which terminology should
be used in each case.
Our main goal moving forward is to provide a qualitative review of the
occurrence of female mating quiescence in arachnids (excluding the orders
Acariformes and Parasitiformes), and to consider its consequences in a repro-
ductive context. Based on the results of an intensive literature search, we
classify female mating quiescence as it occurs at different levels, and propose
a terminology that best represents our findings. Additionally, we reexamine
the question of female quiescence is triggered by male behavior, or whether
it results from females’ decisions. As part of this effort, we also evaluate
if hypotheses proposed for female mating quiescence in some taxa can be
applied to others. Finally, we suggest possible contexts that may favor in
arachnids the evolution of female mating quiescence.
Table 2 Reports of female mating quiescence in the order Araneae. Columns summarize information extracted from each paper (NA ¼ information non-
available).
Mating phase Female body Female
when position during movement Terms used /
quiescence Male behavior before female quiescence during Quiescence Functional
Species occurs quiescence (Quiescence Level) quiescence duration hypothesis Source
Araneomorphae: Agelenadae (RTA)
Agelena Cop+Post Male pulls the female web with his Flexed legs near the NA 30 min C / None 1–3
labyrinthica legs. body (III).
Agelenopsis Cop+Post Male approaches, mounts the Legs lie close to the NA NA Q, C / Female 4–6
aperta female and touches the sides of her body sides, distal sexual
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abdomen “inducing” catalepsy, portions extended aggressiveness
then the male positions the female far outwards, leg IV and cannibalism
on her side. Male “induces” the directly posteriorly female, Female
cataleptic state of the female with and slightly and male genital
short distance pheromones by using outward (III). complexity
the pedipalps or by contact.
Agelenopsis Cop Not reported, but authors mention Legs tucked around NA NA C / None 7
pennsylvanica the action of pheromones. the cephalothorax
(III).
Cabolena Pre + Cop NA Not clear, but in NA 38 min Q1 / None 8
huiztocatl one mating
observation, the
male held the
female with four
legs on the same
side of the
pedipalp used to
copulate (III).
Coelotes atropos Cop NA “Typical position NA NA C / None 3, 9
of agelenids” (III).
Histopona Pre + Cop At the first touch of the male, the “Typical position NA NA C / None 3, 10
torpida female places her legs near the body. of tube web
The male grabs the female body entelegynes” (III).
near the tube retreat.
Hololena adnexa Pre + Cop Male performs vibrations on the Flexed legs near the NA Few seconds C / Female sexual 11
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+Post female’s web. body (III). post copulation aggressiveness
and cannibalism,
Postcopulatory
sexual selection
(CFC)
Hololena curta* Pre + Cop Female becomes cataleptic when Flexed legs (III). None. 48–196 min C / Female sexual 12
the male touches and mounts her. aggressiveness
Also, when female shows and cannibalism,
aggressivity, male tremulates his Postcopulatory
abdomen and grasps her leg at sexual selection
which point she immediately (CFC)
becomes cataleptic.
Wadotes Pre + Cop Male grips the female’s patella/tibia Legs flexed over the NA NA C / None 13
bimucronatus joints with his chelicerae. carapace and below
the sternum (III).
Continued
available).—cont’d
Araneomorphae: Cybaeidae (RTA)
Cedicus Pre + Cop NA Legs close to the NA NA C / None 2, 3
israeliensis body (III).
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Cybaeus Pre + Cop Male suddenly jumps on the female, Rigid body (III). NA NA C / None 3, 14
angustiarum which remains quiet. The male
grabs the female and turns her rigid
body until he is in front of her.
Araneomorphae: Lycosidae (RTA)
Aglaoctenus Post NA Extended legs (II). None. Several minutes M / None 15
oblongus
Alopecosa Cop Male begins courting with Front legs extended Swiveling the NA C / None 16
albofasciata inconspicuous pedipalpal frontwards, the tip abdomen.
movements, which escalates over of the tarsus
time, followed by escalating jerky pointing forward
steps. When courtship peaks, male (III).
movements include leg attracting to
the body. Receptive females
respond becoming cataleptic.
Alopecosa Cop When the male climbs onto the Front legs extended NA NA C / None 16
cuneata female’s back, she becomes frontwards, the tip
cataleptic. of the tarsus
pointing forward
(III).
Alopecosa Cop Male pedipalps are spread and rub Front legs extended NA NA C / None 16
farinosa (Czech the substrate circularly. Male frontwards, the tip
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population) performs jerky body movements. of the tarsus
Male raises his legs. Upon contact, pointing forward
female becomes cataleptic. (III).
Alopecosa Cop NA Front legs extended NA NA C / None 16
farinosa (Italian frontwards, the tip
population) of the tarsus
pointing forward
(III).
Alopecosa Cop Male approaches the female with his Hidden legs, front NA NA C / None 16
inquilina front legs quivering and pointing legs retracted, the
forward. The front leg closer to the tip of the tarsus
female usually quivers more rapidly. points straight
After 15 min, female becomes down or below the
cataleptic. body (III).
Continued
Alopecosa Cop When close to the females, Exposed legs, NA NA C / None 16
mariae male performs sideways steps. anterior legs
After courtship, female becomes towards the front,
cataleptic. tip of the tarsus
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pointing
forward (III).
Alopecosa Cop NA Hidden legs, front NA NA C / None 16
pinetorum legs retracted, the
tip of the tarsus
points straight
down or below the
body (III).
Alopecosa Cop When close to the female, the male Hidden legs, front NA NA C / None 16, 17
psammophila performs cymbial circling: the legs retracted, the
pedipalps are spread and rub the tip of the tarsus
substrate with their prolateral side points straight
while performing circular motions. down or below the
Also, leg quivering is observed and body (III).
stretching of both forelegs laterally,
upwards and outwards, in front
of the female.
Alopecosa Cop Male begins to court with cymbial Exposed legs, NA NA C / None 16, 17
pulverulenta rubbing. Then, he approaches the anterior legs
female with body jerky steps, body towards the front,
crawling, body dropping and tip of the tarsus
abdominal tapping. After courtship, pointing
female becomes cataleptic. forward (III).
Alopecosa Cop Pedipalpal movement is intensive Exposed legs, NA NA C / None 16
ARTICLE IN PRESS
schmidti and escalates in body jerky steps and anterior legs
leg quivering. Female responds towards the front,
offensively, a wrestle occurs before tip of the tarsus
the female becomes cataleptic. pointing
forward (III).
Alopecosa Cop Courting is composed of cymbial Exposed legs, NA NA C / None 16
solitaria circling and abdominal tapping. anterior legs
Subsequently, female becomes towards the front,
cataleptic. tip of the tarsus
pointing
forward (III).
Alopecosa Cop Male begins to court immediately Exposed legs, NA NA C / None 16
striatipes by cymbial rubbing and approaches anterior legs
the female with rapid jerky steps. towards the front,
His front legs are raised, pointing tip of the tarsus
slightly upwards, and perform leg pointing
quivering. forward (III).
Continued
Alopecosa Cop Females often become cataleptic Exposed legs, NA NA C / None 16
sulzeri without being in physical contact anterior legs
with males. Females lay on their side towards the front,
before mating in some cases. tip of the tarsus
pointing
ARTICLE IN PRESS
forward (III).
Alopecosa Cop Male climbs onto the female’s back, Exposed legs, NA NA C / None 16
taeniata which “causes” her to become anterior legs
cataleptic. towards the front,
tip of the tarsus
pointing
forward (III).
Alopecosa Cop Courtship is initiated by escalating Exposed legs, NA NA C / None 16, 17
trabalis pedipalpal movements, which anterior legs
results in body jerky steps with body towards the front,
dropping and abdomen tapping. tip of the tarsus
The front legs are raised, pointed pointing
forward, and occasionally dropped forward (III).
to the substrate.
Arctosa alpigena Cop Male contacts the female Front segs bent, NA NA C / Intersexual 18
lamperti cephalothorax by using his first pair hind legs communication
of legs. Male’s second pair of legs spread (III).
contacts the front legs of the female.
Arctosa littoralis Post Male raises and lowers the pedipalps NA (?). NA 1.5–3.5 min Q / None 19
in an alternate manner without
drumming. This behavior is
followed by a rapid quivering
motion of first pair of legs. Female
responds with the same pattern of
leg movements but to a lesser
height. Pursuit is also observed.
Arctosa lutetiana Cop NA Extended legs, Swiveling the NA Q / None 20
swiveling of abdomen and
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abdomen and erection of
erection of spines spines.
(III).
Diapontia Post NA Motionless in One female Several minutes M / None 21
uruguayensis copulatory position became active
for several minutes five times and
after mating while walked
the males move around with
away (III). the male still
on her
dorsum. The
rest of the
females
remain
motionless.
Geolycosa Post NA NA (?). NA NA Q / None 22
turricola
Hogna gumia Post1 NA NA (?). None. 25.3 17.9 min C / None 23
Continued
Hogna sp. Cop+Post Male advances slowly and courts the Body lies on the NA NA C / Intersexual 24
female raising his forelegs and then ground (III). communication
lowers them slowly while waving
(alternately or simultaneously) the
conspicuous black tarsi. Drumming
of the pedipalps is also observed. At
ARTICLE IN PRESS
the time of mating, male advances
very slowly towards female with his
forelegs extended and semi-raised
and mounts her in one leap.
Lycosa carbonelli Cop+Post Behavior “typical of other Body lies on the At the end of 15 s post C / None 25
lycosids”. ground (III). copulation, copulation
78% of the
females walk
dragging the
male for
10 cm.
Lycosa chaperi Pre + Cop NA NA (?). NA NA C / Female and 26
male genital
complexity
Lycosa inornata Post NA Extended legs, None. 0.2 0.9 min C, S / None 27
swiveling of post copulation
abdomen and
erection of
spines (III).
Lycosa Post NA In the same Total stillness 1.4 2.4 min C / None 23
poliostoma (Arg position as she was of the female
population) during mating (III). until she
reacts
suddenly and
runs away.
Lycosa Cop+Post NA Extended legs (III). None. 4.1 min C / None 28
poliostoma (Uru
ARTICLE IN PRESS
population)
Lycosa thorelli Cop+Post Behavior “typical of other Body lies on the None. NA C / None 25
lycosids”. ground (III).
Pardosa Post Male raises and lowers his pedipalps, NA (?). NA < 1 min Q / None 29
lapidicina skittering legs and pursuing female.
Rabidosa Pre + Cop Male mounts the female by Sternum near the Female 6 min C, I / Female 30, 31
rabida* +Post climbing onto her depressed ground, abdomen rotates her sexual
cephalothorax. in the air, legs abdomen aggressiveness
extended straight about its and cannibalism,
on the ground (III). longitudinal Intersexual
axis during communication
each crossing
by the male,
bringing the
epigynum
within reach
of the male’s
pedipalp.
Continued
Rabidosa Post Male touches the first pair of legs Extended forelegs NA 5 min M / Intersexual 32
santrita of the female. (III). communication
Schizocosa Cop+Post NA Body lies on the Abdomen 6 min A, C / None 33–35
malitiosa ground with rotation and
forelegs extended, convulsive
hindlegs extended movements
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backward, and with body
abdomen slightly elevation and
elevated (III). leg extension.
Schizocosa Cop NA Body lies on the NA NA Q2 / None 36
ocreata ground with
forelegs extended,
hindlegs extended
backward, and
abdomen slightly
elevated (III).
Tigrosa helluo Cop+Post Male touches the first pair of legs First and second None, except 11 min C, Q / Female 37, 38
of the female. pair of legs for short sexual
extended, periods of aggressiveness
depressed low walking. and cannibalism,
cephalothorax. Abdomen Intersexual
Legs remain movements communication
extended (III). may facilitate
the insertion
of pedipalps.
Trochosa robusta Cop+Post Apparently similar to T. terricola. Apparently similar None. 25 s-9 min A / None (but 39
to T. terricola (III). (mean 2.4 min) probably Female
sexual
aggressiveness
and cannibalism)
Trochosa ruricola Cop+Post Apparently similar to T. terricola. Apparently similar None. 10s-8.4 min A / None (but 39
to T. terricola (III). (mean 4.2 min) probably Female
sexual
aggressiveness
and cannibalism)
ARTICLE IN PRESS
Trochosa Cop+Post Apparently similar to T. terricola. Apparently similar None. 10s-12 min A / None (but 39
spinipalpis to T. terricola (III). (mean 5.1 min) probably Female
sexual
aggressiveness
and cannibalism)
Trochosa Cop+Post Male performs abdominal “Disposition None. 10s-10 min A / Female sexual 39
terricola vibrations, swing of front legs, and posture”: legs (mean 5 min) aggressiveness
leg touching. extended forward and cannibalism
and parallel to the
body.
Cephalothorax
inclined forward
(III).
Araneomorphae: Homalonychidae (RTA)
Homalonychus Cop+Post Male touches the female. Front legs flexed; NA NA P, M, 40
selenopoides hind legs extended. QS / None
Nuptial veil is
deposited in the
legs (III).
Continued
Homalonychus Pre + Cop Male approaches the female, drums Motionless on the NA NA M / None 41
theologus +Post his pedipalps in an alternating substrate. Drawn
sequence, and attempts to mount legs close to her
her. Then he moves his front pairs body, the patellae
ARTICLE IN PRESS
of legs alternately up and down, almost touching
tapping the substratum, walking a one another above
few steps, and stopping—mounting the carapace (III).
proceeds while female remains
immobile.
Araneomorphae: Philodromidae (RTA-D)
Thanatus fabricii Cop+Post Male bites the female. Female Legs close to the In four cases 34.6 min Im, T / Female 42
becomes immobile when the body (III). females sexual
nuptial veil is deposited. started to aggressiveness
move after and cannibalism,
the last Intersexual
pedipalp communication,
insertion. Mate choice,
Postcopulatory
sexual selection
Araneomorphae: Desidae (RTA-M)
Metaltella Cop+Post “Pseudo-mounts”. Flexed legs, Gentle 5 min post Q / None 43
iheringi sometimes forelegs movements copulation
extended (III). of chelicerae
and legs.
Metaltella Cop+Post “Pseudo-mounts”. Flexed legs, NA NA Q / None 44
simoni sometimes forelegs
extended (III).
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Araneomorphae: Oxyopidae (RTA-OC)
Oxyopes Pre Female lets herself hang on a string Female remains Female can NA M / None 45
schenkeli when the male tries to mount her. immobile until she instantly
Then, male comes down and starts hangs up, legs are shrug off her
to wrap her, but the female does not nor retracted (III). bridal veil and
shrink her legs. returned to
her original
position on
the blade.
Araneomorphae: Pisauridae (RTA-OC)
Pisaurina mira Pre + Cop Male approaches from posterior and Suspended free in NA NA C / None 46
moves to a location directly above space, face
the female. He moves laterally downward,
across the edge of his perch as she inactive and
brings her legs I and II against her tethered only by a
cephalothorax and descends freely dragline (III).
on her dragline.
Continued
Araneomorphae: Thomiside (RTA-OC)
Xysticus ferox Cop Male grabs the female’s legs with his Flexed legs near the NA NA Im / None 47
legs. body (III).
ARTICLE IN PRESS
Araneomorphae: Hypochilidae (NE)
Hypochilus Cop Leg and pedipalp grooming, Abdomen in 45° NA NA SC / None 48
pococki bobbing, and web-tugging by the with the soil (II).
male. Male and female perform leg
stroking.
Araneomorphae: Theridiidae (ARA)
Argyrodes Pre + Cop Pedipalp and leg touching by the NA (?). NA NA Q / None 49
antipodianus male on the female.
Echinotheridion Cop NA NA (?). NA 4 min C / None 50
gibberosum
Latrodectus Cop Abdominal vibrations. Male NA (?). None NA Im, I / None 51
bishopi touches the first pair of legs of the
female with his legs.
Latrodectus Cop+Post Abdominal vibrations while NA (?). Female can NA M / None 52
geometricus walking, jerking, twanging move and
threads, web biting, and web take away the
reduction. Palpation of female’s legs nuptial veil.
and body with the male’s legs
and pedipalps. Male climbs upon
the female’s venter, palpates,
and drums against her venter and
epigynum. He may also tie the
female with strands of silk.
ARTICLE IN PRESS
Latrodectus NA NA NA (?). None. NA Q / Sperm 53
hasselti transfer
Latrodectus Pre Touches on female’s abdomen and NA (?). 77% of the 22 min before Q / Female 54, 55
hesperus legs. Web reduction. Nuptial veil, females move copulation sexual
abdominal vibrations, body during aggressiveness
shakings, drumming on the female’s quiescence. and cannibalism
abdomen, and attempts to insert
the pedipalps.
Latrodectus Pre + Cop Male cuts threads of silk, then wraps NA (?). NA NA TS / Female 56, 57
mactans the female (nuptial veil). sexual
aggressiveness
and cannibalism
Latrodectus Cop+Post Drumming upon the female threads Ventral side of Leg NA CA, Qt / Female 58
pallidus with the pedipalps. Spinning abdomen upwards movements sexual
of threads around the female. (II). when the aggressiveness
male touch and cannibalism
the female
legs abruptly.
Continued
Latrodectus Cop+Post Male wraps female’s legs and Lowering of the None. NA CS / Female 59
revivensis body (nuptial veil). body (II). sexual
aggressiveness
and cannibalism,
ARTICLE IN PRESS
Mate choice
Latrodectus Cop Male shakes female’s web with his NA (?). NA NA H, CA / None 58
tredecimguttatus legs, swaddles the female with fine
threads, and drums upon her body
until she turns on her back.
Tidarren Cop Pedipalp and leg movements Lowering of the None NA C / None 60
sisyphoides (touching of female abdomen and body (I).
hind legs). Male dies while the
female becomes cataleptic.
Mygalomorphae: Atracidae (AVV)
Atrax robustus Cop Male lifts the female body until Vertical position, Movements NA PS, QS / None 61
she assumes a vertical position. exposing the are only
epigastric region observed
(I). during the
vertical
position of
the female.
Mygalomorphae: Euagridae (AVI)
Euagrus sp. Cop NA Flexed legs (I). NA NA C / None 62
Mygalomorphae: Ischnothelidae (AVI)
Thelechoris Cop+Post Clasping of male’s first tibial Legs and pedipalps Occasional 2 s after CS / None 63
striatipes apophysis with female’s pedipalp. partially flexed quivering or copulation
(except for other
occasional movements.
quivering or other Genital area
ARTICLE IN PRESS
movement) (I). was distended
and more
exposed in
some
specimens.
Mygalomorphae: Mecicobothriidae (ATY)
Mecicobothrium Cop+Post Body vibrations, leg III movements, Legs are flexed, The female NA P / None 64
thorelli pedipalpal drumming, and leg “typical position of can finish the
tapping on the female’s body mygalomorphs” (I). clasping.
and legs. Synchronic
abdominal
movements.
Mygalomorphae: Microhexuridae (AVI)
Microhexura Cop Clasping, male pedipalps vibrations. Legs without None. 11–19 min CS, C / None 65
montivaga movements. Hind
legs and abdomen
rest on the soil (I).
Continued
Mygalomorphae: Pycnothelidae (AVN)
Acanthogonatus Cop+Post Body shaking and tapping on the Over the male, None. 27.1 29.8 s C/ 66, 67
centralis female’s body. “typical position of during Postcopulatory
ARTICLE IN PRESS
mygalomorphs” (I). copulation; sexual selection,
23.7 36.7 s Mate choice
post copulation
Acanthogonatus Pre + Cop Male touches the female with In a 90° between NA NA P / Intersexual 68
tacuariensis his anterior legs. both body axes (I). communication,
Mate choice
Xenonemesia Pre + Cop Only contact is mentioned. Over the male, NA NA M / None 67, 69
platensis +Post “typical position
of mygalomorphs”
(II).
Mygalomorphae: Theraphosidae (AVC)
Aphonopelma Cop+Post Male and female adopt the Abdomen on the NA NA M / None 70
hentzi “typical copulatory position floor, legs
of mygalomorphs”. extended (I).
Eupalaestrus Cop+Post NA First pair of legs and NA NA Im, Q / 71
weijenberghi palps elevated in Postcopulatory
threat position (II). sexual selection
(CFC)
Grammostola Cop+Post Palpation, pedipalpal drumming, The female NA 92.3 113.6 s C, PS / 66, 72
vachoni vibration, and leg tapping. remained quiescent Postcopulatory
with her sexual selection
ARTICLE IN PRESS
cephalothorax and (CFC)
abdomen bent
at an angle of
60°-80° (I).
Plesiopelma Post Palp drumming, body vibrations, Probably similar to NA 30.6 61.2 s C / None 66
longisternale leg tapping on the female body, other
clasping of male’s tibial apophysis mygalomorphs
with female’s pedipalps. with clasping
behavior,
quiescence level
not reported but
inferred (I).
Sickius Cop+Post Male bites the female. Legs NA NA M/ 73
longibulbi semi-extended Postcopulatory
(III). sexual selection
Continued
Vitalius roseus NA Body vibrations, tapping the female NA (?). NA NA C / Female sexual 74
with extended forelegs and aggressiveness
pedipalpal drumming over the and cannibalism,
female’s genital zone. Female and male
ARTICLE IN PRESS
genital
complexity
Species marked with an asterisk indicate papers that provide detailed behavioral description of female quiescence. Abbreviations of major clades are presented in Table 1. Mating phases:
Pre ¼ precopulatory, Cop ¼ copulatory, Post ¼ postcopulatory. Quiescence levels are presented in Table 6. Terms used to describe female mating quiescence in the original papers are:
A ¼ Akinesis, C ¼ Catalepsy, CA ¼ Calm, CS ¼ Cataleptic State, H ¼ Hypnosis, I ¼ Inactivity, Im ¼ Immobility, M ¼ Motionless, P ¼ Passive, PS ¼ Passive State, Q ¼ Quiescence, QS ¼
Quiescent State, Qt ¼ Quiet, S ¼ Stillness, SC ¼ Semi Cataleptic, T ¼ Thanatosis, TS ¼ Trance State. Names of the functional hypotheses are the same as presented in Box 1
(CFC ¼ cryptic female choice). Uppercase notes: (1) According to the ethogram by Schwerdt, Pompozzi, Copperi, and Ferretti (2014); (2) Proposed in this paper based on a video
(see reference in this table).
(1) Preston-Mafham and Preston-Mafham (1993), (2) Gerhardt (1921), (3) Huber (1998), (4) Gering (1953), (5) Singer et al. (2000), (6) Becker, Riechert, and Singer (2005), (7) Pruitt et al.
(2014), (8) Maya-Morales, Jimenez, Murugan, and Palacios-Cardiel (2017), (9) Gerhardt (1928), (10) Gerhardt (1927), (11) Fraser (1987), (12) Xiao et al. (2015), (13) Bennet (1987),
(14) Gerhardt (1933), (15) González and Toscano-Gadea (2020), (16) Just et al. (2018), (17) Chiarle, Kronestedt, and Isaia (2013),(18) Dolejš, Kubcová, and Buchar (2012), (19) Punzo
(2006), (20) Dolejš, Kubcová, and Buchar (2010), (21) Piacentini et al. (2017), (22) Miller and Miller (1987), (23) Schwerdt et al. (2014), (24) Costa and González (2015), (25) Costa
and Capocasale (1984), (26) Sadana (1972), (27) Toscano-Gadea and González (2019), (28) Toscano-Gadea and González (2020), (29) Eason (1969), (30) Rovner (1971),
(31) Schoenberg (2021), (32) Brown (2006), (33) Costa (1979), (34) Aisenberg and Costa (2005), (35) Aisenberg, Estramil, González, Toscano-Gadea, and Costa (2008), (36) Hebets
per. Com. (https://www.youtube.com/watch?v¼Z6UXTq9X004), (37) Nappi (1965), (38) Rypstra and Wilder (2007), (39) Engelhardt (1964), (40) Alvarado-Castro and Jimenez
(2011), (41) Domı́nguez and Jimenez (2005), (42) Sentenská, Šedo, and Pekár (2020), (43) Abregú, Simian, Mattoni, and Peretti (2022), (44) Abregú & Peretti pers. obs.,
(45) Preston-Mafham (1999), (46) Bruce and Carico (1988), (47) Montgomery (1903), (48) Catley (1993), (49) Whitehouse and Jackson (1994), (50) Knoflach (2000), (51) McCrone
and Levi (1964), (52) Knoflach & Van Harter (2002), (53) Snow and Andrade (2004), (54) Scott, Vibert, and Gries (2012), (55) Vibert, Scott, and Gries (2016), (56) Breene and Sweet
(1985), (57) Kaston (1970), (58) Shulov (1940), (59) Berendonck (2003), (60) Knoflach and Benjamin (2003), (61) Frank & Buzatto per. Com., (62) Coyle (1986), (63) Coyle &
O’shields (1990), (64) Costa & Perez-Miles (1998), (65) Coyle (1985), (66) Copperi (2018), (67) Ferretti, Pompozzi, and Perez-Miles (2011), (68) Capocasale and Perez-Miles (1990),
(69) Ferretti, Pompozzi, Copperi, Perez-Miles, and González (2012), (70) Petrunkevitch (1911), (71) Ferretti, Pompozzi, Copperi, González, and Perez-Miles (2013), (72) Ferretti and
Ferrero (2008), (73) Bertani, Fukushima, and da Silva Júnior (2008), (74) Ferretti, Copperi, Schwerdt, and Pompozzi (2015).
Table 3 Reports of female mating quiescence in the order Solifugae. Columns summarize information extracted from each paper (NA ¼ information non-
available).
Mating
phase Female
when Female body position during movement Terms used /
Sexual quiescence Male behavior before female quiescence (Quiescence during Quiescence Functional
Species cannibalism SSD occurs quiescence Level) quiescence duration hypothesis Source
Ammotrechidae
Titanopuga Yes, but Yes Pre + Cop Tapping. Abdomen in vertical None. Females remain in L / None 1
ARTICLE IN PRESS
salinarum apparently a (F > M) +Post position (III). quiescence during
byproduct the entire mating;
of mating lasts under
confinement 4 min.
Eremobatidae
Eremobates Yes Yes Pre + Cop Male advances with elevated Male positions the female Yes3. 2–4 min before Q1 / 2
durangonus (F > M) +Post2 pedipalps and first pair of upward or sideways. release Intersexual
legs. Male chelicerae Abdomen bent upwards communication
extended. He performs (III).
rocking motion. After
stroking over the female, the
female enters in a state of
submission.
Eremobates Yes Yes Pre + Cop Same as E. durangonus. Same as E. durangonus (III). Yes3. NA. S / None 3
marathoni (F > M) +Post2
Eremobates Yes Yes Pre + Cop Same as E. durangonus. Same as E. durangonus (III). Yes3. 2–4 min before Q1 / 2
nodularis (F > M) +Post2 release Intersexual
communication
Continued
Mating
phase Female
3 1
Eremobates No Yes Pre + Cop Same as E. durangonus. Body lowered towards the Yes . 3 min 51 s (mean) Q / None 4, 5
pallipes (F > M) +Post2 substrate, pedipalps,
ARTICLE IN PRESS
chelicerae and legs relaxed,
body bended (peltidium
over abdomen). Abdomen
bent upwards (III).
Eremobates Yes Yes Pre + Cop Same as E. durangonus. Same as E. durangonus (III). Yes3. 2–4 min before Q1 / 6, 2
palpisetulosus (F > M) +Post2 release Intersexual
communication
Galeodidae
Galeodes Yes Yes Pre + Cop Male courts his mate by Lifted up abdomen (III). None. Females remain in L / None 7
arabs (F > M) +Post2 stroking her, then the female quiescence
enters in a state of lethargy. during the entire
mating; mating
lasts under 5 min.
Galeodes Yes Yes Pre + Cop Male advances forward with Abdomen bent dorsally. None. The entire mating Im, PS, L, H, 8–14
caspius (F > M) +Post2 pedipalps raised. The male Legs contracted against the last approximately W, NR, HL,
subfuscus* strikes the female with its body (III). 3 min and the D, P / Female
pedipalps producing her female regains sexual
paralysis. If the female conscious after the aggressiveness
becomes aggressive, the mating is finished and cannibalism
male touches her
propeltidium with the
suction organs and bites the
lateral part of the female’s
ARTICLE IN PRESS
propeltidium and chews it.
In this movement, he also
clasps the female pedipalps.
After the bite, the female
becomes immobile.
Galeodes No Yes Pre + Cop Male strokes the female. Abdomen in vertical None. NA. P / None 10
granti (F > M) +Post2 The male merely touches position (III).
the female with his pedipalps
in reply to which she lifts her
abdomen and allows him to
grasp her with his chelicerae.
Galeodes sp. Yes Yes Pre + Cop Male approaches the female. Lifted up abdomen (III). None. NA. L / Intersexual 13
(F > M) +Post2 The male gets responses communication
from the females as he
continues stroking her with
his pedipals. At that time the
female lifts her abdomen for
copulation.
Continued
Mating
phase Female
Galeodes No Yes Pre + Cop Male approaches the female In lateral position, with the None. NA. P / None 3, 15
sulfuripes (F > M) +Post2 with pedipalps lifted, the prosoma bent over the
ARTICLE IN PRESS
male raises and lowers his opisthosoma at a 90° angle,
pedipalps several times. legs flexed against the body.
Suddenly the male touches Abdomen in vertical
the pedipalps of the female position (III).
with his pedipalps. She falls
into paralysis. As if receiving
an electrical shock, she falls
into a convulsion and her
whole body appears to be
paralyzed.
Othoes Yes Yes Pre + Cop Male grabs the female using Laterally, with the prosoma None. NA F, Im, S / 16, 17
saharae* (F > M) +Post2 his pedipalps and chelicerae. bent over the opisthosoma at Female sexual
a 90° angle, legs flexed aggressiveness
against the body. Abdomen and cannibalism
over prosoma (III).
Solpugidae
Metasolpuga Yes, but Yes Pre + Cop Male grabs the posterior end Legs against the body, palps None. 5–6 min Q, T, C / 18
picta apparently a (F > M) +Post2 of the female opisthosoma against chelicerae Abdomen Intersexual
byproduct with his chelicerae while in vertical position (III). communication
of touching the female
confinement prosoma and chelicerae with
his pedipalps.
Species marked with an asterisk indicate papers that provide detailed behavioral descriptions of female quiescence. SSD ¼ Sexual size dimorphism (F ¼ female; M ¼ Male). Mating phases: Pre ¼ precopulatory,
ARTICLE IN PRESS
Cop ¼ copulatory, Post ¼ postcopulatory. Quiescence levels are presented in Table 6. Terms used to describe female mating quiescence in the original papers are: C ¼ Catatony, D ¼ Defenseless state, F ¼ Freezing,
Im ¼ Immobility, H ¼ Hypnosis, HL ¼ Hypnosis-Like, L ¼ Lethargy, NR ¼ No Response, P ¼ Paralysis, PS ¼ Passivity, Q ¼ Quiescence, S ¼ Submission, T ¼ Torpor, W ¼ Willnessless. The names of the functional
hypotheses are the same as presented in Box 1 (CFC ¼ cryptic female choice). Uppercase notes: (1) Eremobates females may submit or accept a male without prolonged quiescence; one was even observed to feed
on termites while being mated. The author uses submission when describing behavior (Muma, 1966a). For E. palpisetulosus, Punzo (1997) uses lethargic citing Heymons (1902). (2) After the attack phase, throughout
the copulatory stage, and after the post transfer massages. (3) If the female is not completely quiescent, she can struggle or fight. During quiescence females may walk or even prey (Punzo, 1998b). One female was observed
feeding on a termite during copulation (Muma, 1966a).
(1) Peretti pers. obs., (2) Muma (1966a), (3) Punzo (1998b), (4) Muma and Muma (1988), (5) Rowsell and Cushing (2020), (6) Punzo (1997), (7) Cloudsley-Thompson (1961), (8) Heymons (1902), (9) Berland (1932),
(10) Cloudsley-Thompson (1967), (11) Cloudsley-Thompson (1977), (12) Hrušková-Martišová, Pekár, and Gromov (2008), (13) Pandram and Sharma (2018), (14) Hrušková-Martišová, Pekár, and Bilde (2010),
(15) Amitai, Levy, and Shulov (1962), (16) Junqua (1962), (17) Junqua (1966), (18) Wharton (1987).
Table 4 Reports of female mating quiescence in the order Scorpiones. Reports were classified as follows: (A) Immobility with postural change during courtship
initiation stage, and (B) Immobility without postural change during sexual sting behavior.
Female body
Mating Female body position during Female
phase when Male behavior Male behavior position quiescence movement Terms used /
Sexual quiescence before female during female before (Quiescence during Quiescence Functional
Species cannibalism SSD occurs quiescence quiescence quiescence Level) quiescence duration hypothesis Source
(A) Immobility with postural change: female motionless during courtship initiation stage
Bothriuridae
Bothriurus No Absent Pre Grasping, Reorientation, None Totally docile, NA Several minutes SP, P / 1–4
bonariensis* or slight vibratory vibratory (receptive legs relaxed, Intersexual
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(F > M) behaviors. behaviors, females) / pedipalps communication,
cheliceral Sting rubbing backwards, Postcopulatory
movements. (intermediate metasoma sexual selection
receptive extended (CFC)
females). backwards (III).
Bothriurus No Absent Pre Grasping. Reorientation. NA Pedipalps next to NA NA SP / Intersexual 5
buecherli or slight the body, communication,
(F > M) metasoma Postcopulatory
supported on the sexual selection
mesosoma, legs (CFC)
withdrawn
towards the body
(III).
Bothriurus No Absent Pre Grasping, Reorientation, None Immobile and NA NA SP / Intersexual 4
burmeisteri* or slight vibratory vibratory (receptive relaxed body communication,
(F > M) behaviors. behaviors. females) / (III). Postcopulatory
Sting rubbing sexual selection
(intermediate (CFC)
receptive
females).
Bothriurus No Absent Pre Grasping, Reorientation, None Totally docile, NA NA SP / Intersexual 2, 4
chacoensis* or slight vibratory vibratory (receptive legs relaxed, communication,
(F > M) behaviors. behaviors, females) / pedipalps, Postcopulatory
cheliceral Sting rubbing backwards sexual selection
movements. (intermediate metasoma (CFC)
receptive extended
Bothriurus No Absent Pre Grasping, Reorientation, None Immobile and NA NA SP / Intersexual 4, 6
cordubensis* or slight metasoma vibratory (receptive relaxed body communication,
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(F > M) shaking. behaviors. females) / (III). Postcopulatory
(intermediate (CFC)
receptive
females)
Bothriurus No Absent Pre Grasping, Reorientation, None Totally docile, Metasoma NA SP / Intersexual 2, 4
flavidus* or slight metasoma metasoma (receptive pedipalps erect, communication,
(F > M) shaking. shaking. females) / backwards, legs pulsing. Postcopulatory
Sting rubbing relaxed, elevated sexual selection
(intermediate metasoma (III). (CFC)
receptive
females).
Bothriurus noa* No Absent Pre Grasping, Reorientation, None Immobile and NA NA SP / Intersexual 4, 6
or slight metasoma vibratory (receptive relaxed body communication,
(F > M) shaking. behaviors. females) / (III). Postcopulatory
(intermediate (CFC)
receptive
females).
Continued
initiation stage, and (B) Immobility without postural change during sexual sting behavior.—cont’d
Female body
Bothriurus No Absent Pre Grasping, Reorientation, None Totally docile, NA NA SP / Intersexual 2, 4, 6

prospicus* or slight vibratory metasoma (receptive legs relaxed, communication,
(F > M) behaviors, shaking. females) / pedipalps Postcopulatory
metasoma Sting rubbing backwards, sexual selection
shaking. (intermediate metasoma (CFC)
receptive extended
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Brachistosternus No Absent Pre Grasping, Reorientation, None Totally docile, NA NA SP / Intersexual 2, 4
ferrugineus* or slight intense vibratory (receptive legs relaxed, communication,
(F > M) metasoma behaviors, females) / pedipalps, Postcopulatory
shaking. cheliceral Sting rubbing backwards sexual selection
movements. (intermediate metasoma (CFC)
receptive extended
Brachistosternus No Absent Pre Grasping, Reorientation, None Immobile and NA NA SP / Intersexual 4
pentheri* or slight vibratory vibratory (receptive relaxed body communication,
(intermediate (CFC)
receptive
females).
Timogenes No Absent Pre Grasping, Reorientation, None Immobile and NA NA SP / Intersexual 4
dorbignyi* or slight vibratory vibratory (receptive relaxed body communication,
(intermediate (CFC)
receptive
females).
Timogenes No Absent Pre Stridulation, Reorientation, None Docile, pedipalps NA NA SP / Intersexual 2, 4
elegans* or slight grasping, vibratory (receptive backwards, legs communication,
(F > M) metasoma behaviors, females) / relaxed, Postcopulatory
shaking. cheliceral Sting rubbing metasoma sexual selection
movements. (intermediate extended (CFC)
receptive backwards,
females). chelicerae more
or less extended
(III).
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Urophonius No Absent Pre Grasping, Reorientation, None Immobile, Minimal leg 2–60 min M / Intersexual 7
achalensis or slight vibratory vibratory (receptive pedipalps and (mean ¼ 16 min) communication,
(F > M) behaviors, behaviors, females) / backwards, legs pedipalp Postcopulatory
metasoma metasoma Sting rubbing relaxed, movements. sexual selection
touching. touching. (intermediate metasoma (CFC)
receptive extended
Urophonius No Absent Pre Grasping, Reorientation, None Immobile, Minimal leg 4–20 min M, SP, Qt / 2, 4,
brachycentrus* or slight vibratory vibratory (receptive pedipalps and (mean ¼ 10 min) Intersexual 7, 8
(F > M) behaviors, behaviors, females) / backwards, legs pedipalp communication,
metasoma metasoma Sting rubbing relaxed, movements. Postcopulatory
touching. touching, (intermediate metasoma sexual selection
cheliceral receptive extended (CFC)
movements. females). backwards (III).
Urophonius No Absent Pre Grasping, Reorientation, Not reported. Totally docile, NA NA SP / Intersexual 2
iheriingi* or slight vibratory vibratory legs relaxed, communication,
(F > M) behaviors. behaviors, pedipalps Postcopulatory
metasoma backwards sexual selection
shaking. metasoma (CFC)
extended
backwards (III).
Continued
Female body
Buthidae
Centruroides NA Yes Pre Vibratory Reorientation, None. NA (?). NA NA M / None 12
margaritatus (F > M) behaviors. vibratory
behaviors.
Leiurus No Absent Pre Grasping, Reorientation, Clubbing, Relaxed (but not Minimal leg At least 1 min 33 s PQ / None 9
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quinquestriatus or slight vibratory vibratory pushes. completely), and (see source)
(F > M) behaviors. behaviors, leg pedipalps next to pedipalp
4th grasping. the body, movements.
metasoma
supported
completely on
the mesosoma
(III).
Parabuthus No Absent Pre Grasping. Reorientation. Not reported. NA (?). NA NA P / Intersexual 13
planicauda or slight communication
(F > M)
Rhopalurus rochai No Yes Pre NA Walks over the Not reported. NA (?). NA NA Q / None 14
(F > M) back of the
quiescent female
(over-ride).
Tityus NA Yes Pre Grasping, Reorientation. Contact with Immobile with None. NA P, I / Intersexual 10
magnimanus (F > M) pedipalp grip, male. body withdraws communication
vibratory against their own
behaviors body (III).
(with
non-receptive
females).
Zabius fuscus No Yes Pre Grasping, Reorientation. None Relaxed, but not Female NA PQ, SP / 2, 4
(F > M) vibratory (receptive completely (III). almost Intersexual
behaviors, females) / motionless. communication,
male probing. Sting rubbing Postcopulatory
(intermediate sexual selection
receptive (CFC)
females).
Diplocentridae
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Nebo No Absent Pre Grasping, Reorientation. Not reported. NA (?). NA NA P / None 15
hierochonticus or slight vibratory
(F > M) behaviors.
Iuridae
Hadrurus No Absent Pre Juddering, Reorientation, NA Immobile with Not 1–10 min (3 h P, I / None 11
arizonensis or slight aggressive vibratory body withdraws reported. exception).
(F > M) grasping, and behaviors. against their own
lunges. body (III).
Scorpionidae
Opisthophthalmus No NA Pre Grasping, Reorientation. NA NA (?). Female NA SP, M / 16
latimanus vibratory almost Intersexual
behaviors. motionless communication
(III).
(B) Immobility without postural change: female motionless during sexual sting
Bothriuridae
Bothriurus No Absent Pre Sexual sting. Sexual sting. NA Immobile with NA 1–10 min M / Intersexual 17
bonariensis or slight no apparent communication
(F > M) resistance (I).
Continued
Female body
Urophonius No Absent Pre Sexual sting Sexual sting. None / Immobile with Minimal leg 2–20 min SSQ / 7
achalensis or slight (1–6 times). female no apparent movements. (7.5 min) Intersexual
(F > M) resistance. resistance, communication,
metasoma Postcopulatory
mesosoma (I). (CFC)
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Urophonius No Absent Pre Sexual sting Sexual sting. None / Immobile with Minimal leg 7–30 min SSQ / 7
brachycentrus or slight (1–4 times). female no apparent movements. (mean ¼ 15.5 min) Intersexual
(F > M) resistance. resistance, communication,
metasoma Postcopulatory
mesosoma (I). (CFC)
Euscorpiidae
Euscorpius No No Pre Sexual sting Sexual sting, may Not reported Both sexes still NA 13.2 3.0 min St / None 18
flavicaudis (0–10 times). twist sting in the (I).
wound
Megacormus NA Absent Pre Sexual sting, Sexual sting, Not reported. Passive with no None. 4–50 min P / Intersexual 19, 20
gertschi or slight pedipalp grip reorientation, observable (mean ¼ 6.8 min). communication,
(F > M) (up to 4 times). vibratory reaction. Postcopulatory
behaviors, Metasoma curled sexual selection
cheliceral to the left and (CFC)
movements, resting on the
ventral substrate (I).
stimulation by
the male’s first
pair of legs.
Scorpiops luridus NA Absent Pre Grasping, Sexual sting. Not reported. Passive/ NA 2–5 min P, M / 21
or slight sexual sting. motionless Intersexual
(F > M) female (I). communication,
Postcopulatory
sexual selection
(CFC)
Iuriidae
Hadrurus No Absent Pre Sexual sting Sexual sting, None. Passive/ NA 1–35 min P / None 11
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arizonensis or slight (1–14 times). vibratory motionless
(F > M) behaviors. female (I).
Columns summarize information extracted from each paper (NA ¼ information non-available). Species marked with an asterisk indicate papers that provide detailed behavioral description of female quiescence. Quiescence duration in immobility without
postural change ¼ sexual sting duration (sexual sting occurs in other species, but only those in which immobility during this behavior has been described are cited here). SSD ¼ Sexual size dimorphism (F ¼ female; M ¼ Male). Mating phases:
Pre ¼ precopulatory. Quiescence levels are presented in Table 6. Terms used to describe female mating quiescence in the original papers are: I ¼ Immobile, M ¼ Motionless, P ¼ Passive, PQ ¼ Partial female Quietude, Q ¼ Quiescent, Qt ¼ Quietude,
SP ¼ Submission Posture (or submissive female), St ¼ Stillness, SSQ ¼ Sexual Sting female Quiescence. The names of the functional hypotheses are the same as presented in Box 1 (CFC ¼ Cryptic female choice). 1Attack or cannibalism is frequent in
non-receptive females in a non-sexual context (Peretti et al., 1999).
(1) Covelo de Zolessi (1956), (2) Peretti (1993), (3) Peretti (1995), (4) Peretti (2001), (5) Toscano-Gadea (2010), (6) Peretti, Acosta, and Martinez (2000), (7) Oviedo-Diego, M.O. personal obs., (8) Maury (1968), (9) https://www.youtube.com/watch?
v¼yfMMV4S-4t4 (4:24 min, estimated time of the beginning of quiescence), (10) Ross (2009), (11) Tallarovic, Melville, and Brownell (2000), (12) Briceño and Bonilla (2009), (13) Alexander (1959), (14) Chantall-Rocha and Japyassú (2017), (15) Rosin and
Shulov (1963), (16) Alexander (1957), (17) Olivero, Vrech, Oviedo-Diego, Mattoni, and Peretti (2019), (18) Benton (1993), (19) Francke (1979), (20) Olguı́n-Perez, Francke, and Carbajal-Saucedo (2021), (21) Jiao and Zhu (2010).
Table 5 Reports of female mating quiescence in the orders Opiliones (harvestmen), Amblypygi (whip spiders), and Pseudoscorpiones.
when Male-female behavior position during movement Terms used /
Sexual quiescence before female quiescence during Quiescence Functional
Species cannibalism SSD occurs quiescence (Quiescence Level) quiescence duration hypothesis Source
OPILIONES: Gonyleptidae
Iporangaia No Yes Post After copulation, the Prostrated (II). None. Until 5 h Q / None 1
pustulosa (F > M) male remains with his
second pair of legs
extended frontward,
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touching female’s legs
(mate guarding).
Neosadocus No Yes Cop Female helps the male In copulatory Pulling back 2 min Qt / None 2
bufo (F < M) to achieve the position (face-to- and hitting
copulatory position. face), female slightly the male
inclines up so that with the legs
the male can access II after the
the gonopore (I). initial
contact.
Pachyloides No No Cop In mating position, In copulatory Moving 654 s Im / None 3
thorellii male raises his third position (face-to- their legs II
pair of legs until he face), female slightly slowly II
touches female’s inclines up so that before
second pair of legs. the male can access separation.
Then male inserts the the gonopore (I).
penis.
OPILIONES: Nomoclastidae
Quindina No Yes Pre Female touches male In front of the male None. 1–20 min Q / None 4
albomarginis (F > M) with her second pair of (I).
legs
OPILIONES: Phalangiidae
Phalangium No Yes Cop Male rubs the distal In copulatory Motionless NA Q/ 5
opilio (F > M) portion of the penis position (face-to- or bouncing Communication
between the face), female slightly movements.
latero-posterior lobes inclines up so that
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of female’s mouth. the male can access
the gonopore (I).
OPILIONES: Sclerosomatidae
Leiobunum No Yes Cop Mutual pedipalp In copulatory NA NA Q / None 6
vittatum (F > M) drumming and position (face-to-
cheliceral grab while face), female slightly
the male copulates. inclines up so that
the male can access
the gonopore (I).
AMPBLYPYPI: Phrynichidae
Damon No NA Pre Male touches the The female NA NA M/ 7
variegatus female’s body with the remained crouched, Communication
first pair of legs and allowing the male to
threat her extending push with their
his pedipalps. pedipalps (III).
Receptive females
respond with leg
touching.
Continued
Table 5 Reports of female mating quiescence in the orders Opiliones (harvestmen), Amblypygi (whip spiders), and Pseudoscorpiones.—cont’d
when Male-female behavior position during movement Terms used /
Sexual quiescence before female quiescence during Quiescence Functional
Species cannibalism SSD occurs quiescence (Quiescence Level) quiescence duration hypothesis Source
Phrynus No NA Pre Same as previous. Same as previous NA NA M/ 7*
barbadensis (III). Communication
PSEUDOSCORPIONES: Cheliferidae
Dactylochelifer No NA Cop The female shows Standing immobile NA 20 min M / None 8
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latreillii resistance. in front of the male
latreillii (I).
PSEUDOSCORPIONES: Chernetidae
Dendochernes No NA NA NA NA (?). NA NA C / None 9
cyrneus
Columns summarize information extracted from each paper (NA ¼ information non-available). SSD ¼ Sexual size dimorphism (F ¼ female; M ¼ Male). Mating phases: Pre ¼ precopulatory,
Cop ¼ copulatory, Post ¼ postcopulatory. Quiescence levels are presented in Table 6. Terms used to describe female mating quiescence in the original papers are: C ¼ Catalepsy, I ¼ Immob-
ile, M ¼ Motionless, Q ¼ Quiescence Qt ¼ Quiet. The names of the functional hypotheses are the same as presented in Box 1.
(1) Requena and Machado (2014), (2) Willemart, Osses, Chelini, Macı́as-Ordóñez, and Machado (2009), (3) Stanley, Francescoli, and Toscano-Gadea (2016), (4) Mora (1990), (5) Willemart et al.
(2006), (6) Fowler-Finn, Triana, and Miller (2014), (7) Alexander (1962), (7*) Alexander (1962) under Admetus barbadensis, (8) Kirchmair and Raspotnig (2021), (9) Cuthbertson (1984).
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Fig. 3 (A) Terms used in the literature to refer to female quiescence depending on the
arachnid order, with their frequency indicated by horizontal bars. (B) Number of articles
in which any of the terms used to refer to female quiescence was employed, classified
according to the arachnid order. (C) Number of articles in which any of the terms used to
refer to female quiescence were employed, classified according to the arachnid order
and the decade in which they were published.
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2. Literature search
We conducted an extensive literature review of quiescence in arach-
nids using the Scopus, Google Scholar, and ResearchGate databases. The
following terms were searched for in titles, keywords, or abstracts of articles
published until June 2022: “arachnid + sexual + catalepsy,” “arachnid +
sexual + quiescence,” “arachnid + sexual + stillness,” and “arachnid +
sexual + motionless,” “arachnid + sexual + immobility,” “arachnid + sexual
+ inactivity,” and “arachnid + sexual + passive”. We also replaced the word
“arachnid” with the name of each arachnid order or with the common name
of relevant taxa (e.g., spiders, scorpions, harvestmen). During our review,
we identified a host of terms used for female mating quiescence, such as
lethargy, torpor, catatonic state, akinesis, hypnosis, trance state, paralysis,
submission, and quiet. Although these terms were not employed in the ini-
tial survey, they were considered in the results (see below). For our analysis,
we limited our review to articles written in English and Spanish. Articles in
other languages were considered if they were cited in publications included
in our search. In those cases, they were translated with free translation
websites (Google Translator or DeepL Translator), so descriptions of some
behaviors may be imprecise. Despite the great diversity of terms used in the
literature, we will use quiescence throughout the text because we argue that
this is the only term that explicitly contemplates the possibility of voluntary
decisions of the female, even when she remains immobile (see Section 3).
We extracted the following information from each article: terminology
employed by the author, male and female behaviors performed immediately
before immobility, female quiescence duration, phases in which it occurs
(i.e., precopulatory, copulatory, or postcopulatory), occurrence of female
postural changes or movements during quiescence, reports of sexual canni-
balism (in the article and the species in general), sexual size dimorphism, and
functional hypothesis tested or proposed by the author(s) to explain female
quiescence (Box 1), which we then discussed critically. Other information
was extracted only for specific taxa, such as the type of web and genital com-
plexity in spiders.
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BOX 1 Contexts that may have favored the evolution

of female quiescence in arachnids.
Female sexual aggressiveness and cannibalism: female mating quiescence is a
behavior induced by the males to prevent or decrease the chances of being
attacked and/or cannibalism during sexual interactions. This hypothesis has been
originally proposed for spiders (e.g., Fraser, 1987; Kaston, 1970; Shulov, 1940) but
also applies to solifuges, which exhibit sexual cannibalism. An assumption of this
hypothesis is that female mating quiescence is under male’s control.
Female and male genital complexity: female mating quiescence is a behav-
ior that allows the correct insertion of morphologically complex genital struc-
tures. This hypothesis has been originally proposed for spiders (Gering, 1953)
but may also apply to other groups with direct sperm transfer like solifuges.
An assumption of this hypothesis is that female mating quiescence is under
female’s control.
Sperm transfer: female mating quiescence is a behavior that allows sperm
transfer. This hypothesis has been originally proposed for spiders (e.g., Snow &
Andrade, 2004), but may also apply to solifuges or harvestmen. An assumption
of this hypothesis is that female mating quiescence is under female’s control.
Copulation substrate constraints: female mating quiescence is a behavior
that can only occur in those taxa in which individuals’ mate on a firm substrate,
such as non-web builder spiders, solifuges, scorpions, harvestmen, whip spiders,
and pseudoscorpions.
Intersexual communication and mate choice: female mating quiescence is
a behavior that allows the female to communicate, for example, her sexual recep-
tivity after mate choice. An assumption of this hypothesis is that female mating
quiescence is under female’s control.
Postcopulatory sexual selection and female quiescence: female mating
quiescence is a behavior that may evolve in a postcopulatory sexual selection
context by cryptic female choice or sperm competition. An assumption of this
hypothesis is that female mating quiescence is under female’s control.
3. Terminologies, history, and other concerns

Heymons (1902) penned a pioneering report of female immobility
during mating in arachnids. The study species was the solifuge Galeodes
caspius subfuscus, in which a drastic female change in posture is seen during
the precopulatory phase: the abdomen is bent dorsally against the cephalo-
thorax, and the legs remain contracted and motionless. The terms used for
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this female behavior by Heymons (1902) were “passive state,” “will-less

state,” “lethargy,” “hypnosis,” and “mindless state.” However, several prob-
lems can be noted in this article and others that came after. First, many
authors have made anthropocentric interpretations of female mating quies-
cence, using terms with medical and psychological implications or that refer
to physical or physiological states that have not been actually evaluated but
assumed subjectively. Such terms include “hypnosis,” “torpor-like state,”
“catalepsy,” “lethargy,” and “paralysis” (e.g., Amitai et al., 1962; Bennett,
1987; Coyle, 1985; Heymons, 1902; Pandram & Sharma, 2018; Rovner,
1971; Wharton, 1987).
A second common problem plaguing many past studies is a gender-biased
implication that males impose or force females into mating quiescence. For
example, in the case of G. caspius subfuscus mentioned above, it seems that
males catching the female is a necessary stimulus for female quiescence.
However, Heymons (1902) himself obtained quiescence in females when
he imitated the male movement with fine tweezers, at least in “young, mature,
freshly caught females,” and not with “older, fertilized ones.” In fact, older,
fertilized females tried to free themselves by “violent” movements.
Heymons’s (1902) interpretation of this difference was that the development
of “sexual glands” (perhaps he was referring to gonads) is a precondition to
female immobility. In these descriptions there is no consideration of female
autonomy, nor of mate choice, sexual cooperation, or inter-sexual commu-
nication. Certainly, these interpretations mirror attitudes in the society of that
time. Although Darwin had already postulated the concept of mate choice in
1871, the theory of sexual selection, especially his idea of female choice, was
generally ignored for about 60 years until Fisher (1930) took it up again.
The notion that males control female mating quiescence in arachnids is,
however, challenged by key anecdotes and details in publications. For exam-
ple, it has been reported that females can perform different movements during
their apparent state of “immobility,” including “subtle” bouncing or slow leg
and pedipalp movements in spiders (Berland, 1932), solifuges
(Cloudsley-Thompson, 1967, 1977; Hrušková-Martišová, Pekár, & Bilde,
2010; Hrušková-Martišová, Pekár, & Gromov, 2007), and harvestmen
(Stanley et al., 2016; Willemart et al., 2006, 2009). Also, females that are oth-
erwise immobile can still conduct movements that expose her genitalia, such
as swiveling the abdomen or protruding and distending it in wolf spiders
(Lycosidae) and tarantulas (Mygalomorphae) (Coyle & O’Shields, 1990;
Engelhardt, 1964; Just et al., 2018; Rovner, 1971). Even more, females in
a state of immobility can “wake up” and even walk with the male still in
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the mating position, as in some wolf spiders ( Just et al., 2018; Piacentini et al.,
2017) and solifuges (Peretti, Vrech, & Hebets, 2021), or interrupt mating and
abandon the male, as in some cobweb spiders (Theriidae) and solifuges
( Junqua, 1962, 1966; Shulov, 1940). These examples clearly challenge the
commonly held assumption in the literature that females are completely at
the mercy of males with regard to quiescence.
Beyond anthropocentric and gender-biased interpretations of female
mating quiescence, a third problem is that female quiescence is often men-
tioned only briefly, and in terms that are overly vague and non-descript,
and that differ markedly from study to study. In some cases, quiescence is pres-
ented as females just remaining immobile in the position they had adopted
before courtship, as reported for some spiders and a pseudoscorpion, yet in
other cases constituting a drastic change in female posture, as reported in other
spiders as well as in scorpions and some solifuges (Tables 1-5).
A fourth problem concerns all the terms used to refer to female mating
quiescence, not just among different orders but even within the same ones.
The diversity of terms used to refer to female quiescence in arachnids is vast
(Fig. 3A). The most considerable diversity of terms occurs in spiders, in
which 11 terms refer to any type of female immobility during mating
(Table 2). Linked to this last problem, we emphasize that multiple terms
for quiescence were sometimes used as synonyms, even when they are
not. The fifth and final problem is linked to the two previous problems.
Because descriptions of female quiescence behavior are often poor in details,
and because there is no consensus on the terminology used, it is difficult to
establish homologies that would enable a detailed analysis of these phenom-
ena in a phylogenetic context.
We thus below propose a single universal term, with the goal of producing
a precise and objective description of female quiescence. In this pursuit, it is
essential to consider that when naming a behavior, we can unintentionally
formulate a hypothesis, for example, by selecting words that reveal or foster
misinterpretations of the term or a bias in its use. Accordingly, we note the
need to describe behaviors in detail and objectively, without invoking untested
assumptions. An interesting illustration of this is when researchers refer to
thanatosis, a behavior that also implies immobility, but which occurs in survival
rather than in sexual contexts. The simple and parsimonious assumption that
individual animals control their own thanatosis is never questioned, nor is
it supposed that it is under the control of other animals ( Jones et al., 2011;
Machado & Pomini, 2008; Pereira, Elpino-Campos, Del-Claro, &
Machado, 2004). In fact, evidence in flour beetles, Tribolium castaneum, shows
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that thanatosis duration is not only under individual control but is also heritable
(Miyatake et al., 2004). Moreover, when males of the gift-giving spider Pisaura
mirabilis perform thanatosis to avoid being cannibalized during mating, it is
described as a “male strategy,” even though it occurs when both males and
females are handling the gift (Bilde, Tuni, Elsayed, Pekár, & Toft, 2006;
Hansen, Gonzales, Toft, & Bilde, 2008). Thus, unless rigorous observations
or experimentation prove the contrary, female immobility should be consid-
ered by default as a behavior directly under female control.
4. General results of the survey

Our survey flagged 120 articles containing at least one combination of
searched terms: 70 papers with spiders, 19 with scorpions, 21 with solifuges,
six with harvestmen, and four with other arachnid orders (Figs. 3B-C). These
data showed no consistency in the use of the terms even within given taxa.
For example, for the wolf spiders of the family Lycosidae, five terms were used
to describe the same phenomenon (“quiescence,” “catalepsy,” “motionless,”
“inactivity,” and “akinesis”). Moreover, different authors used different terms
for the same species (e.g., Rovner, 1971; Schoenberg, 2021 for Rabidosa
rabida; Costa, 1979 and Aisenberg & Costa, 2005 for Schizocosa malitiosa).
After compiling the available information, we defined and classified female
quiescence as occurring at different levels, according to the degree of females’
body postural change, and the mating phase when it takes place (Table 6).
Table 6 Definitions of the operational levels of female mating quiescence.

Quiescence
level Definition Occurrence
Level I No postural change. The female may be Precopulatory and
quiescent during the precopulatory phase and copulatory phases
remain in this state during sperm transfer. The
female moves immediately after copulation.
Level II No postural change. The female remains Copulatory and/or
quiescent, even after the sexual interaction has postcopulatory
concluded. phases
Level III Evident postural change. It may include Precopulatory,
modification in the initial position of the body copulatory and
and/or appendages (flexion or extension) or body postcopulatory
relaxation. Quiescence may begin in the phases
precopulatory phase and remain after the sexual
interaction.
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In the following sections, we compile different aspects of the biology of each

arachnid order, along with the occurrence of female quiescence, and group
them according to hypotheses proposed by the authors of the articles reviewed
(Tables 1–5). Finally, we discuss the generality of these hypotheses among the
arachnid orders sampled here.
5. Quiescence and mating phase

Our analysis shows that spiders are the taxon with the most reports of
species exhibiting female quiescence. It has been reported in 84 species, dis-
tributed in 18 families (Tables 1–2). In 42 species, quiescence occurs in only
one phase: 7.1% during the precopulatory phase (n ¼ 3), 69% during the
copulatory phase (n ¼ 29), and 22.7% during the postcopulatory phase
(n ¼ 10). Quiescence may also occur during more than one phase in spiders,
and it has been reported in 40 species. In 35% of the species (n ¼ 14), female
quiescence begins during the precopulatory phase and continues during the
copulatory phase. In 55% of the species (n ¼ 22), female quiescence starts
during the copulation phase and continues during the postcopulatory phase.
Finally, in 10% of the species (n ¼ 4), female quiescence begins during the
precopulatory phase and ends during the postcopulatory phase. In two spe-
cies of 84 (2.4%), the phase in which quiescence occurred was not specified.
In 12 of 13 species of solifuges, female quiescence begins after the initial
phase of the encounter between sexes (precopulatory phase or “attack
phase,” following Muma, 1966a). Once the male approaches the female
and touches her with his pedipalps or even grasps her with his chelicerae,
the female becomes quiescent. Females start moving again only after the
sperm transfer massage phase (Peretti et al., 2021).
Scorpions, in contrast, are characterized by the occurrence of quiescence
only at the precopulatory phase. It occurs at the beginning of courtship,
when the male touches the female, performs body vibrations called
juddering (Olivero et al., 2017; Polis & Sissom, 1990), or even sting the female
with his telson (stinging behavior, Polis & Sissom, 1990; Peretti, 2014,
Table 4, Fig. 2F). The occurrence of female quiescence in the precopulatory
phase makes sense: females need to move at the beginning of the locomotion
phase (also known as mating dance or promenade à deux) when males search for
an appropriate place to deposit the spermatophore on the substrate. Then, the
female places herself on the spermatophore to take the sperm (Polis & Sissom,
1990). It is important to note that female quiescence only occurs when the
female is sexually receptive (Peretti, 2001); otherwise, she does not adopt this
posture, and male and female may move away in opposite directions.
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In harvestmen, female quiescence has been described at the pre-

copulatory, copulatory, and postcopulatory phases. However, most of the
observed cases of quiescence (66.7%) occur during copulation (n ¼ 4 of 6).
There is only one species with precopulatory quiescence: Quindina
albomarginis, a species with non-traditional sex roles in which males build a
mud nest where females lay their eggs and leave them under paternal protec-
tion (Mora, 1990). Iporangaia pustulosa (Fig. 2C), another species with exclu-
sive paternal care, is the only species we found with postcopulatory
quiescence (Requena & Machado, 2014).
For whip spiders, female quiescence has been described in Damon vari-
egatus and Phrynus barbadensis in a precopulatory context (Alexander,
1961, 1962). In pseudoscorpions, female quiescence has also been reported
for only two species. In Dactylochelifer latreillii latreilliid, the female remains qui-
escent in front of the male during the sperm transfer phase (Cuthbertson,
1984; Kirchmair & Raspotnig, 2021). In Dendochernes cyrneus, the data related
to quiescence is anecdotal since it is based on field observation of a single cou-
ple during the mating dance (Cuthbertson, 1984); thus, the quiescence phase
cannot be determined.
6. Arachnids and quiescence levels

For operational purposes, we recognize three quiescence levels
(Table 6). Level I is characterized by no apparent change in female body pos-
ture. In this level, the female remains quiescent during the precopulatory
phase, and the behavior may continue during sperm transfer. The female
moves immediately after the male separates. As for Level I, Level II is char-
acterized by no sign of body postural change. In this level, females remain
quiescent during the copulatory and/or postcopulatory phases. In some
cases, females may remain quiescent long after copulation has finished.
Finally, Level III is characterized by a drastic female postural change that
may include extension or flexion of legs and sometimes total body relaxa-
tion. This behavior occurs in any of the three phases of mating.
Spiders present the three quiescence levels: 13.1% of the species
(n ¼ 11 of 84) show Level I, 7.14% (n ¼ 6 of 84) show Level II, and
63.1% (n ¼ 53 of 84) show Level III. Unfortunately, 16.7% (n ¼ 14 of 84)
of the spider species for which quiescence has been reported could not be
assigned to one of the three levels because the articles do not present clear
descriptions of the body and appendages’ position during quiescence.
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Female quiescence in solifuges has been reported in 13 species distributed

in four families (Table 3). All can be classified as quiescence Level III. The
most controversial case in the order occurs in the family Eremobatidae since
there is no consensus between the authors on the occurrence of quiescence
because females can perform some movements while quiescent (Peretti et al.,
2021). For this revision, we will consider that eremobatids present quiescence,
making the corresponding clarifications of the level when necessary.
Although not explicitly described in the literature as quiescence per se, a
period of quietude, immobility, or female passivity has also been reported in
many scorpions (Benton, 1993; Oviedo-Diego, 2022; Peretti, 1995, 2001;
Polis & Sissom, 1990). Quiescence Level I occurs in seven species from three
families, and Level III in 19 species from three families (Table 4). Thus, 26%
of the species (n ¼ 7 of 27) showed female quiescence level I, whereas 70%
(n ¼ 19 of 27) showed female quiescence Level III. In five species, the level
of quiescence was not determined because female body posture during qui-
escence in the initial stage was not reported. Both quiescence levels may
appear in the same species in different moments of the mating interaction,
first during the initial stage (Table 4A) and then during the sexual sting
behavior (Table 4B).
Despite being a relatively well-studied order concerning their reproduc-
tive biology (e.g., Machado & Macı́as-Ordóñez, 2007; Machado, Requena,
Toscano-Gadea, Stanley, & Macı́as-Ordóñez, 2015; Macı́as-Ordóñez et al.,
2010), reports of female quiescence are scarce in harvestman (Table 5). In
addition, this female behavior is not described in detail. Female quiescence
has been reported in six species distributed in four families, with the
Gonyleptidae concentrating records for 3 species. All harvestman species
can be classified as showing quiescence Level I, except for Iporangaia pustulosa
(Fig. 2C), which shows quiescence Level II. Thus, harvestmen show only
two quiescence levels: 83.3% of the species (n ¼ 5 of 6) show Level I and
16.7% (n ¼ 1 of 6) Level II. Finally, whip spiders show female quiescence
Level III (n ¼ 2), whereas pseudoscorpions (n ¼ 1) show female quiescence
Level I (Table 5).
7. Behavioral and morphological traits associated

with quiescence
In this section, we present and discuss some aspects of the arachnids’
life and their relationship with quiescence. The following issues are summa-
rized in Tables 1–5.
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7.1 Quiescence and female body position

As we have already described, postural changes are notorious in quiescence
Level III. For example, in wolf spiders, funnel web spiders (Agelenidae), crab
spiders (Thomisidae and Philodromidae), and desids (Desidae), females flex
their legs below the sternum with the patellae above the cephalothorax, very
close to the body line (e.g., Montgomery, 1903; Sentenská et al., 2020; Xiao
et al., 2015). On the contrary, in some species of wolf spiders, females extend
their legs forward and backward, and sometimes the cephalothorax is placed
near the substrate in an almost flat position (e.g., Costa & Capocasale, 1984;
Toscano-Gadea & González, 2020; Wilder & Rypstra, 2004). These pat-
terns are evident in wandering spiders or species that build a platform-like
web with a firm surface. Levels I and II were reported in several spider
species. For example, in cobweb spiders, ambiguous terms such as “body
lowering” or “abdomen upward” have been used (Berendonck, 2003;
Knoflach & Benjamin, 2003; Shulov, 1940; but see Andrade & MacLeod,
2015). Such descriptions indicate a mere positioning of the female body
during copulation without the dramatic postural change observed in other
spiders (Andrade & MacLeod, 2015). It is hard to imagine how females could
perform a remarkable postural change while still hanging from the web.
Thus, it seems unlikely that orb weavers and 3D-web spiders perform the
postural changes characteristic of ground species, with more drastic postural
changes of quiescence Level III. All tarantulas show Levels I and II of female
quiescence, except for Sickius longibulbi (Theraphosidae), which shows a
modification of the stereotyped copulatory position of all mygalomorphs
(Fig. 2G). In this species, the female lies directly with her dorsum on the
substrate while the male holds her tightly against it in a vertical position
(Bertani et al., 2008).
Postural changes of females during quiescence are evident in solifuges. All
species included in our dataset (n ¼ 13) follow a common pattern: the female
raises her abdomen (this is probably due to a joint action between the male and
the female), forming an angle of approximately 90° with their cephalothorax
(Fig. 2B). In some species such as Galeodes caspius subfuscus the abdomen
may be rotated to the dorsal side while in Othoes saharae the abdomen is over
the cephalothorax of the female (Berland, 1932; Cloudsley-Thompson,
1967, 1977; Heymons, 1902; Hrušková-Martišová, Pekár, & Bilde, 2010;
Hrušková-Martišová et al., 2007; Junqua, 1962, 1966). This pattern is not
the only postural change reported in solifuges. Several species also change
the position of legs and pedipalps (Table 3). Changes in leg position have
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been reported in representatives of all families (except Eremobatidae), with

females flexing their legs against the body. However, changes in the position
of the pedipalps have been reported only in representatives of Galeodidae
and Solpugidae. Interestingly, the behavioral postures are not the same
for both families. In G. caspius subfuscus, females remain with their pedipalps
stretched in front of their body during quiescence (Berland, 1932; Cloudsley-
Thompson, 1967, 1977; Heymons, 1902; Hrušková-Martišová, Pekár, &
Bilde, 2010; Hrušková-Martišová et al., 2007), whereas in Metasolpuga picta
the pedipalps remain against their chelicerae (Wharton, 1987).
In scorpions, quiescence may or may not involve female postural changes
(Table 4A and B). Quiescence Level I is present in 26% of the species (n ¼ 7
of 27) and this type of quiescence always involves female immobility during
the sexual sting. Only females of Megacormus gertschi bend her metasoma
(“tail”) to one side, placing it on the substrate (Olguı́n-Perez et al.,
2021). However, this body position is commonly adopted by scorpions
when they are resting or a common position during the mating dance; thus,
we consider it also a case of quiescence Level I. Quiescence Level III, which
occurs in 70% of the species (n ¼ 19 of 27), involves female immobility with
pedipalps and metasoma extended backward, and relaxed legs in 42% (n ¼ 8
of 19) of the species, with a case with the same body posture but exhibiting
an elevated metasoma. In 11% of the species (n ¼ 2 of 19), females withdraw
the pedipalps to a contracted position close to their body. In one case, the
female has an immobile posture with pedipalps next to the body, metasoma
supported on the mesosoma, and legs withdrew towards the body. In 26%
of the species (n ¼ 5 of 19), the female is reported immobile, with the
body relaxed without detailing the position of the appendages or metasoma
(Table 4).
In contrast, none of the harvestman species surveyed adopts a noticeable
postural change. The only subtle postural change we can mention occurs
during the copulatory phase, in which the female is positioned in front of
the male, in the same position as he takes. It seems likely that the female’s
quiescence as well as the position that she adopts during copulation is a
prerequisite for sperm transfer (see Section 9.3).
In whip spiders, a clear postural change occurs in Damon variegatus and
Phrynus barbadensis (Alexander, 1962). Females of both species rest their
bodies on the ground and spread their legs out sideways. The pedipalps
are folded from the semi-extended position into a rest position, and the
antenniform legs become motionless. This change in body posture is
consistent with the classification of quiescence Level III.
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Finally, in the pseudoscorpion Dactylochelifer latreillii latreillii the female

remains quiescent in front of the male during the sperm transfer phase with
no evident postural change, which corresponds to quiescence Level I
(Kirchmair & Raspotnig, 2021).
7.2 Quiescence and female movements

Besides being quiescent, females may execute subtle movements that indi-
cate their willingness to mate. These movements may be brief, but appar-
ently crucial to facilitating the intromission of the male copulatory organ.
For example, some wolf spiders such as Trochosa terricola, Rabidosa rabida,
and several Alopecosa species (Engelhardt, 1964; Just et al., 2018; Rovner,
1971), swivel the abdomen while quiescent. This distinctive behavior
may allow the male to achieve the female genital opening and insert his ped-
ipalps. According to Foelix (2011), abdominal swiveling is performed by
females in all “modern” wandering spiders (e.g., Clubionidae, Lycosidae,
Salticidae, and Thomisidae) (but see Engelhardt, 1964; Rovner, 1971). In
the tarantula Telechoris striatipes, quiescent females perform occasional body
quivering. Also, genitalia protruding is frequent, allowing exposure to the
genital opening (Coyle & O’Shields, 1990). The female is in control of her
movements during quiescence; in fact, she can interrupt quiescence at any
time. For example, among the wolf spiders Alopecosa trabalis and Diapontia
uruguayensis, females may end the quiescence and walk with the male on their
dorsum ( Just et al., 2018; Piacentini et al., 2017). In the white widow spider,
Latrodectus pallidus, the female usually moves their legs while quiescent and, in
some cases, she can liberate herself from the threads spun around her by the
male and stop copulation (Shulov, 1940).
In solifuges, females become completely quiescent (Level III), as reported
for the families Galeolidae, Solpugidae, and some Amotrechidae. A striking
feature of quiescence in several species of galeodids are the involuntary
“spasms,” which consist of slight movements of the females’ pedipalps and
legs. Although the authors have used the term involuntary, no experimental
evidence excludes a female control over the spasms, which could have evolved
in a context of inter-sexual communication (see Section 9.5). In Eremobates
palpisetulosus, females may perform some movements, such as walking in tan-
dem in front of the male during mating, or even prey on insects in the middle
of the mating sequence (Muma, 1966a; Peretti et al., 2021; Punzo, 1997).
Females may also move the second and third pairs of legs, and chelicerae
before sperm transfer (Peretti et al., 2021 and references therein). The position
of the abdomen is also different between species with total quiescence and
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eremobatids. While the abdomen is bent forward more than 90° in the first
ones (e.g., Galeodidae), the angle formed by the abdomen and the body line of
eremobatid females is around 90° (Peretti et al., 2021). In general, females
showing quiescence Level III move much less during mating than those
species that never become quiescent, such as Oltacola chacoensis (Peretti &
Willemart, 2007) or Gluvia dorsalis (Hrušková-Martišová, Pekár, &
Cardoso, 2010), in which females seem to struggle to free themselves from
the males during the entire mating, especially at the end of the sperm transfer
phase. Future studies are necessary to determine the occurrence of these
movement differences in these taxa and whether they may involve a female
sexual strategy.
In most scorpion species there were no reports of female movements
during quiescence level I (Table 4A); only Urophonius achalensis and
U. brachycentrus females performed subtle leg movements. Similarly, during
quiescence Level III (Table 4A), most studies do not describe any female
movement. However, females of U. achalensis, U. brachycentrus, and
Leiurus quinquestriatus performed subtle leg and pedipalp movements, and
females of Bothriurus flavidus performed metasoma erection (Table 4).
Female movements during quiescence have been observed in three har-
vestman species. These behaviors range from slow leg movements before sep-
aration in Pachyloides thorellii (Stanley et al., 2016) to strong pulling back
movements and intense leg taps on the male in Neosadocus bufo (Willemart
et al., 2009). These authors described P. thorellii females are “almost” immobile.
We did not find any reports of female movements during quiescence in
whip spiders and pseudoscorpions.
7.3 Quiescence and male behaviors

Some authors suggested that female quiescence is triggered by males, appar-
ently without any female cooperation. In the spiders Agelenopsis aperta and
A. pennsylvanica, quiescence Level III appears to be induced by male pher-
omones (Becker et al., 2005; Pruitt et al., 2014; Singer et al., 2000). Singer
et al. (2000) speculate that males can also become quiescent by contacting
their own pheromones in the female web. However, no study has demon-
strated whether pheromones play a role in male quiescence. Another inter-
esting behavior occurs when the male bites the female before quiescence.
This has been reported in the spiders Wadotes bimucronatus, Rabidosa rabida
(Rovner, 1971; Schoenberg, 2021), Thanatus fabricci (Sentenská et al.,
2020), and Sickius longibulbi (Bertani et al., 2008), all with quiescence
Level III. Apparently, this quiescence level could be a response to venom
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injection (Sentenská et al., 2020) or purely a mechanical stimulus, as has been

reported for scorpions (Peretti, 2014). In many spider species, female quies-
cence Level III appears to respond simply to male mounting during copu-
lation, as in A. aperta and Hololena curta (Gering, 1953; Singer et al., 2000;
Xiao et al., 2015), perhaps triggered by male tactile stimulation (Becker
et al., 2005; Singer et al., 2000). A similar behavior has also been reported
in two species of wolf spiders, Alopecosa taeniata and A. cuneata ( Just
et al., 2018).
Except for the hypothetical cases of pheromones, some contact between
the mating pair seems to be necessary before the female becomes quiescent
(regardless of the quiescence level). This contact may include different body
parts. Leg and pedipalp touching have been reported in Arctosa alpigena lamperti
(Dolejš et al., 2012), A. aperta, (Gering, 1953), Acanthogonatus tacuariensis
(Capocasale & Perez-Miles, 1990), and Latrodectus hesperus (Vibert et al.,
2016). Clasping between the legs of both sexes occurs in Xysticus ferox
(Montgomery, 1903, under X. stomachosus), or between the legs and the ped-
ipalps as in Thelechoris striatipes and Microhexura montivaga (Coyle, 1985;
Coyle & O’Shields, 1990). It is beyond the scope of this review to detail
all behaviors of each species. However, it seems that some level of female col-
laboration is required for the occurrence of quiescence and some authors have
proposed that quiescence may reflect mate choice or copulation collaboration
from the female (Fraser, 1987; Sadana, 1972; Wilder & Rypstra, 2004; Xiao
et al., 2015). For more details on the behaviors performed by males prior to
quiescence and their relationship with the quiescence level, see Table 2.
As we mentioned before, a male-biased view has influenced the study of
sexual behavior in solifuges. Female quiescence is considered to be a
male-induced behavior, in which the term to induce has a connotation, even
implicit, of manipulation or coercion. Although there are variations among
species, a prerequisite for the onset of female quiescence is that males touch
females on some region of their body (this may vary according to species, see
Peretti et al., 2021). For example, in Galeodes spp., the male approaches from
the frontal region and touches the female with his pedipalps. The female
become quiescent, and then the male grabs the lateral area of the female’s
abdomen with his chelicerae. The exception to this general pattern within
the family Galeodidade occurs in Othoes saharae, in which female quiescence
begins after the male bites the female with his chelicerae ( Junqua, 1962,
1966). Unlike most spiders, solifuges do not have venom glands associated
with their chelicerae (Punzo, 1998a), so the bite could only produce a
mechanical stimulus to the female. A stunning case has been reported for
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Galeodes sulfuripes, mainly because of the description of how females become

quiescent. Amitai et al. (1962) describes it as follows: “...as if receiving an
electrical shock, she fell down in a convulsion and her whole body appeared
to be paralyzed”. In this description the authors attribute a predominant role
for the male, who generates female quiescence, and she responds passively. It
is evident that the female’s quiescent response is not considered a behavior
under her own control. However, females in this and other species of solifuges
becomes quiescent only as long as they are sexually receptive, otherwise they
reject the male, attacking him or simply moving away (Peretti et al., 2021).
The case of Eremobatidae is more challenging. Muma (1966a), Muma and
Muma (1988), and Punzo (1997) describe the males as striking females with
their pedipalps during the attack phase, a pattern similar to that which occurs
in Galeodes. Likewise, in Eremobates pallipes, the female has been described as
becoming quiescent after the male contacts her, either by touching her with
his pedipalps or chelicerae (Rowsell & Cushing, 2020). However, Punzo
(1998b) suggests that this stroke in Galeodidae and Solpugidae is not per-
formed at all by males in Eremobatidae. Therefore, although there is no
consensus, some degree of contact between sexes seems to be necessary for
female cooperation, including quiescence, during mating. Before quiescence
Level I, scorpion males perform a sexual sting behavior in 100% of the species
(Peretti, 2014; Polis & Sissom, 1990). Male sexual sting is characterized by
placing or inserting the stinger of the telson in some area of the female’s body.
If males perforate the female’s tegument in all species or inoculates venom is
unclear (Benton, 1993; Francke, 1979; Peretti, 2014; Polis & Sissom, 1990;
Toscano-Gadea, 2010). However, Olguı́n-Perez et al. (2021) confirmed cuti-
cle piercing and venom inoculation in M. gertschi. Although this behavior can
be considered as “aggressive”, the female remains immobile (Francke, 1979;
Jiao & Zhu, 2010; Olivero et al., 2019; Olguı́n-Perez et al., 2021; M. Oviedo-
Diego pers. obs.). On the other hand, before quiescence Level III takes place,
in 68% of the species with this type of quiescence (n ¼ 13 of 19) the male
grasps a female body region and/or performs vibratory behaviors. In 37%
of the species (n ¼ 7 of 19) the male performs grasping and metasoma shaking
(Table 4A).
As in the other arachnid orders, we can find particular behaviors per-
formed by males before quiescence in harvestmen. A peculiar case is that
of Quindina albomarginis, probably due to the nontraditional sexual roles.
In this species, males build mud nests where the females deposit their eggs,
which are protected by the males (Mora, 1990). Females initiate courtship by
touching the substrate inside the nest with their first pair of legs, then
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touching the male with their second pair of legs. At different times during
this sequence, the female becomes quiescent for up to 20 min until the male
starts touching her with his first pair of legs, and courtship resumes (Mora,
1990). Although the author does not explain the function of female quies-
cence, it could be a case of intersexual communication (see Section 9.5). In
species with copulatory quiescence, female quiescence begins after penis
intromission. This suggests that penis intromission is responsible for trigger-
ing female quiescence (Fowler-Finn et al., 2014; Stanley et al., 2016;
Willemart et al., 2006, 2009). In Iporangaia pustulosa, the only harvestman
species with postcopulatory quiescence, males perform postcopulatory mate
guarding of the female, touching her first pair of legs with their second pair
of legs throughout the process. The female can remain quiescent for up to
5 h near the male (Requena & Machado, 2014).
Finally, during female quiescence of whip spiders, the male can push the
female’s body with their pedipalps (Alexander, 1961). It does not appear to
occur any specific male behavior preceding female quiescence. The court-
ship in Damon variegatus and Phrynus barbadensis consists of gentle strokes by
the male’s first legs on the female’s body, even touching her ventral region
near her genital opening. The male also “threatens” her with his extended
pedipalps. The female may attack him or try to escape but may respond
with mutual touching (Alexander, 1961, 1962). Female quiescence in whip
spiders starts after male courtship (Alexander, 1962).
7.4 Quiescence, aggression, and cannibalism

For some spiders, aggression and cannibalism are relevant, given that females
can impose strong selection on male mating and fertilization strategies
(Dharmarathne & Herberstein, 2022; Tuni, Schneider, Uhl, & Herberstein,
2020). In this context, some authors have proposed that quiescence is a way
for females to respond to male behaviors during courtship, such as pedipalp
drumming, leg flexing, and abdominal vibrations in the funnel web spiders
Hololena adnexa, H. curta, and Agelenopsis aperta, as well as in the wolf spider
Rabidosa rabida (Becker et al., 2005; Fraser, 1987; Schoenberg, 2021; Xiao
et al., 2015). These repertoires may minimize the chances of the male being
cannibalized during courtship (Xiao et al., 2015). In fact, we found reports
of females attacking or cannibalizing males during mating in 38.1% of the spider
species (n ¼ 32 of 84) in which females show quiescence. In contrast, 61.9% of
the spider species (n ¼ 52 of 84) with quiescence are non-cannibalistic or there
is no mention of females attacking males during mating.
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Interestingly, cannibalism and sexual attacks are present in spiders with

different types of quiescence levels, being common among cobweb spiders,
which show quiescence Levels I and II. Within Level III, the families
Agelenidae and Homalonichidae contain more species with occurrence of
sexual cannibalism (Table 1). Overall, the data reveal that female quiescence
is not necessarily a characteristic behavior of cannibalistic or aggressive spiders.
However, it is possible that it has been one of the most common explanations
because spiders are so often considered to be highly aggressive and cannibal-
istic. This conception is held today only for some taxa (e.g., Araneidae and
Theridiidae) (Dharmarathne & Herberstein, 2022). However, we do not
reject the possibility that some male behaviors, such as the production of pher-
omones or bites (Becker et al., 2005; Sentenská et al., 2020), may trigger
female quiescence in some species.
A remarkable case of aggression and cannibalism occurs in solifuges.
They are voracious arachnids, and attacks and cannibalism between sexes
have been frequently reported (Punzo, 1998a). However, there are no
observations or studies that clearly document cannibalism during mating
in the field. Many of these reports could be laboratory artifacts, particularly
if the individuals were confined in small containers during behavioral obser-
vations (Peretti et al., 2021). As we mentioned, sperm transfer in solifuges is
mediated by the male chelicerae, which are inserted into the female genitalia
to deposit and manipulate the spermatophore. Junqua (1966) mentioned
that during this process, females can be injured and eventually consumed
by the male (the “nutrition instinct” prevails over reproduction). In contrast,
females may also attack males (Punzo, 1998b). We found that reports of
female quiescence are more common among species in which females attack
or cannibalize males during mating (76.9%, n ¼ 10 of 13) than in species in
which no aggressive behaviors between sexes have been reported (23.1%,
n ¼ 3 of 13).
Although cannibalism is common in scorpions (Benton, 2001; Polis &
Sissom, 1990), articles documenting species showing female quiescence
Level I rarely mention sexual cannibalism (Peretti, Acosta, & Benton,
1999; Tallarovic et al., 2000; Table 4B). In the same way, female quiescence
involving drastic body postural changes (Level III) seems unrelated to can-
nibalism or sexual cannibalism since data revealed no aggressiveness between
males and receptive females, except some reports in L. quinquestriatus and
Nebo hierochonticus during quiescence Level III (Brantley, 2017; Rosin &
Shulov, 1963; Table 4A). When cannibalism occurs in a non-sexual
context females are typically non-sexually receptive or deprived from food
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(Peretti et al., 1999). Consequently, avoidance of cannibalism or attacks,

although an attractive hypothesis, is not supported as a general explanation
for female quiescence in scorpions.
Unlike other arachnids, such as spiders and solifuges, cannibalism in a
mating context or sexual attacks do not occur in harvestmen (Machado &
Macı́as-Ordóñez, 2007; but see Parisot, 1962 for a dubious report under con-
fined laboratory conditions). In this way, aggressiveness can be ruled out as a
possible explanation for female quiescence in this order. Reports of attacks can
be found in Iporangaia pustulosa (Requena & Machado, 2014), in which caring
males reject and are aggressive toward visiting females that may consume some
eggs prior or during oviposition (Requena & Machado, 2014). Such aggres-
sive behavior may be related to egg defense by the males, and it has also been
reported for Quindina albomarginis (Mora, 1990).
7.5 Quiescence and sexual size dimorphism

Among species in which females attack or cannibalize males during sexual
interactions, we also explored the relationship between sexual size dimor-
phism (SSD) and female quiescence. Can tiny males be more likely cannibal-
ized by large females? If so, are they using any strategy to “induce” female
quiescence to avoid cannibalism? The general pattern in spiders is a
female-biased SSD (McLean, Garwood, & Brassey, 2018), with some extreme
cases, such as those reported in the families Thomisidae and Araneidae
(Hormiga, Scharff, & Coddington, 2000). This female-bias SSD has a strong
relationship with sexual cannibalism, meaning that the greater the difference
in size between sexes, the more frequent is sexual cannibalism (Wilder,
Rypstra, & Elgar, 2009). Based on our analysis, we establish that 15.5% of
the spider species (n ¼ 13 of 84) with reports of female quiescence show a mar-
ked SSD, while 84.5% do not show a marked SSD (n ¼ 71 of 84). Moreover,
female-biased SSD seems to be more common in species showing quiescence
Levels I and II (Table 1).
In solifuges, females of all species included in our dataset are larger than
males, following the general pattern for the entire order (McLean et al.,
2018). This would give the female the option to cannibalize the males with-
out further risk. However, there are no studies that correlate SSD to sexual
cannibalism, nor with female quiescence in this arachnid order.
Although female-biased SSD is a general pattern in scorpions (McLean
et al., 2018), there were no reports of SSD (or females slightly larger than
males) in species showing quiescence Level I (Table 4B). In species showing
quiescence Level III (Table 4A), females were similar or slightly larger than
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males in 89% of the species (n ¼ 17 of 19), while in only two species females
were clearly larger than males.
In harvestmen, the direction of SSD depends on the family (McLean
et al., 2018). Nevertheless, females of most of the species included in our
dataset are larger than males. Moreover, the species also tend to be sexually
dimorphic in other morphological structures, such as legs, pedipalps, and
chelicerae (McLean et al., 2018). However, these characteristics were not
considered here.
In whip spiders and pseudoscorpions, SSD is female-biased (McLean
et al., 2018), but no information related to SSD is available for the species
showing quiescence.
8. Quiescence and other biological traits in spiders

In this section, we describe quiescence in relation to unique biological
traits in spiders, such as their hunting strategies, copulation position, and
female genital complexity (Table 7).
8.1 Quiescence and hunting strategies

The hunting strategies of spiders can be broadly divided into three types
(Foelix, 2011): those that build webs (web builders), those that ambush
or chase the prey without using catching webs (wandering spiders), and
those that live in burrows or under rocks without abandoning it and that
do not use silk threads or web for catching prey (sedentary spiders). Our data
show that 48.8% of the species (n ¼ 41 of 84) with female quiescence occur
in wandering spiders, mainly among wolf spiders (n ¼ 35). In contrast, web
builders represent 40.5% of the species (n ¼ 34 of 84), mainly in the cobweb
spiders (n ¼ 11). Finally, sedentary spiders represent 10.7% of the species
(n ¼ 9 of 84), mainly in tarantulas of the family Theraphosidae (n ¼ 6).
Quiescence Levels I and II were observed in web builders, mainly in cobweb
spiders and sedentary tarantulas. In turn, Level III was observed in web
builders with a different web morphology (i.e., sheet webs of the families
Agelenidae and Desidae), and among the wandering spiders only in wolf
spiders (Tables 1 and 7).
8.2 Quiescence and copulation position

In spiders, von Helversen (1976) and later Foelix (2011) proposed three types
of copulatory positions. Type I is common in more “primitive” spiders such as
segmented trapdoor spiders (Mesothelae), tarantulas (Mygalomorphae), and
Table 7 Spiders’ life traits and values according to the three levels of female mating quiescence proposed in this review.
Cannibalism or Sexual size
Hunting strategies sexual attack dimorphism (SSD) Mating position Genital complexity
Quiescence Web
levels builder Wandering Sedentary Yes No Yes No Type I Type II Type III Entelegyne Haplogyne
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Level I 9.52% 0% 3.57% 9.38% 15.38% 7.69% 14.08% 66.67% 7.69% 0% 1.45% 66.67%
(n ¼ 8) (n ¼ 0) (n ¼ 3) (n ¼ 3) (n ¼ 8) (n ¼ 1) (n ¼ 10) (n ¼ 10) (n ¼ 1) (n ¼ 0) (n ¼ 1) (n ¼ 10)
Level II 5.95% 0% 2.38% 12.5% 3.85% 15.38% 5.63% 20% 15.38% 3.57% 5.8% 20%
(n ¼ 5) (n ¼ 0) (n ¼ 2) (n ¼ 4) (n ¼ 2) (n ¼ 2) (n ¼ 4) (n ¼ 3) (n ¼ 2) (n ¼ 2) (n ¼ 4) (n ¼ 3)
Level III 15.48% 44.05% 2.38% 53.13% 69.23% 23.08% 70.42% 6.67% 15.38% 87.50% 73.91% 6.67%
(n ¼ 13) (n ¼ 37) (n ¼ 2) (n ¼ 17) (n ¼ 36) (n ¼ 3) (n ¼ 50) (n ¼ 1) (n ¼ 2) (n ¼ 49) (n ¼ 51) (n ¼ 1)
Unspecified 9.52 4.76% 2,38% 25% 11.54% 58.85% 9.86% 6.67% 61.54% 8.93% 18.84% 6.67%
(n ¼ 8) (n ¼ 4) (n ¼ 2) (n ¼ 8) (n ¼ 6) (n ¼ 7) (n ¼ 7) (n ¼ 1) (n ¼ 8) (n ¼ 5) (n ¼ 13) (n ¼ 1)
Total 40.47% 48.81% 10.71% 38.09% 61.9% 15.48% 84.52% 17.86% 15.48% 66.67% 82.14% 17.86%
(n ¼ 34) (n ¼ 41) (n ¼ 9) (n ¼ 32) (n ¼ 52) (n ¼ 13) (n ¼ 71) (n ¼ 15) (n ¼ 13) (n ¼ 56) (n ¼ 69) (n ¼ 15)
The table includes information on 84 spider species for which we obtained data female mating quiescence.
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haplogyne spiders (see definition in Section 8.3). In this position, the male
approaches from the front, the female lifts the cephalothorax, and the male
inserts one or both pedipalps into the female genital opening. Type II is pre-
sent in web builders, and it is a derivation of the type I, as the spiders copulate
in the same position but hanging upside down from the web. Type III is wide-
spread in wandering spiders, such as wolf spiders, jumping spiders, and flower
crab spiders. The male climbs onto the female’s cephalothorax and then turns
to the right or left side of the female’s abdomen. The female rotates her abdo-
men so the male can insert one of the pedipalps. Each time one of the ped-
ipalps is used, the male switches to the other side of the female’s abdomen.
Among the spider species included in our data set, 17.9% (n ¼ 15 of 84)
copulate in Type I position, 15.5% (n ¼ 13 of 84) in Type II position, and
66.7% (n ¼ 56 of 84) in Type III position. When mating positions were
related with the levels of female quiescence, 66.7% of the species
(n ¼ 10 of 15) with quiescence Level I copulated in Type I position. In con-
trast, 87.5% of the species (n ¼ 49 of 56) with quiescence Level III copulated
in Type III position. Finally, an equal percentage of 15.4% of the species
(n ¼ 2 of 13) with quiescence Levels II and III copulated in Type II position
(Table 7).
8.3 Quiescence and genitalia

Gering (1953) proposed that the “cataleptic” state he observed in females of
agelenid spiders represents an adjustment to the complexity of male and
female genitalia and mechanics of copulation: unless the female is immobile,
the complex male genitalia cannot fit appropriately the female genitalia.
Even subtle movements of the female during certain phases of copulation
could preclude the chances of correct coupling of genital structures.
Following the same reasoning, Coyle and O’Shields (1990), proposed that
the female, “even in a cataleptic state,” may provide a mechanical challenge
to the male’s copulatory ability, monitoring his performance, and adjusting
her behavior and/or physiology to maximize her fitness. Based on this
hypothesis, we evaluated whether quiescence is linked with male and
female genital complexity taking as a reference the classical definition of
haplogyne and entelegyne genitalia of spiders (Foelix, 2011; Figs. 4-5). In
general, the entelegyne genital morphology tends to be more complex than
the haplogyne, both in females and males. The female haplogyne genitalia
have a single genital opening where the sperm travel both to and from
the spermatheca. The male pedipalps are inserted in the primary genital
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Fig. 4 Male spider genitalia of haplogyne and entelegyne types. (A) Haplogyne pedi-
palp of a male of Ariadna boesenbergi (Segestriidae). (B) Entelegyne pedipalp of a male
of Trachelas mexicanus (Trachelidae). Note the complexity of structures in the
entelegyne model. Credits: Cristian Grismado.
opening (the opening through which the eggs are laid). In contrast, the
female entelegyne genitalia have a pair of extra copulatory openings that
connect to the spermatheca, where the male inserts their pedipalps
during copulation. These additional openings are near the primary genital
opening and are located in a sclerotized plate called the epigynum. The
epigynum co-evolves as a counterpart of the male pedipalps and their
complexity (Eberhard & Huber, 2010). We found that 82.1% of the
species (n ¼ 69 of 84) with female quiescence were entelegyne spiders,
mainly wolf spiders (n ¼ 39). Haplogyne spiders comprises 17.9% of the spe-
cies (n ¼ 15 of 84) with female quiescence reports, mainly tarantulas of the
family Theraphosidae (n ¼ 6). Although these results show a tendency
toward an association of female quiescence with more complex entelegyne
genitalia, they are probably biased due to the high number of studies on these
spiders, which comprises 39,923 species, i.e., 79% of all spider diversity
(Magalhaes & Ramı́rez, 2022, I. Magalhães pers. comm.). Classical models
for studies on sexual behavior among entelegyne spiders include families
such as Lycosidae (e.g., Aisenberg & Costa, 2005) and Theridiidae (e.g.,
Andrade & MacLeod, 2015). Among entelegyne spiders, we found mostly
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Fig. 5 Female spider genitalia showing the internal organization of the haplogyne
(A) and entelegyne (B) types. In the haplogyne type, copulation occurs via a unique gen-
ital opening of the female genitalia, and the sperm is stored in the spermatheca that
is connected to a single duct. The sperm uses the same duct during copulation and
fertilization (black arrows). In the entelegyne type, copulation occurs through the
copulatory orifices. An additional copulatory duct is connected to the spermatheca.
Fertilization of eggs (orange dots) occurs in a single way trajectory when the sperm
leaves the spermatheca through a different duct.
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quiescence Level III, except for Theridiidae in which there are only cases of
quiescence Levels I and II. In turn, haplogyne spiders tend to show almost
exclusively quiescence Levels I and II, with the only exception of the taran-
tula Sikius longibulbi, an haplogyne spider with quiescence Level III (Bertani
et al., 2008) (Tables 1–2 and 7).
Another pattern from our analysis is that quiescence Level III is found
almost exclusively in representatives of the RTA clade (Table 1). In this clade,
the genitalia of adult males have a retrolateral tibial apophysis (RTA) in the
pedipalps (Huber, 1995a; Wheeler et al., 2016). This apophysis is the first male
structure to lock against in particular areas of the epigynum during genital
coupling, allowing the male pedipalp to align in the appropriate position with
respect to the female genital opening (Huber, 1995a; Poy, Ramı́rez,
Michalik, & Piacentini, 2019). We will revisit this topic in Section 9.2.
9. Contexts that may have favored the evolution

of female quiescence
Multiple functions have been proposed for female quiescence, espe-
cially in spiders (Box 1). Therefore, there may not be a single evolutive pres-
sure that has favored its appearance in arachnids, but rather a combination
of behavioral, ecological, and evolutionary contexts acting simultaneously
or sequentially in each species. Based on information provided in the
previous sections, we evaluate several contexts that may have favored the
evolution of female quiescence in arachnids. These hypothetical contexts
are: (1) female sexual aggressiveness and cannibalism, (2) female and male
genital complexity, (3) sperm transfer, (4) copulation substrate constraints,
(5) intersexual communication and mate choice, and (6) postcopulatory
sexual selection (Box 1).
9.1 Female sexual aggressiveness and cannibalism

Although not a rule, there is a consensus that many arachnids, such as scor-
pions, solifuges, and some spiders, are more aggressive than other arthropods
(Elgar, 1992). In general, most arachnids are predatory and solitary animals
with a variable repertoire of aggressiveness, including cases of sexual canni-
balism. Sexual cannibalism can be interpreted as an extreme case of sexual
conflict resolution (Wilder et al., 2009). Several explanations for its evolution
have been suggested, such as female foraging, mate rejection, male sacrifice,
and genetic constraints (Andrade, 1998; Elgar, 1992; reviewed in Wilder
et al., 2009). The hypothesis of sexual aggressiveness and cannibalism as an
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explanation for female quiescence has been suggested for several families of
spiders (Agelenidae: Becker et al., 2005; Fraser, 1987; Xiao et al., 2015;
Theridiidae: Berendonck, 2003; Breene & Sweet, 1985; Kaston, 1970;
Shulov, 1940; Vibert et al., 2016; Philodromidae: Sentenská et al., 2020;
Theraphosidae: Ferretti et al., 2013). In solifuges, males and females have
generally been classified as highly aggressive (Punzo, 1998a). From a male
perspective, female quiescence may prevent males from being cannibalized.
As early 20th century articles suggested, female quiescence is necessary for
copulation in these “bloodthirsty” animals (Heymons, 1902), so that appeas-
ing the female is a requirement of males to avoid being cannibalized. From
a female perspective, quiescence may have evolved to prevent females
from being injured during mating. This is the most frequent explanation
for the evolution of female quiescence in solifuges (e.g., Heymons, 1902;
Hrušková-Martišová, Pekár, & Bilde, 2010; Junqua, 1966; Peretti et al.,
2021; Peretti & Willemart, 2007). For other taxa, our analysis of the literature
does not support cannibalism or sexual attacks as a cause of female quiescence.
Indeed, female quiescence has been reported in orders with absence of
cannibalism or potentially harmful sexual attacks, such as harvestmen, whip
spiders, and pseudoscorpions (Table 5).
In spiders, although most of the species with female quiescence do not
show cannibalism or sexual attack, there do seem to be some examples
supporting this explanation. For instance, in the wolf spider Rabidosa rabida
females that do not become quiescent are more likely to attack their sexual
partners (Schoenberg, 2021). In Hololena curta, when a female becomes
aggressive towards a courting male, he immediately vibrates its abdomen
and grabs her leg after which the female becomes quiescent (Xiao et al.,
2015). In Hololena adnexa, the male vibrates the female’s web, which causes
the female to become quiescent. If the female does not recognize the male
after the vibrations, she immediately attacks him (Fraser, 1987). Whether
and how the female recognizes the male is not clear. After copulation is fin-
ished, the female remains quiescent for a couple of seconds, giving the male
time to leave (Fraser, 1987). Agelenosis aperta is the only example of males
using pheromones to, according to the authors, “induce” female quiescence
and prevent cannibalism (Becker et al., 2005; Fischer, 2019). Apparently,
A. aperta is a particularly aggressive spider because it inhabits deserts
areas with limited prey availability (Becker et al., 2005; Hammerstein &
Riechert, 1988; Riechert & Tracy, 1975). This extreme aggressiveness
may lead females to frequently cannibalizing courting males (Riechert,
Singer, & Jones, 2001).
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Other examples in favor of the sexual aggressiveness and cannibalism

hypothesis can be found in tarantulas of the family Theraphosidae.
Historically, sexual cannibalism in species of this family was thought to be
the rule; even B€ ucherl (1952) proposed “slaughter” of females toward males.
Whether this is indeed a rule (Costa-Schmidt et al., 2017), it has been
suggested that female quiescence could allow the males to withdraw after
copulation without the risk of being attacked (Ferretti et al., 2013).
Among species of the family Theridiidae, in which sexual cannibalism has
direct implications for sperm transfer (Snow & Andrade, 2004), there is also
evidence that female quiescence may prevent cannibalism. In the widow
spiders Latrodectus pallidus and L. tredecimguttatus, for example, females are
prone to attack males after quiescence (Shulov, 1940). In L. revivensis, in
which 30% of males are cannibalized during or after copulation (most
frequently smaller males), the quality or quantity of male courtship may
induce female quiescence by decreasing female aggressiveness and thus
avoiding cannibalism (Berendonck, 2003). However, no study has evaluated
whether female quiescence is more frequent when females are paired with
smaller males. A relation between male behavior and quiescence has been
proposed for L. mactans and L. hesperus, in which the behavior of cutting
threads in the female’s web and the abdominal vibrations of the male before
copulation “induce” female quiescence and hence reduce cannibalism
(Breene & Sweet, 1985; Scott et al., 2012).
Female quiescence is not the only mechanism that could prevent sexual
cannibalism in spiders. The “bridal veil,” a term coined by Bristowe (1931),
is a series of silk threads spun by the male to anchor a female’s legs or ceph-
alothorax to the substrate and it has been reported in several spider families
(e.g., Theridiidae: Berendonck, 2003; Scott et al., 2012; Shulov, 1940;
Philodromidae: Sentenská et al., 2020). Although its function is still unclear
(Scott, Andersson, & Andrade, 2018), it has been suggested that bridal veil
gives males an extra couple of seconds to avoid being cannibalized (Breene &
Sweet, 1985; Kaston, 1970; Scott et al., 2018), especially in the context of
coercive copulations (Breene & Sweet, 1985). Additionally, the bridal veil
may also reduce female aggressiveness by means of male pheromones
(Sentenská et al., 2020). Although females can escape from the bridal veil
(Scott et al., 2018; Sentenská et al., 2020), they must be quiescent for the
male to be able to spin over her.
Finally, an additional aspect for spiders is the relationship between female
copulatory quiescence, male mating position, and cannibalism. Wilder et al.
(2009) proposed that cannibalism is more frequent when the copulation
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position of males is dangerously close to the female chelicerae or to the

region of the spinnerets and fourth pair legs, where the male can be wrapped
very quickly by the female. Following this logic, cannibalism should be
more frequent in spiders that copulate in Type I and II positions, such as
cobwebs and tarantulas, than in those spiders that copulate in Type III posi-
tion, such as wolf spiders. Although this relationship holds in general, female
quiescence has been reported in taxa with all copulation positions, which
indicates that cannibalism should not be taken as the sole explanation for
female quiescence in spiders.
In solifuges, most of the species in which quiescence has been reported
show cannibalism or sexual attacks. Although it has not been explicitly pro-
posed as a functional hypothesis, at least one example supports the hypothesis
of sexual aggressiveness and cannibalism in solifuges. In Galeodes caspius sub-
fuscus, when a female becomes aggressive, the male touches her cephalothorax
with the suction organs of his pedipalps and grasps her with his chelicerae.
After the grasp, the female becomes quiescent. This description could be
evidence that female quiescence plays a role in reducing her aggression and,
therefore, the chances of sexual cannibalism (Berland, 1932; Cloudsley-
Thompson, 1967, 1977; Heymons, 1902; Hrušková-Martišová, Pekár, &
Bilde, 2010; Hrušková-Martišová et al., 2008). However, it should be
remembered that cases of sexual cannibalism or attacks have been reported
only in laboratory conditions (Peretti et al., 2021). For example, Wharton
(1987) mentions that cannibalism in Metasolpuga picta is an effect of confine-
ment and something similar occurs in Titanopuga salinarum (A.V. Peretti and
F. Cargnelutti pers. obs.). Field observations suggest that males may escape
from sexually unreceptive or recently copulated females.
Regarding scorpions, sexual cannibalism can occur in some species
(Benton, 2001; Polis & Sissom, 1990). This phenomenon occurs mainly
during the precopulatory phase, especially in the first instances of mate rec-
ognition, and is extremely rare in the copulatory and postcopulatory phases
(Peretti et al., 1999). Precopulatory sexual cannibalism usually occurs when
there is a marked size difference in the mating pair (McLean et al., 2018).
Therefore, males are expected to have strategies to avoid being cannibalized
(Elgar, 1991; Polis & Farley, 1979). Avoidance of cannibalization or attacks,
although an attractive hypothesis, is not supported as a general explanation for
female quiescence in scorpions given its low frequency (Brantley, 2017;
Rosin & Shulov, 1963). The same argument applies for harvestmen, whip
spiders, and pseudoscorpions, in which no case of unequivocal sexual canni-
balism or harmful attacks during mating interactions has ever been reported.
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9.2 Female and male genital complexity

It has been hypothesized for spiders that female quiescence could be related
to genital complexity of both sexes. In fact, spiders have a great diversity of
genital morphologies, with taxa with relatively simpler genitalia and others
with more complex ones. Our results show that female quiescence has been
reported in 82.1% of the species (n ¼ 69 of 84) with complex genitalia
(entelegyne), and 17.9% of the species (n ¼ 15 of 84) with simple genitalia
(haplogyne). This pattern also occurs when we analyze the three quiescence
levels (Table 7). These results may be influenced by the higher diversity of
entelegyne spiders against haplogyne spiders, and by the fact that haplogyne
spiders are not generally considered as models in sexual selection studies.
Moreover, we are aware that classifying the genital complexity of spiders using
the dichotomy entelegyne and haplogyne is an oversimplification, as there are
examples showing that female genitalia of haplogyne spiders is more complex
than previously thought (Burger et al., 2006; Burger, Nentwig, & Kropf,
2003). However, in the absence of a more precise way of measuring complex-
ity, we will use entelegyne and haplogyne as proxies of morphological
complexity of male and female genitalia in spiders and address the limitations
of this approach below.
The genital complexity hypothesis has been defended by some authors
(Abregú et al., 2022; Coyle & O’Shields, 1990; Gering, 1953). In Metaltella
iheringi, for instance, the male has an extremely long embolus and females
present long copulatory ducts (D.A. Abregú and A.V. Peretti unpub. data).
Therefore, the authors suggest that female quiescence (Level III) may help
the successful insertion and removal of the long embolus during copulation
(Abregú et al., 2022). However, species showing quiescence Levels I and II,
such as the widow spiders of the genus Latrodectus, also have males with
extremely long embolus and female genitalia with long ducts (Snow,
Abdel-Mesih, & Andrade, 2006). Thus, although female quiescence may
be necessary for a correct genital coupling of complex genitalia, this female
behavior seems not to be correlated with quiescence level. Moreover, in
some species female quiescence may continue once the male has completed
intromission. If the main function of female quiescence is to favor genital
coupling, why would females remain quiescent once the male has finished
sperm transfer and removed his genitalia?
An interesting pattern is found in spiders belonging to the RTA clade
(see Section 8.3): all cases of quiescence Level III have been reported in spe-
cies from this clade (Table 2). Is there a relationship between quiescence
level III and the presence of the retro-tibial apophysis (RTA) on male ped-
ipalp? Is quiescence necessary for a precise genital coupling between sexes in
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these spiders? Currently, there are not enough studies to answer these ques-
tions. However, the relationship between the RTA and female quiescence is
challenging to establish. First, many representatives of this clade have sec-
ondarily lost the RTA, such as the wolf spiders (Griswold, 1993) and some
ghost spiders of the family Anyphaenidae (Ramı́rez, 2003), which also show
quiescence (D. Poy pers. comm.). Second, evidence supporting the function
of the RTA as a “anchor” is not conclusive. For example, in the funnel-web
spider Histopona torpida, the male pedipalp is fixed in the female epigyne gen-
ital opening with a patellar apophysis instead of the RTA (Huber, 1994). In
Textrix denticulata, it is the RTA that is locked to the female genitalia (Huber,
1994). In Allagelena similis (Osterloh, 1922, under Agelena gracilens, p. 348),
the patellar apophysis serves in the internal arrestation of the bulb, whereas in
Agelenopsis spp. (Gering, 1953, p. 13), the “RTA serves to limit the extent of
cymbial flexure” (summarized in Huber, 1994). In wolf spiders, the RTA is
absent, and the locking mechanism is accomplished by different sclerites of
the copulatory bulb (Poy et al., 2019). Notably, in Agalenocosa pirity, a species
with female quiescence (D. Poy pers. comm.), a basal tibial process (BTP)
is present but, as in the case of RTA of some agelenid spiders, it does not
participate in the genital coupling (Poy et al., 2019). Although the RTA
may not contact female genital structures in all species, a function during
the process of male pedipalp insertion is not entirely excluded. For
example, the RTA may allow the correct expansion of the bulb mem-
branes. Moreover, the RTA could have different functions in different
species demonstrating its high functional divergence (Huber, 1994,
1995a, 1995b). Whether quiescence has evolved as a female behavior that
allows the correct insertion of genital structures in the RTA clade will
need further evidence, particularly from “non-model” species outside wolf
spiders, flower crab spiders, and funnel web spiders. Finally, the multiple
losses of the RTA in spiders can be used to test the relationship between
this male trait and female quiescence in a broad phylogenetic context.
As mentioned above, we used the classification of entelegyne and
haplogyne as a proxy for male and female genital complexity. However,
are all entelegyne spiders equally complex and all haplogyne equally simple?
Is it valid to compare the genitalia between phylogenetically distant taxa or
with different reproductive strategies? The survey of this review showed that
the relationship between male and female morphological genital complexity
and Level III of female quiescence may not hold because many taxa of
spiders with complex genitalia show different quiescence levels or no
quiescence at all. Thus, to address the genital complexity hypothesis in spiders,
it is important to develop accurate methods to evaluate genital complexity
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(see examples in Azevedo, Griswold, & Santos, 2018; Kuntner et al., 2016;
Ramı́rez & Michalik, 2014).
The situation is more complicated in the other arachnid orders. However,
there are certain criteria for classifying genital complexity. In other taxa with
direct sperm transfer, such as solifuges, there are detailed descriptions of
the male chelicerae, the structure used for sperm transfer in most families
(Bird, Wharton, & Prendini, 2015). Again, data related to female genitalia
are scarce. However, some articles describing the genital operculum of the
females mention that it may have grooves, hollows, and furrows (e.g.,
Maury, 1980, 1984). Moreover, possible correlations between male chelicerae
shape and female genital operculum form have been established (Maury, 1980,
1984; Peretti et al., 2021; Peretti & Willemart, 2007). For example, in
Titanopuga salinarum the tip of the male movable cheliceral finger fits into
particular areas on the female genital sternite (A.V. Peretti pers. obs.).
In harvestmen the intromittent male genitalia may be very complex
(Macı́as-Ordóñez et al., 2010; Perez-González & Werneck, 2018).
Concerning females, the information gap is remarkable. Although some
studies have focused on female morphology, no study has focused on the
interaction between the genitalia of the sexes (but see Perez-González &
Werneck, 2018), which makes it challenging to interpret the relation-
ship between female quiescence and female genital complexity in this
order. As in other arachnid orders, clear criteria to quantify female genital
complexity have not been established, so it is difficult to rule out the
genital complexity hypothesis.
In orders with indirect sperm transfer, this complexity has been partly
analyzed through focusing on spermatophore morphology. In scorpions,
for instance, the spermatophores can be classified as flagelliform (relatively
simple) and lamelliform (more complex) (Francke, 1979; Peretti, 2010).
Although Peretti (2010) analyzed the female genitalia of scorpions with both
types of spermatophores, no attempt to classify their level of complexity
was made. A similar analysis of spermatophore complexity was performed
by Weygoldt (1969) on pseudoscorpions. In this case, the author classifies
spermatophores only as simple and complex. Interestingly, the simplest sper-
matophores occur in species with no pair formation, whereas the most com-
plex occurs in species in which there is pair formation for sperm transfer.
Unfortunately, we do not have information about female genital complexity
in pseudoscorpions. However, in taxa with indirect sperm transfer it is easy
to discard the genital complexity hypothesis because females must actively
collaborate to take up the spermatophore for sperm transfer to occur.
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9.3 Sperm transfer

Female quiescence has been reported in arachnids with both direct and indi-
rect sperm transfer. Given that each species has a unique repertoire of move-
ments during mating, it may be possible that female quiescence occurs in
different phases depending on the sperm transfer mode. For example, in
scorpions, pseudoscorpions, and whip spiders, which transfer sperm indi-
rectly, female quiescence is not expected to appear during the copulatory
phase, given that active female participation is necessary to position herself
over the spermatophore and pick up the sperm (Peretti, 2003, 2010;
Weygoldt, 1969, 1990, 2002). In contrast, in taxa with direct sperm transfer,
such as spiders and solifuges, female quiescence may occur during copulatory
phase. However, this does not prevent female quiescence from occurring at
other phases of the mating process.
In spiders, the evolution of quiescence as a mechanism allowing sperm
transference has been proposed for the black widow Latrodectus hasselti
(Snow & Andrade, 2004), and the tarantulas Grammostola vachoni (Copperi,
2018) and Eupalaestrus weijenberghi (Ferretti et al., 2013). In L. hasselti, the only
proposed hypothesis is that quiescence could favor sperm transfer, without
further details (Snow & Andrade, 2004). Within the tarantulas, the sperm
transference requires individuals facing each other, and male clasping the
female fangs with their first pair of legs and raising her to his pedipalps to reach
the genital opening (Costa-Schmidt et al., 2017 and references therein).
Therefore, it is possible that female quiescence without postural change is nec-
essary for the correct positioning of the pair. However, an exception occurs in
the theraphosid Sikius longibulbi, a species that does not follow the tarantula’s
typical copulation pattern mentioned above (for more details see Costa-
Schmidt et al., 2017). In this species, the female becomes quiescent after
the male bites her, possibly injecting venom, and touches her hind legs with
his third pair of legs. Then, the male pushes her so that she falls with her dor-
sum on the substrate, and he then positions his body at a 90° angle to the
female’s body, so sperm transfer takes place (Bertani et al., 2008). Since females
do not have spermatheca, quiescence in this position may favor sperm transfer
to their oviduct (Bertani et al., 2008).
Solifuges are an exciting group for studying female quiescence in a sperm
transfer context because there are two sperm transfer patterns in the order.
Species in most families use their chelicerae (Punzo, 1998a), whereas in
eremobatids sperm transfer occurs between the contact of genital openings
(Muma, 1966b). In both types of transferences, the male chelicerae are
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inserted at some point of the copulation into the female genital openings in a
behavior that has been called “chewing behavior” (Punzo, 1998a). One of
the functions of the chelicerae insertion is sperm transfer, but it may play a role
in genital stimulation or even in sperm competition (Peretti et al., 2021). For
example, in eremobatids, it has been proposed that males can place sperm
within the female genitalia, and also remove sperm from rival males (Bird
et al., 2015; Rowsell & Cushing, 2020). Has quiescence evolved in solifuges
to prevent the female from injuries during this mating phase? Although this
idea is attractive, some considerations are necessary. First, we can find some
species (e.g., Oltacola chacoensis) in which females do not become quiescent
during the entire mating but stay quiet only for the precise moment of sperm
transfer (Peretti & Willemart, 2007). Second, quiescence in eremobatids is
partial, lasting the few seconds it takes the sperm transference from genital
opening to genital opening. Quiescence probably helps to prevent the female
from being injured by the energetic movements of the male chelicerae
inside her genital atrium during sperm transfer. Although this idea cannot
be excluded, longer quiescence duration (as occurs in most families) can be
explained by other reasons (see Section 9.5).
In harvestmen, when mating takes place, the female faces the male in a
slightly tilted upwards position, and quiescence is necessary for the male to
access the female genital opening. Although, there are no explicit data show-
ing that sperm transfer explains female quiescence, it is evident that female
cooperation is necessary for sperm transfer in this order, as females may reject
male attempts to insert the penis by lowering the body and obstructing their
genital opening (Macı́as-Ordóñez et al., 2010). Moreover, even when the
female become quiescent, eversion of the ovipositor is required for the inser-
tion of the penis (Machado & Macı́as-Ordóñez, 2007).
In orders that use spermatophores, female quiescence (mainly Level III)
occurs during the precopulatory phase in scorpions and whip spiders
(Tables 4A and 5). In whip spiders, once the spermatophore is deposited
on the substrate, the female interrupts quiescence and positions herself over
the spermatophore by lowering her body and grasping it with her genital
opening (Alexander, 1961, 1962). The female jerks forward against the sper-
matophore in a quick fashion, to finally stand up with masses of sperm caught
under her genital opening (Alexander, 1961). The absence of female quies-
cence during copulation may be related to an active female participation
during sperm transfer.
Female quiescence occurs after the male positions the female on the sper-
matophore in pseudoscorpions (Kirchmair & Raspotnig, 2021). The male
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then uses the modified tarsi of his forelegs to open the genital atrium of the
females and favor the uptake of the spermatophore (Kirchmair & Raspotnig,
2021; Weygoldt, 1969). While we cannot rule out the female having a role
in this facilitation of the opening of her genital operculum in this instance,
the female remains quiescent until sperm transfer is complete.
9.4 Copulation substrate constraints

Spider species that copulate in a firm substrate seem to show the three qui-
escence levels, whereas species that copulate in unstable surfaces, like webs,
tend to show only quiescence Levels I and II. Indeed, quiescence Levels I
and II are especially clear in web builders, such as cobwebs, since they do
not have a stable platform suitable for body postural changes, typical of qui-
escence Level III (Table 7). Moreover, to perform postural changes of quies-
cence Level III in the web would imply a loss of stability of the female body
and hence a loss of control of the delicate movements of copulation while still
hanging. Among other web builders, we highlight species that build sheet and
tubular webs and show quiescence Level III (n ¼ 13 species of 85). These
webs, such as those in the family Agelenidae, have a horizontal platform with
a tube retreat (Eberhard, 1990; Herberstein & Tso, 2011), providing a firm
surface where postural change can occur. In other arachnids, species that cop-
ulate in firm substrate (e.g., soil or tree bark) are present in scorpions, solifuges,
harvestmen, scorpions, whip spiders, and pseudoscorpions.
9.5 Intersexual communication and mate choice

Animals use complex signals to communicate with conspecifics and other
species in various contexts and for multiple reasons (Hebets et al., 2016;
Hebets & Anderson, 2018). In arthropods, communication between sexes
can occur through multiple channels, such as visual or chemical, and indi-
viduals are able to perceive the information with multiple sensory channels
(Huber, 2005; Partan, 2013). One message that females may communicate is
their willingness to copulate. In this context, chemical communication plays
an essential role in mate recognition, female receptivity, and mate choice as
has been observed in spiders (Fischer, 2019; Huber, 2005; Roberts & Uetz,
2005) and scorpions (Oviedo-Diego, Costa-Schmidt, Mattoni, & Peretti,
2021; Romero-Lebrón, Oviedo-Diego, Elias, Vrech, & Peretti, 2019).
Other signals, such as acoustic, are also used by some female spiders to com-
municate their sexual receptivity (e.g., Dutto, Calbacho-Rosa, & Peretti,
2011). In scorpions, both female resistance and quiescence during mating
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can be used by females to communicate their willingness to copulation

(Peretti, 2001).
Can female quiescence be considered a form of communication between
sexes? If so, what are females communicating? In spiders, intersexual commu-
nication has been proposed as a possible selective pressure for the evolution of
female quiescence behavior in only a few species of wolf spiders (Brown,
2006; Costa & González, 2015; Dolejš et al., 2012; Nappi, 1965). In this sense,
it is possible that females become quiescent after evaluating male behaviors
during courtship that may indicate his quality. For example, in the redback
widow Latrodectus hasselti and in the white widow L. pallidus, abdominal
tremulation performed by males can be used to communicate its quality to
females (Uhl & Elias, 2011). In L. hesperus, in particular, Scott et al. (2012)
observed that males that perform web-reduction behavior (i.e., cutting and
bundling sections of the female web and wrapping them) “induce” quiescence
of females more quickly. This male behavior has been suggested to indicate
male quality (Scott et al., 2012). In Schizocosa ocreata, a wolf spider showing
quiescence Level III, substrate-borne signals may function as quality indicators
(Gibson & Uetz, 2008). However, other traits possibly related to male quality,
such as size, weight, and body condition, are not correlated with female qui-
escence (E. Hebets pers. comm.). In the funnel web spider Agelenopsis aperta,
males court females by a combination of abdominal waggle and web flex
tremulations (Singer et al., 2000; Uhl & Elias, 2011). After web courtship,
the male strokes both sides of the female abdomen with his legs and then
she becomes quiescent (Becker et al., 2005; Singer et al., 2000; Singer &
Riechert, 1995). Females may select males based on vibrational signals
(Singer et al., 2000; Singer & Riechert, 1995). Also, males could “induce”
female quiescence by volatile pheromone emission (Becker et al., 2005).
The authors speculate that males would use pedipalp drumming and waving
to direct the pheromones to the female during the precopulatory courtship.
These authors also argue that male pheromones “induce” female quiescence
to decrease the likelihood of sexual cannibalism. This phenomenon (i.e.,
“catalepsy-inducing pheromones,” according to Fischer, 2019) has not been
reported in other spiders and therefore needs further investigation. In sum, in a
communication context, male spiders may use specific behaviors to indicate
their quality to females who can analyze them and respond with quiescence to
indicate receptivity. For more details on male behaviors related to intersexual
communication see Table 2 and Uhl and Elias (2011).
What do we know about the other arachnid orders? In scorpions, female
quiescence has been interpreted in a sexual communication context related
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to female receptivity (Maury, 1968; Peretti, 1993, 1995, 2001; Toscano-

Gadea, 2010). Female quiescence could be considered a period when the
female could screen for male stimulatory behaviors, such as juddering or
other vibratory patterns, accepting his courtship if she is receptive
(Peretti, 2001; Rosin & Shulov, 1963). Studies focused on intersexual com-
munication in solifuges are scarce and the evidence is anecdotal. For exam-
ple, Wharton (1987) suggests that males of the solpugid Metasolpuga picta find
females in their burrows through chemical or acoustic signals. However, the
author does not specify what kind of signals are involved. Also, it has been
observed that males of this species, upon finding a female, dig close to her
burrow entrance to stimulate her appearance (Wharton, 1987). However,
there is no experimental evidence to support these interpretations. What
can female quiescence imply from an intersexual communication perspec-
tive in solifuges? One hypothesis is that quiescence is related to communi-
cating sexual receptivity, as suggested in many species from different families
(Muma, 1966a; Pandram & Sharma, 2018; Punzo, 1997; Wharton, 1987).
Also, as before, it may prevent females from being seriously injured during
copulation (Peretti & Willemart, 2007). Another possibility is that it com-
municates to males that they will not be attacked during sperm transfer.
In the whip spiders Damon variegatus and Phrynus barbadensis, as already
mentioned, females become quiescent only if male courtship is successful.
At this moment, the male deposits the spermatophore, and the female stops
quiescence to begin the sperm transfer phase only when the male quivers
(Alexander, 1961, 1962). In harvestmen, studies are not conclusive, but at
least in Phalangium opilio quiescence may act in an intersexual communica-
tion context and mate choice (Willemart et al., 2006), especially because the
position adopted by the female’s during quiescence is a prerequisite for
sperm transfer (Machado & Macı́as-Ordóñez, 2007).Intersexual communi-
cation becomes more evident if we can observe female movements during
quiescence. For example, the abdominal swiveling behavior in several wolf
spiders (Engelhardt, 1964; Just et al., 2018; Rovner, 1971) or the distention
or protrusion of the genital area in the tarantulas of the family Ischnothelidae
(Coyle & O’Shields, 1990). Particularly, the female swiveling has been
suggested to be produced in cooperation between the sexes (Rovner,
1971; but see Engelhardt, 1964) or actively produced by the female
(Coyle & O’Shields, 1990 for Ischnothelidae; Foelix, 2011 and Just et al.,
2018 for wolf spiders). In both cases, a female participation is implicit, which
may indicate that she is active and capable of perceiving the male behaviors
and responding accordingly.
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Extreme cases of female movements during quiescence occur in some

spiders. Females may interrupt the quiescence during copulation and start
to move even while males are still on the dorsum of her abdomen, as in some
wolf spiders ( Just et al., 2018; Piacentini et al., 2017). In addition, intrasexual
variability in the occurrence or duration of female quiescence would also be
considered evidence of communication or a signal that quiescence is not a
fixed behavior. Moreover, it may indicate that females decide when to inter-
rupt quiescence. This variation in quiescence duration has been reported in
several spiders; in the tarantula Acanthogonatus centralis, for example, just
63.3% of the females remained quiescent after copulation (Copperi, 2018).
Also, in the wolf spider Lycosa thorelli, 77.8% of females walked with males
on their dorsum whereas 5.6% remained quiescent after mating (Costa &
Capocasale, 1984). Other examples indicate that this variation in quiescence
duration or occurrence is more widespread in spiders (M. González pers.
comm.; see also Berendonck, 2003; Costa & González, 2015; Coyle &
O’Shields, 1990; Nappi, 1965; D. Abregú and A.V. Peretti unpub. data).
Although variation in female quiescence duration in spiders seems to be
evident, there is no information to explain it. However, we can formulate a
series of questions that may inspire future investigations. For example, are
females interrupting quiescence to penalize males according to certain char-
acteristics that reflect their poor quality as mating partner? Is the copulatory
courtship of males reflecting unwanted characteristics for the female? Is it pos-
sible that females evaluate males during copulation, and only the most efficient
males benefit from the total duration of female quiescence? Studies on taran-
tulas provide some insights. For example, in Xenonemesia platensis, male cop-
ulatory courtship results in female quiescence during copulation, which can
increase her resistance to subsequent matings (Ferretti et al., 2012, 2013).
Thus, female quiescence could be a male quality-testing behavior in this spe-
cies. In Acanthogonatus centralis (Fig. 2H), the absence of quiescence is related to
copulation interruption by the female. Therefore, quiescence would indicate
to males that females accept to continue copulation, an interpretation that
could be framed in the context of mate choice (Copperi, 2018). In
Grammostola vachoni, female quiescence after copulation may be associated
with a body position for sperm to be stored appropriately (Copperi, 2018).
In this sense, the function would be linked to cryptic female choice.
Finally, an anecdotal but interesting event occurred during a mating experi-
ment in Mecicobothrium thorelli. Costa and Perez-Miles (1998) reported that
they intentionally disturbed a mating male to take a picture, and at that
moment, the female interrupted the quiescence, attacked, and killed the male.
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This could indicate a female rejection of a male with a poor display during
copulation.
Intersexual communication in a mate choice context in solifuges occurs in
Othoes saharae. In this species, female quiescence is generated as a reflex trig-
gered by “male violence or aggression”; the male grabs her with his chelicerae
and wraps the female with pedipalps ( Junqua, 1966). According to the author,
if this “attack” is not made with sufficient “vigor” or “conviction,” which
would be reflecting male’s condition, the female does not become quiescent,
but defends herself from the male ( Junqua, 1966). This example suggests that
female cooperation is necessary for quiescence to occur, and some female eval-
uation of male behavior is present.
In the scorpions Bothriurus bonariensis, Urophonius achalensis, and
U. brachycentrus, it has been reported that females can choose males based
on their body condition. That selection may be reflected in the female
showing quiescence Level III, at least, as a first acceptance filter since it does
not necessarily imply that sperm transfer occurs (Olivero, 2014;
Oviedo-Diego et al., 2021). Additionally, it is possible that male condition
is related to the male juddering behavior, which can be considered a dem-
onstration of his vigor and used by the female as a signal to become quies-
cence (Oviedo-Diego et al., 2021; Peretti, 1993, 1995). Another behavior
possibly related to mate choice is the sexual sting. A recently published arti-
cle in a scorpion species that shows quiescence Level I, M. gertschi, suggests
that during the sexual sting, the female can perceive male attributes, such as
the duration of this behavior, a mechanical signal of piercing, and chemical
signals of the venom, using them to male evaluation (Olguı́n-Perez et al.,
2021). In some buthid scorpions, male walks over the back of female and
remains there for a time (“override”) (Lira, Pordeus, Rego, Iannuzzi, &
Albuquerque, 2018; Outeda-Jorge, 2010), and particularly in Jaguajir rochae
the female remains quiescent during this male behavior (Chantall-Rocha &
Japyassú, 2017). This may be a situation of mutual mate choice during which
males recognize chemical signals and sense female receptivity, while females
evaluate morphological characteristics of the male (Outeda-Jorge, 2010).
In harvestmen, just one report in Phalagium opilio could be related to
female mate choice. In this species, when dealing with smaller males, the
female is more likely to bounce and try to escape than when faced to a larger
male (Willemart et al., 2006). Therefore, females may accept to mate with a
larger male, becoming quiescent. This example reveals female quiescence as
a key element in a complex intersexual communication process in a mate
choice context.
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Finally, one of the cases that best exemplifies quiescence in a mate choice
context in whip spiders is that females only become quiescent if male court-
ship is successful (Alexander, 1961, 1962). Although one of the most studied
aspects of whip spiders is reproductive behavior (Chapin & Hebets, 2016),
there are no data relating courtship parameters to male quality for this order
(E. Hebets pers. comm.).
9.6 Postcopulatory sexual selection

As is well known, postcopulatory sexual selection extends Darwin’s ideas of
precopulatory sexual selection (Andersson, 1994; Darwin, 1871).
Postcopulatory sexual selection also has its own evolutionary mechanisms:
sperm competition (Parker, 1970), cryptic female choice (Eberhard,
1985, 1996), and sexual antagonistic coevolution (Arnqvist & Rowe,
2005). The first two can be understood as equivalent to the evolutionary
mechanisms proposed by Darwin at the precopulatory level, intrasexual
competition or direct competition between males and females’ choice,
respectively (Eberhard, 2015).
Could female quiescence have evolved in the context of postcopulatory
sexual selection? Our literature review found no data to help answering this
question, but still, it is a plausible hypothesis. We can start to analyze this
behavior from a cryptic female choice perspective. In the beetle Tribolium
castaneum, sperm transfer occurs mainly during female quiescence and the dura-
tion of female quiescence is related to increased sperm transfer (Bloch-Qazi,
2003). In turn, quiescence may be a mechanism used by the female to control
sperm transfer and use during fertilization. A similar example can be observed in
the wolf spider Schizocosa malitiosa, a spider showing quiescence Level III, in
which females can control the amount of sperm stored during copulation
(Albo & Costa, 2017). Although it is not clear how such control would be
exerted, the female may eject or kill the spermatozoa (Albo & Costa, 2017;
Eberhard, 1996). As in the example with T. castaneum, sperm transfer occurs
during quiescence, and whatever the cryptic female choice mechanism is used
to control the amount of sperm stored by the females, it should occur during
quiescence (see Albo & Costa, 2017 for more details).
There are further examples in which quiescence may be involved in a
cryptic female choice context. In the redback widow spider Latrodectus
hasselti, paternity is related to copulation duration (Andrade, 1996), although
sperm transfer requires only a fraction of the total copulation time (Snow &
Andrade, 2004). As in the example of S. malitiosa, sperm transfer in L. hasselti
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occurs while female is quiescent, and during this period, the female may bias
male paternity (Snow & Andrade, 2004). In this species, the male has short
bursts of activity during copulation, withdrawing for periods and then
resuming copulation. Is it possible that during this time, the female exerts
control over the transferred spermatozoa?
A key aspect of a cryptic female choice is copulatory courtship. The uni-
versality of copulatory courtship in animals is currently not disputed
(Eberhard & Lehmann, 2019). In spiders showing quiescence, copulatory
courtship takes place during this stage. In several species of the genus
Latrodectus, intersexual interactions can be extensive, and males perform
multiple copulatory courtship behaviors, which may indicate that cryptic
female choice is exerted during this period (Andrade & MacLeod, 2015).
Although empirical data to support these statements are scarce, the role of
quiescence in a framework of cryptic female choice is a topic that deserves
further study.
Turning to sperm competition, is it possible that female quiescence allows
more sperm transfer? In some species males can block the female genital open-
ing with secretions or part of their body or genitalia (i.e., mating plugs). In
Latrodectus spiders mating plugs are originated from the male pedipalp, i.e.,
a broken piece of the embolus (Uhl, Nessler, & Schneider, 2010). In this case,
apparently, the female does not need to be quiescent. However, in Agelena
labyrinthica, a species showing quiescence Level III, the mating plug is gener-
ated by male and female secretions (Uhl et al., 2010 and references therein). In
other species, females can bias paternity by controlling the genital plug forma-
tion (Aisenberg, Barrantes, & Eberhard, 2015; Andrade & MacLeod, 2015;
Schneider, Uhl, & Herberstein, 2015). Can females control the formation
of the mating plug during quiescence?
Finally, in the context of sexually antagonistic coevolution, quiescence as a
response to conflict is against this theory. For example, in Latrodectus spiders,
signs of aggression rather than quiescence should be expected because sexual
size dimorphism is highly female-biased, so that females can easily attack and
even consume males without risk of injury (Andrade & MacLeod, 2015).
10. Limitations of the present study

A first limitation of our review is that those publications that do not
provide a fine-scale description of reproductive interactions may have been
left out of the literature search, producing an underestimation of numbers
of species showing female mating quiescence. This could be the case,
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for instance, of spiders in which a bridal veil has been described. In these
spiders, it is common that the female remains quiescent during this male
behavior (see Section 9.1).
A second limitation is that some examples of female mating quiescence
discussed here are problematic to interpret. This is the case of quiescence that
occurs during the copulatory phase in spiders and harvestmen. The problem
lies in how to differentiate the immobility necessary for sperm transfer from
quiescence itself in the context of the proposed scenarios discussed in
Section 7.1 (see also the hypothesis “Sperm transfer” in Box 1). Although
we chose to include these cases as a type of quiescence of its own (Level
I, Table 6), the available behavioral information is not sufficiently detailed
to allow a solid conclusion.
A third limitation is that the term quiescence is used to describe a con-
tinuum of female behaviors. In one extreme there are quiescent females fully
capable of reacting to external stimuli (i.e., “weak quiescence”), and in
another extreme there are quiescent females unable to react to external stim-
uli (i.e., “strong quiescence”). If we want to understand the evolution of
female quiescence, it would be important to classify the species along this
continuum. The question is what should be considered as a stimulus and
how reliable the female response to this stimulus would be. Although it
would be possible to propose certain criteria, such as manipulating the
female during quiescence and observing her ability to react to the stimulus,
the number of species that have been manipulated, such as spiders (Costa &
Perez-Miles, 1998; Rovner, 1971; Xiao et al., 2015), solifuges (Heymons,
1902; Junqua, 1966), and scorpions (Peretti, 2001), is very small compared
to the total number of species included in our review. As an example, only
two spider species of 84 were manipulated in some way during quiescence.
Thus, any attempt to classify the species along a continuum of female control
over quiescence is hampered by lack of empirical data.
A fourth limitation is that it was often impossible to establish a precise
temporal sequence between male behaviors and female quiescence. This
was due, at least in part, to the absence of detailed ethograms or fine-scale
descriptions of reproductive interactions in most species included in
the review (in some species there may be no consistent sequence). We have
attempted to present the best approximation including all male behaviors
that the authors suggested occurred before quiescence in Tables 2–4. This
will provide a basis for stimulating future studies based on fine observations
of the male-female flow interactions in the context of the quiescence
behavior.
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Finally, we did not test our hypotheses linking quiescence to other vari-
ables in a phylogenetic framework. Although we showed that quiescence is
frequent in some taxa (e.g., the RTA clade in spiders), linking quiescence to
other variables needs to be tested by quantitative evidence. For example, one
question in spiders is whether quiescence results from the lifestyle: has qui-
escence evolved in wandering spiders and then reverted in web builders?
Our survey also indicates that quiescence may have multiple origins, but tests
using phylogenetic evidence are lacking.
11. Conclusions and prospects

The present review shows that female quiescence may be a multi-
causal phenomenon. Concerning the possible hypotheses explaining the
evolution of female quiescence in arachnids, intersexual communication
and mate choice accumulate the highest number of examples in their favor.
However, alternative hypotheses, such as female and male genital complex-
ity or female aggressiveness, are also supported by observations of some few
species. Based on the general results of our review, we present below a list of
general suggestions that can improve our knowledge on female quiescence:
• We highlight the importance of carefully naming behavioral units when
describing sexual interactions (see Peretti & Córdoba-Aguilar, 2007)
because they could be taken as indirect hypotheses that are often related
to authors’ biases. We encourage researchers to avoid the “cultural
boomerang,” defined as the scenario in which cultural preconceptions
are used to interpret nature and then turn around and use cultural depic-
tions of nature to denote what is natural (Ah-King & Ahnesj€ o, 2013;
Ganetz, 2004).
• We emphasize the need to describe objectively and in detail each behavior
mentioned during sexual interactions, in this case, female quiescence. The
publication of behaviors without their pertinent description can lead to
misinterpretations and biased conceptions by the readers (for more details
on how to describe behavior, see Peretti & Córdoba-Aguilar, 2007).
• It is necessary to give relevance to anecdotes, negative results, or outliers
during experimentation since they may represent behavioral variability,
for which alternative interpretations and hypotheses can be postulated
(see Peretti, 2014). For example, examine in detail whether female qui-
escence varies, even subtly, in duration or posture in some of the females
analyzed in the sample (e.g., by individual size or weight).
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• We suggest that researchers publish as supplementary information all

the sexual behaviors observed during their experiences, even if the data
exceeds the original topic of the investigation. We are aware that
female quiescence is more widespread in arachnids than we have been
able to report here. For example, quiescence has been observed in several
species of wolf spiders and ghost spiders (E. Hebets pers. comm.; D. Poy
pers. comm., C. Toscano-Gadea and M. González pers. comm.;
M. Izquierdo pers. obs.) but these data still need to be published.
• We encourage researchers to establish (within a particular genus for
instance) morphological complexity criteria for both male and female
genitalia and evaluate whether this complexity is related to the presence
of female quiescence during mating.
• To easily evaluate whether female quiescence is related to sperm transfer
beyond the transfer systems, in those taxa with direct sperm transfer, it is
possible to assess whether the duration of female quiescence limits the
amount of sperm transferred by males and stored by females. The meth-
odological knowledge to perform spermatozoa counting in various taxa
of arachnids is vast (e.g., Abregú, Peretti, & González, 2019; Cargnelutti,
Calbacho-Rosa, Córdoba-Aguilar, & Peretti, 2018; Snow & Andrade,
2004; Vrech, Oviedo-Diego, Olivero, & Peretti, 2019).
• Although the hypotheses that are best supported are intersexual commu-
nication and mate choice, empirical studies are scarce and assays can be
designed to test these hypotheses formally. For example, it is possible to
modify the quality of males through their body condition (i.e., by alter-
ing their feeding regime) and compare whether females remain quiescent
in equal proportion among both groups of males (well fed vs. poorly fed).
Different parameters of female quiescence, such as latency and duration
variation, could also be examined in conjunction with data on male body
condition.
• Regarding precopulatory and copulatory male courtship and its relation-
ship to female quiescence, it would be useful to assess the natural variability
of male courtship and its effect on female quiescence. Also, male courtship
behaviors can be experimentally modified by suppressing or altering the
structures involved in the male behavior, or by suppressing the sensory
capabilities of the female to perceive such stimuli.
• It is not yet clear why there is variation in the consistency of female qui-
escence. In experiments of successive copulations with different males,
the female’s quiescence consistency can be tested.
• More studies focused on investigating the mechanisms behind quies-
cence are necessary. Specifically, what triggers such behavior in females
ARTICLE IN PRESS
and what happens to them from a physiological perspective. These stud-

ies would allow for better interpretations of the observed patterns, as well
as the possible involvement of the male in female quiescence.
• Finally, our review showed that quiescence occurs in different arachnid
taxa, suggesting multiple phylogenetic origins. If quiescence is a character-
istic of particular clades, such as the RTA clade of spiders, the occurrence
of quiescence may be related to morphological structures or behaviors and
mapped in the phylogeny. We recommend the study of reproductive
behaviors of non-model species or families to illustrate the occurrence
of female quiescence from a more integrative perspective.
Acknowledgments
We are very grateful to Glauco Machado for his invitation to write this review and for his
valuable suggestions on the manuscript. We also thank William G. Eberhard and Eileen
Hebets for critically reviewing previous versions. We are grateful to Sofı́a Copperi,
Gabriel Pompozzi, Leonela Schwerdt, Nelson Ferretti, Cristian Grismado, Andres Porta,
Gustavo Requena, Macarena González, Carlos Toscano-Gadea, Anita Aisenberg, Petr
Dolejš, Luis Piacentini, Hernán Iuri, Dante Poy, Ivan Magalhães, Bruno Buzatto, Luciana
Baruffaldi, Catherine Scott, Sean McCann, Brandi Pessman, and Rogelio Bertani who
have kindly shared data from their personal observations and even videos and photos of
female quiescence in different arachnid species. Financial support was provided by the
Consejo Nacional de Investigaciones Cientı́ficas y Tecnicas (PIP-CONICET), Secretarı́a
de Ciencia y Tecnica de la Universidad Nacional de Córdoba (SECyT-UNC), and
Fondo para la Investigación Cientı́fica y Tecnológica (FONCyT).
References
Abregú, D. A., Peretti, A. V., & González, M. (2019). Male performance and associated costs
in successive sexual encounters in a polygynous web wolf spider. Acta ethologica, 22(3),
175–186. https://doi.org/10.1007/s10211-019-00323-9.
Abregú, D. A., Simian, C., Mattoni, C. I., & Peretti, A. V. (2022). Sexual behavior of
Metaltella iheringi (Keyserling, 1891) (Araneae: Desidae): Sexual patterns, female quies-
cence and comparisons with other spiders. The Journal of Arachnology, 50(1), 81–89.
https://doi.org/10.1636/JoA-S-20-045.
Adler, N. T., & Toner, J. P., Jr. (1986). The effects of copulatory behavior on sperm transport
and fertility in rats. Annals of the New York Academy of Sciences, 474(1), 21–32. https://doi.
org/10.1111/j.1749-6632.1986.tb27995.x.
Ah-King, M. (2007). Sexual selection revisited — Towards a gender-neutral theory and
practice. A response to Vandermassen’s ’sexual selection: A tale of male bias and feminist
Denial’. European Journal of Women’s Studies, 14(4), 341–348. https://doi.org/10.
1177/1350506807081883.
Ah-King, M. (2011). Female sexual selection in light of the Darwin–Bateman paradigm.
Behavioral Ecology, 22(6), 1142–1143. https://doi.org/10.1093/beheco/arr109.
Ah-King, M., & Ahnesj€ o, I. (2013). The “sex role” concept: An overview and evaluation.
Evolutionary Biology, 40(4), 461–470. https://doi.org/10.1007/s11692-013-9226-7.
Aisenberg, A., & Costa, F. G. (2005). Females mated without sperm transfer maintain high
sexual receptivity in the wolf spider Schizocosa malitiosa. Ethology, 111(6), 545–558.
https://doi.org/10.1111/j.1439-0310.2005.01077.x.
ARTICLE IN PRESS
Aisenberg, A., Barrantes, G., & Eberhard, W. G. (2015). Post-copulatory sexual selection in
two tropical orb-weaving Leucauge spiders. In Cryptic female choice in arthropods
(pp. 79–108). Cham: Springer.
Aisenberg, A., Estramil, N., González, M., Toscano-Gadea, C. A., & Costa, F. G. (2008).
Silk release by copulating Schizocosa malitiosa males (Araneae, Lycosidae): A bridal veil.
The Journal of Arachnology, 36(1), 204–206. https://doi.org/10.1636/St07-12SC.1.
Albo, M. J., & Costa, F. G. (2017). Female wolf spiders exert cryptic control drastically reduc-
ing ejaculate size. Ethology, 123(9), 659–666.
Alexander, A. J. (1957). The courtship and mating of the scorpion, Opisthophthalmus
latimanus. Proceedings of the Zoological Society of London, 128(4), 529–544. https://doi.
org/10.1111/j.1096-3642.1957.tb00274.x.
Alexander, A. J. (1959). Courtship and mating in the buthid scorpions. Proceedings of the
Zoological Society of London, 133, 145–169. https://doi.org/10.1111/j.1469-7998.1959.
tb05557.x.
Alexander, A. J. (1961). Courtship and mating in amblypygids (Pedipalpi, Arachnida).
Proceedings of the Zoological Society of London, 138 (3), 379-383). https://doi.org/10.
1111/j.1469-7998.1962.tb05705.x.
Alexander, A. J. (1962). Biology and behavior of Damon variegatus Perty of South Africa
and Admetus barbadensis Pocock of Trinidad, WI (Arachnida, Pedipalpi). Zoologica,
47, 25–37.
Alvarado-Castro, J. A., & Jimenez, M. L. (2011). Reproductive behavior of Homalonychus
selenopoides (Araneae: Homalonychidae). The Journal of Arachnology, 39(1), 118–127.
https://doi.org/10.1636/Hi10-35.1.
Amitai, P., Levy, G., & Shulov, A. (1962). Observations on mating in a solifugid Galeodes
sulfuripes Roewer. Bulletin of the Research Council of Israel, 3, 156.
Andersson, M. (1994). Sexual selection. Princeton, New Jersey: Princeton Univ. Press.
Andrade, M. C. (1996). Sexual selection for male sacrifice in the Australian redback spider.
Science, 271(5245), 70–72.
Andrade, M. C. (1998). Female hunger can explain variation in cannibalistic behavior despite
male sacrifice in redback spiders. Behavioral Ecology, 9(1), 33–42. https://doi.org/10.
1093/beheco/9.1.33.
Andrade, M. C., & MacLeod, E. C. (2015). Potential for CFC in black widows (genus
Latrodectus): Mechanisms and social context. In Cryptic female choice in arthropods
Arnqvist, G., & Rowe, L. (2005). Sexual conflict. New Jersey: Princeton University Press.
Azevedo, G. H., Griswold, C. E., & Santos, A. J. (2018). To complicate or to simplify?
Phylogenetic tests of complexity trends and genital evolution in ground spiders
(Araneae: Dionycha: Gnaphosidae). Zoological Journal of the Linnean Society, 184(3),
673–694. https://doi.org/10.1093/zoolinnean/zly016.
Ballesteros, J. A., Santibáñez-López, C. E., Baker, C. M., Benavides, L. R., Cunha, T. J.,
Gainett, G., et al. (2022). Comprehensive species sampling and sophisticated algorithmic
approaches refute the monophyly of Arachnida. Molecular Biology and Evolution, 39(2),
msac021. https://doi.org/10.1093/molbev/msac021.
Becker, E., Riechert, S., & Singer, F. (2005). Male induction of female quiescence/catalepsis
during courtship in the spider, Agelenopsis aperta. Behaviour, 142, 57–70. https://doi.org/
10.1163/1568539053627767.
Bennett, R. G. (1987). Systematics and natural history of Wadotes (Araneae, Agelenidae).
Journal of Arachnology, 91–128. http://www.jstor.org/stable/3705512.
Benton, T. G. (1993). Courtship behaviour of the scorpion, Euscorpius flavicaudis. Bulletin of
the British Arachnological Society, 9(5), 137–141.
Benton, T. G. (2001). Reproductive biology. In P. Brownell, & G. A. Polis (Eds.), Scorpion
biology and research (pp. 278–301). Oxford University Press: New york.
ARTICLE IN PRESS
Berendonck, B. (2003). Reproductive strategies in Latrodectus revivensis (Araneae; Theridiidae):

Functional morphology and sexual cannibalism. (PhD dissertation, D€ usseldorf: Heinrich
Heine University D€ usseldorf.
Berland, L. (1932). Les Arachnides (Biologie, Systematique). Encyclopedic entomologique A, 16,
52. Paris: Lechevalier.
Bertani, R., Fukushima, C. S., & da Silva Júnior, P. I. (2008). Mating behavior of Sickius
longibulbi (Araneae, Theraphosidae, Ischnocolinae), a spider that lacks spermathecae.
The Journal of Arachnology, 36(2), 331–335. https://doi.org/10.1636/CSt07-100.1.
Bilde, T., Tuni, C., Elsayed, R., Pekár, S., & Toft, S. (2006). Death feigning in the face of
sexual cannibalism. Biology Letters, 2(1), 23–25. https://doi.org/10.1636/CSt07-100.1.
Bird, T. L., Wharton, R. R. A., & Prendini, L. (2015). Cheliceral morphology in Solifugae
(Arachnida): Primary homology, terminology, and character survey. Bulletin of the
American Museum of Natural History, no. 394.
Birkhead, T. R., & Pizzari, T. (2002). Postcopulatory sexual selection. Nature Reviews
Genetics, 3, 262–273. https://doi.org/10.1038/nrg774.
Bloch-Qazi, M. C. (2003). A potential mechanism for cryptic female choice in a flour beetle.
Journal of Evolutionary Biology, 16, 170–176.
Brantley, J. D. 2017 [Jason D. Brantley] (26 oct 2017). egyptian deathstalker mating 1/2
[Video]. https://www.youtube.com/watch?v¼yfMMV4S-4t4.
Breene, R. G., & Sweet, M. H. (1985). Evidence of insemination of multiple females by
the male black widow spider, Latrodectus mactans (Araneae, Theridiidae). Journal of
Arachnology, 331–335.
Briceño, R. D., & Bonilla, F. (2009). Substrate vibrations in the scorpion Centruroides
margaritatus (Scorpiones: Buthidae) during courtship. Revista de Biologıá Tropical, 57,
267–274.
Bristowe, W. S. (1931). 68. The mating habits of spiders: A second supplement, with the
description of a new Thomisid from Kritkatau. In Proceedings of the Zoological Society of
London (pp. 1401–1412). Oxford, UK: Blackwell Publishing Ltd. https://doi.org/10.
1111/j.1096-3642.1931.tb01070.x. Vol. 101., No. 4.
Brown, C. A. (2006). Observations on courtship and copulation of the wolf spider Rabidosa
santrita (Araneae, Lycosidae). The Journal of Arachnology, 34(2), 476–479. https://doi.org/
10.1636/S05-55.1.
Bruce, J. A., & Carico, J. E. (1988). Silk use during mating in Pisaurina mira (Walckenaer)
(Araneae, Pisauridae). The Journal of Arachnology, 1–4. https://www.jstor.org/stable/
3705799.
ucherl, W. (1952). Instintos maternais nas aranhas brasileiras. Dusenia, 3, 57–74.
B€
Burger, M., Michalik, P., Graber, W., Jacob, A., Nentwig, W., & Kropf, C. (2006).
Complex genital system of a haplogyne spider (Arachnida, Araneae, Tetrablemmidae)
indicates internal fertilization and full female control over transferred sperm. Journal of
Morphology, 267(2), 166–186. https://doi.org/10.1002/jmor.10394.
Burger, M., Nentwig, W., & Kropf, C. (2003). Complex genital structures indicate
cryptic female choice in a haplogyne spider (Arachnida, Araneae, Oonopidae,
Gamasomorphinae). Journal of Morphology, 255(1), 80–93. https://doi.org/10.1002/
jmor.10048.
Capocasale, R. M., & Perez-Miles, F. (1990). Behavioural ecology of Acanthogonatus
tacuariensis (Perez & Capocasale)(Araneae, Nemesiidae). Studies on Neotropical Fauna
and Environment, 25(1), 41–47. https://doi.org/10.1080/01650529009360800.
Cargnelutti, F., Calbacho-Rosa, L., Córdoba-Aguilar, A., & Peretti, A. V. (2018). Patterns of
sperm transfer behavior in a pholcid spider with two distinct copulatory phases. Journal of
Insect Behavior, 31(6), 616–628.
Carranza, J. (2010). Sexual selection and the evolution of evolutionary theories. Animal
Behaviour, 79(3), e5. https://doi.org/10.1016/j.anbehav.2009.08.010.
ARTICLE IN PRESS
Catley, K. M. (1993). Courtship, mating and postoviposition behaviour of Hypochilus pococki

Platnick (Araneae, Hypochilidae). Memoirs of the Queensland Museum, 33(46), 9–474.
Chantall-Rocha, S., & Japyassú, H. F. (2017). Diffuse resistance courtship in the scorpion
Rhopalurus rochai (Scorpiones: Buthidae). Behavioural Processes, 135, 45–55. https://doi.
org/10.1016/j.beproc.2016.11.017.
Chapin, K. J., & Hebets, E. A. (2016). The behavioral ecology of amblypygids. Journal of
Arachnology, 1–14. https://doi.org/10.1636/V15-62.1.
Chiarle, A., Kronestedt, T., & Isaia, M. (2013). Courtship behavior in European species of
the genus Pardosa (Araneae, Lycosidae). Journal of Arachnology, 41(2), 108–125.
Cloudsley-Thompson, J. L. (1961). Observations on the natural history of the ‘camel-spider’,
Galeodes arabs C. L. Koch (Solifugae: Galeodidae) in the Sudan. Entomological Monthly
Magazine, 97, 145–152.
Cloudsley-Thompson, J. L. (1967). Reproduction in Solifugae. Turtox News, 45, 212–215.
Cloudsley-Thompson, J. L. (1977). Adaptational biology of Solifugae (Solpugida). Bulletin of
the British Arachnological Society, 4, 61–71.
Cohen, C. (2010). Darwin on woman. Comptes Rendus Biologies, 333, 157–165. https://doi.
org/10.1016/j.crvi.2009.12.003.
Copperi, S. (2018). Biologıá reproductiva de arañas migalomorfas (Araneae, Mygalomorphae),
patrones y mecanismos de selección sexual. (PhD dissertation, Argentina: Universidad
Nacional del Sur.
Costa, F. G. (1979). Análisis de la cópula y de la actividad post-copulatoria de Lycosa malitiosa
(Tullgren) (Araneae, Lycosidae). Revista Brasileira de Biologia, 39(2), 361–376.
Costa, F. G., & Capocasale, R. M. (1984). La producción de ootecas de Lycosa malitiosa
Tullgren (Araneae: Lycosidae). Importancia de la muda de maturacion sobre la primera
oviposición. Aracnologia, 2, 1–8.
Costa, F. G., & González, M. (2015). El comportamiento sexual de Hogna bivittata y Hogna
sp.(Araneae, Lycosidae), dos especies simpátricas del sur de Uruguay. Boletıń de la Sociedad
Zoológica del Uruguay, 24, 91–102.
Costa, F. G., & Perez-Miles, F. (1998). Behavior, life cycle and webs of Mecicobothrium thorelli
(Araneae, Mygalomorphae, Mecicobothriidae). Journal of Arachnology, 317–329. https://
www.jstor.org/stable/3706238.
Costa-Schmidt, L. E., Albo, M. J., Bollatti, F., Cargnelutti, F., Calbacho-Rosa, L.,
Copperi, S., et al. (2017). Sexual selection in neotropical spiders: Examples from selected
groups. In C. Viera, & M. Gonzaga (Eds.), Behaviour and ecology of spiders (pp. 303–350).
Cham, Switzerland: Springer. https://doi.org/10.1007/978-3-319-65717-2_12.
Covelo de Zolessi, L. (1956). Observaciones sobre el comportamiento sexual de Bothriurus
bonariensis (Koch) (Scorpiones, Bothriuridae). Boletıń-Facultad de Agronomıá. Uruguay:
Universidad de la República. 35–pp12.
Coyle, F. A. (1985). Observations on the mating behaviour of the tiny mygalomorph spider,
Microhexura montivaga Crosby & Bishop (Araneae, Dipluridae). Bulletin of the British
Arachnological Society, 6, 328–330.
Coyle, F. A. (1986). Courtship, mating, and the function of male-specific structures in the
mygalomorph spider genus Euagrus (Araneae, Dipluridae). Proceedings of the Ninth
International Congress of Arachnology, 33–38.
Coyle, F. A., & O’Shields, T. C. (1990). Courtship and mating behavior of Thelochoris karsh
(Araneae, Dipluridae), an African funnel web spider. Journal of Arachnology, 18, 281–296.
https://www.jstor.org/stable/3705434.
Cuthbertson, D. R. (1984). Catalepsy and phoresy in pseudoscorpions. Newsletter of the British
Arachnological Society, 39, 3.
Darwin, C. (1871). The descent of man and selection in relation to sex. London, England: John
Murray.
ARTICLE IN PRESS
Dharmarathne, W. D. S. C., & Herberstein, M. E. (2022). Limitations of sperm transfer in the

complex reproductive system of spiders. Biological Journal of the Linnean Society, 135(3),
417–428. https://doi.org/10.1093/biolinnean/blab158.
Dolejš, P., Kubcová, L., & Buchar, J. (2010). Courtship, mating, and cocoon maintenance of
Tricca lutetiana (Araneae: Lycosidae). The Journal of Arachnology, 38(3), 504–510. https://
doi.org/10.1636/Hi09-29.1.
Dolejš, P., Kubcová, L., & Buchar, J. (2012). Reproduction of Arctosa alpigena lamperti
(Araneae: Lycosidae)–where, when, how, and how long? Invertebrate Reproduction and
Development, 56(1), 72–78. https://doi.org/10.1080/07924259.2011.617072.
Domı́nguez, K., & Jimenez, M. L. (2005). Mating and self-burying behavior of Homalonychus
theologus Chamberlin (Araneae, Homalonychidae) in Baja California sur. The Journal of
Arachnology, 33(1), 167–174. https://doi.org/10.1636/M03-4.
Dutto, M. S., Calbacho-Rosa, L., & Peretti, A. V. (2011). Signalling and sexual conflict:
Female spiders use stridulation to inform males of sexual receptivity. Ethology,
117(11), 1040–1049. https://doi.org/10.1111/j.1439-0310.2011.01957.x.
Eason, R. R. (1969). Life history and behavior of Pardosa lapidicina Emerton (Araneae:
Lycosidae). Journal of the Kansas Entomological Society, 339–360. https://www.jstor.org/
stable/25083804.
Eberhard, W. G. (1985). Sexual selection and animal genitalia. In Sexual Selection and Animal
Genitalia Harvard University Press.
Eberhard, W. G. (1990). Function and phylogeny of spider webs. Annual Review of Ecology
and Systematics, 341–372. https://www.jstor.org/stable/2097029.
Eberhard, W. G. (1996). Female control: Sexual selection by cryptic female choice. Princeton
University Press: Princeton, New Jersey.
Eberhard, W. G. (2009). Postcopulatory sexual selection: Darwin’s omission and its
consequences. Proceedings of the National Academy of Sciences, 106(Supplement_1),
10025–10032.
Eberhard, W. G. (2015). Cryptic female choice and other types of post-copulatory sexual
selection. In A. V. Peretti, & A. Aisenberg (Eds.), Cryptic female choice in arthropods
Eberhard, W. G., & Huber, B. A. (2010). Spider genitalia: Precise maneuvers with a
numb structure in a complex lock. In J. L. Leonard, & A. Córdoba-Aguilar (Eds.),
Evolution of primary sexual characters in animals (pp. 249–284). Oxford, UK: Oxford
University Press.
Eberhard, W. G., & Lehmann, G. U. (2019). Demonstrating sexual selection by cryptic
female choice on male genitalia: What is enough? Evolution, 73(12), 2415–2435.
Eisner, T., & Meinwald, J. (1995). The chesmistry of sexual selection. Proceedings of the
National Academy of Sciences, 92, 50–55. https://doi.org/10.1073/pnas.92.1.50.
Elgar, M. A. (1991). Sexual cannibalism, size dimorphism, and courtship behavior in
orb-weaving spiders (Araneidae). Evolution, 45(2), 444–448. https://doi.org/10.2307/
2409679.
Elgar, M. A. (1992). Sexual cannibalism in spiders and other invertebrates. In M. A. Elgar, &
B. J. Crespi (Eds.), Cannibalism: Ecology and evolution among diverse taxa (pp. 129–156).
Oxford: Oxford University Press.
Engelhardt, W. (1964). Die mitteleueopaischen arten der gattung Trochosa C. L. Koch, 1848
(Araneae, Lycosidae). Morphologie, Chemotaxonomie, Biologie, Autokologie.
Zeitschrift f€ €
ur Morphologie und Okologie der Tiere, 54, 219–392. https://doi.org/10.1007/
BF00390678.
Fedina, T. Y. (2007). Cryptic female choice during spermatophore transfer in Tribolium
castaneum (Coleoptera: Tenebrionidae). Journal of Insect Physiology, 53, 93–98. https://
doi.org/10.1016/j.jinsphys.2006.10.011.
ARTICLE IN PRESS
Ferretti, N., Copperi, S., Schwerdt, L., & Pompozzi, G. (2015). First record of Vitalius roseus
(Mello-Leitão, 1923) (Araneae: Theraphosidae: Theraphosinae) in Argentina:
Distribution map, natural history, and sexual behaviour. Arachnology, 16(7), 241–243.
https://doi.org/10.13156/arac.2015.16.7.241.
Ferretti, N. E., & Ferrero, A. A. (2008). Courtship and mating behavior of Grammostola schulzei
(Schmidt 1994) (Araneae, Theraphosidae), a burrowing tarantula from Argentina. The
Journal of Arachnology, 36(2), 480–483. https://doi.org/10.1636/CSt08-09.1.
Ferretti, N., Pompozzi, G., Copperi, S., González, A., & Perez-Miles, F. (2013). Sexual
behaviour of mygalomorph spiders: When simplicity becomes complex; an update of
the last 21 years. Arachnology, 16(3), 85–93. https://doi.org/10.13156/100.016.0303.
Ferretti, N., Pompozzi, G., Copperi, S., Perez-Miles, F., & González, A. (2012). Copulatory
behavior of Microstigmatidae (Araneae: Mygalomorphae): A study with Xenonemesia
platensis from Argentina. Journal of Arachnology, 252–255. https://doi.org/10.1636/
Hi11-74.1.
Ferretti, N., Pompozzi, G., & Perez-Miles, F. (2011). Sexual behavior of Acanthogonatus
centralis (Araneae: Mygalomorphae: Nemesiidae) from Argentina, with some notes on
their burrows. The Journal of Arachnology, 39(3), 533–536. https://doi.org/10.1636/
Hi09-72.1.
Fischer, A. (2019). Chemical communication in spiders–a methodological review. The
Journal of Arachnology, 47(1), 1–27. https://doi.org/10.1636/0161-8202-47.1.1.
Fisher, R. A. (1930). The genetical theory of natural selection. New York: Oxford University
Press.
Foelix, R. (2011). Biology of spiders. New York: Oxford University Press.
Fowler-Finn, K. D., Triana, E., & Miller, O. G. (2014). Mating in the harvestman Leiobunum
vittatum (Arachnida: Opiliones): From premating struggles to solicitous tactile engage-
ment. Behaviour, 151(1213), 1663–1686. https://www.jstor.org/stable/2452644.
Francke, O. F. (1979). Observations on the reproductive biology and life history of
Megacormus gertschi Diaz (Scorpiones: Chactidae; Megacorminae). Journal of
Arachnology, 223–230. https://www.jstor.org/stable/3705146.
Fraser, J. B. (1987). Courtship and copulatory behavior of the funnel-web spider, Hololena
adnexa (Araneae, Agelenidae). Journal of Arachnology, 257–262. https://www.jstor.org/
stable/3705736.
Ganetz, H. (2004). Familiar beasts: Nature, culture and gender in wildlife films on television.
Nordicom Review, 25, 197–213. https://doi.org/10.1515/nor-2017-0281.
Gerhardt, U. (1921). Vergleichende Studien u €ber die Morphologie des m€annlichen
Tasters und die Biologie der Kopulation der Spinnen. Archiv f€ ur Naturgeschichte, 87(4),
78–247.
Gerhardt, U. (1927). Neue biologische Untersuchungen an einheimischen und
ausl€andischen Spinnen. Zeitschrift f€ €
ur Morphologie und Okologie der Tiere, 8(1/2), 96–186.
Gerhardt, U. (1928). Biologische Studien an Griechischen, Corsischen und Deutschen
Spinnen. Zeitschrift f€ €
ur Morphologie und Okologie der Tiere, 10, 576–675.
Gerhardt, U. (1933). Neue Untersuchungen zur Sexualbiologie der Spinnen, insbesondere
an Arten der Mittelmeerl€ander und der Tropen. Zeitschrift f€ €
ur Morphologie und Okologie der
Tiere, 27, 1–75.
Gering, R. L. (1953). Structure and function of the genitalia in some American agelenid
spiders. Smithsonian miscellaneous. Collections.
Gibson, J. S., & Uetz, G. W. (2008). Seismic communication and mate choice in wolf spiders:
Components of male seismic signals and mating success. Animal Behaviour, 75(4),
1253–1262. https://doi.org/10.1016/j.anbehav.2007.09.026.
González, M., & Toscano-Gadea, C. A. (2020). In Cruce de caminos: comportamiento sexual de la
araña lobo sudamericana Aglaoctenus oblongus (Araneae, Lycosidae). (presentación oral) VI
Congreso Latinoamericano de Aracnologıá (VICLA), virtual: 137.
ARTICLE IN PRESS
Griswold, C. E. (1993). Investigations into the phylogeny of the lycosoid spiders and their kin
(Arachnida: Araneae, Lycosoidea). Smithsonian Contributions to Zoology, 539, 1–39.
Hammerstein, P., & Riechert, S. E. (1988). Payoffs and strategies in territorial contests: ESS
analyses of two ecotypes of the spider Agelenopsis aperta. Evolutionary Ecology, 2(2),
115–138. https://doi.org/10.1007/BF02067272.
Hansen, L. S., Gonzales, S. F., Toft, S., & Bilde, T. (2008). Thanatosis as an adaptive male
mating strategy in the nuptial gift–giving spider Pisaura mirabilis. Behavioral Ecology, 19(3),
546–551. https://doi.org/10.1093/beheco/arm165.
Hebets, E. A., & Anderson, A. (2018). Using cross-disciplinary knowledge to facilitate
advancements in animal communication and science communication research. Journal
of Experimental Biology, 221(18), jeb179978. https://doi.org/10.1242/jeb.179978.
Hebets, E. A., Barron, A. B., Balakrishnan, C. N., Hauber, M. E., Mason, P. H., &
Hoke, K. L. (2016). A systems approach to animal communication. Proceedings of the
Royal Society B: Biological Sciences, 283(1826), 20152889. https://doi.org/10.1098/
rspb.2015.2889.
von Helversen, O. (1976). Gedanken zur Evolution der Paarungsstellung bei den Spinnen
(Arachnida: Araneae). Entomologica germanica, 13–28. https://doi.org/10.1127/entom.
germ/3/1976/13.
Herberstein, M. E., & Tso, I. M. (2011). Spider webs: Evolution, diversity and plasticity. In
M. E. Herberstein (Ed.), Spider behaviour: Flexibility and versatility (pp. 57–98).
Cambridge, United Kingdom: Cambridge University Press. https://doi.org/10.1017/
CBO9780511974496.004.
Heymons, R. (1902). Biologische Beobachtungen an asiatischen Solifugen, nebst Beitragen zur
Systematik derselben. Verlag der K€ onigl. Akademie der Wissenschaften, in Commission
bei Georg Reimer.
Hormiga, G., Scharff, N., & Coddington, J. A. (2000). The phylogenetic basis of sexual size
dimorphism in orb-weaving spiders (Araneae, Orbiculariae). Systematic Biology, 49(3),
435–462. https://doi.org/10.1080/10635159950127330.
Hrušková-Martišová, M., Pekár, S., & Bilde, T. (2010). Coercive copulation in two sexually
cannibalistic camel-spider species (Arachnida: Solifugae). Journal of Zoology, 282(2),
91–99. https://doi.org/10.1111/j.1469-7998.2010.00718.x.
Hrušková-Martišová, M., Pekár, S., & Cardoso, P. (2010). Natural history of the Iberian
solifuge Gluvia dorsalis (Solifuges: Daesiidae). The Journal of Arachnology, 38(3),
466–474. https://doi.org/10.1636/Hi09-104.1.
Hrušková-Martišová, M., Pekár, S., & Gromov, A. (2007). Biology of Galeodes caspius sub-
fuscus (Solifugae, Galeodidae). Journal of Arachnology, 546–550. https://doi.org/10.1636/
JOA/06-64.1.
Hrušková-Martišová, M., Pekár, S., & Gromov, A. (2008). Analysis of stridulation in
Solifuges (Arachnida: Solifugae). Journal of Insect Behavior, 21, 440–449. https://doi.
org/10.1007/s10905-008-9141-4.
Huber, B. A. (1994). Copulatory mechanics in the funnel-web spiders Histopona torpida and
Textrix denticulata (Agelenidae, Araneae). Acta Zoologica, 75(4), 379–384. https://doi.org/
10.1111/j.1463-6395.1994.tb00974.x.
Huber, B. A. (1995a). The retrolateral tibial apophysis in spiders - shaped by sexual selection?
Zoological Journal of the Linnean Society, 113, 151–163. https://doi.org/10.1111/j.1096-
3642.1995.tb00931.x.
Huber, B. A. (1995b). Genital morphology and copulatory mechanics in Anyphaena accentuata
(Anyphaenidae) and Clubiona pallidula (Clubionidae: Araneae). Journal of Zoology, 235,
689–702.
Huber, B. A. (1998). Spider reproductive behaviour: A review of Gerhardt’s work from
1911–1933, with implications for sexual selection. Bulletin of the British Arachnological
Society, 11, 81–91.
ARTICLE IN PRESS
Huber, B. A. (2005). Sexual selection research on spiders: Progress and biases. Biological
Reviews, 80(3), 363–385. https://doi.org/10.1017/S1464793104006700.
Jiao, G. B., & Zhu, M. S. (2010). Courtship and mating of Scorpiops luridus Zhu Lourenço &
Qi, 2005 (Scorpiones: Euscorpiidae) from Xizang province, China. Journal of Venomous
Animals and Toxins Including Tropical Diseases, 16(1), 155–165. https://doi.org/10.1590/
S1678-91992010005000010.
Jones, T. C., Akoury, T. S., Hauser, C. K., Neblett, M. F., Linville, B. J., Edge, A. A., et al.
(2011). Octopamine and serotonin have opposite effects on antipredator behavior in the
orb-weaving spider, Larinioides cornutus. Journal of Comparative Physiology A, 197(8),
819–825. https://doi.org/10.1007/s00359-011-0644-7.
Junqua, C. (1962). Biologie-Donnees sur la Reproduction dun Solifuge-Othoes Saharae
Panouse. Comptes rendus hebdomadaires des seances del academie des sciences, 255(20), 2673.
Junqua, C. (1966). Recherches biologiques et hystrophysiologiques sur un Solifuge saharien : Othoes
saharae panouse (p. 124p). Paris: Museum national d’Histoire naturelle (Memoires du
Museum national d’Histoire naturelle, Ser. A – Zoologie (1950–1992); 43).
Just, P., Opatova, V., & Dolejš, P. (2018). Does reproductive behaviour reflect phylogenetic
relationships? An example from central European Alopecosa wolf spiders (Araneae:
Lycosidae). Zoological Journal of the Linnean Society, 185(4), 1039–1056. https://doi.
org/10.1093/zoolinnean/zly060.
Kaston, B. J. (1970). Comparative biology of American black widow spiders. Transactions of
the San Diego Society of Natural History, 16(3), 33–82.
Kirchmair, G., & Raspotnig, G. (2021). Mating behavior of Dactylochelifer latreillii latreillii
(Pseudoscorpiones: Cheliferidae): A quantitative study. The Journal of Arachnology,
49(2), 198–204. https://doi.org/10.1636/JoA-S-20-057.
Knoflach, B. (2000). Copulation and emasculation in Echinotheridion gibberosum (Kulczynski,
1899) (Araneae, Theridiidae). European arachnology, 139–144.
Knoflach, B., & Benjamin, S. P. (2003). Mating without sexual cannibalism in Tidarren
sisyphoides (Araneae, Theridiidae). The Journal of Arachnology, 31(3), 445–448. https://
doi.org/10.1636/S02-56.
Knoflach, B., & van Harten, A. (2002). The genus Latrodectus (Araneae: Theridiidae) from
mainland Yemen, the Socotra Archipelago and adjacent countries. Fauna of Arabia,
19, 321–362.
Kuntner, M., Cheng, R. C., Kralj-Fišer, S., Liao, C. P., Schneider, J. M., & Elgar, M. A.
(2016). The evolution of genital complexity and mating rates in sexually size dimorphic
spiders. BMC Evolutionary Biology, 16(1), 1–9. https://doi.org/10.1186/s12862-016-
0821-y.
Lira, A. F., Pordeus, L. M., Rego, F. N., Iannuzzi, K., & Albuquerque, C. M. (2018). Sexual
dimorphism and reproductive behavior in the Brazilian scorpion Tityus pusillus
(Scorpiones, Buthidae). Invertebrate Biology, 137(3), 221–230. https://doi.org/10.1111/
ivb.12221.
Machado, G., & Macı́as-Ordóñez, R. (2007). Reproduction. In R. Pinto-da-Rocha,
G. Machado, & G. Giribet (Eds.), Harvestmen: The biology of Opiliones (pp. 414–454).
Cambridge: Harvard University Press.
Machado, G., & Pomini, A. M. (2008). Chemical and behavioral defenses of the neotropical
harvestman Camarana flavipalpi (Arachnida: Opiliones). Biochemical Systematics and
Ecology, 36(5–6), 369–376. https://doi.org/10.1016/j.bse.2007.12.006.
Machado, G., Requena, G. S., Toscano-Gadea, C., Stanley, E., & Macı́as-Ordóñez,
R. (2015). Male and female mate choice in harvestmen: General patterns and inferences
on the underlying processes. In A. V. Peretti, & A. Aisenberg (Eds.), Cryptic female choice
in arthropods: Patterns, mechanisms and prospects (pp. 169–201). Cham, Switzerland:
Springer. https://doi.org/10.1007/978-3-319-17894-3_7.
ARTICLE IN PRESS
Macı́as-Ordóñez, R., Machado, G., Perez-González, A., & Shultz, J. W. (2010). Genitalic
evolution in Opiliones. In J. Leonard, & A. Córdoba-Aguilar (Eds.), The evolution of
primary sexual characters in animals (pp. 285–306). New York: Oxford University Press.
Magalhaes, I. L., & Ramı́rez, M. J. (2022). Phylogeny and biogeography of the ancient spider
family Filistatidae (Araneae) is consistent both with long-distance dispersal and vicariance
following continental drift. Cladistics, 38(5), 538–562. https://doi.org/10.1111/cla.
12505.
Matthews, M., & Adler, N. T. (1977). Facilitative and inhibitory influences of reproductive
behavior on sperm transport in rats. Journal of Comparative and Physiologial Psychology, 91,
727–741. https://doi.org/10.1037/h0077364.
Matthews, M., & Adler, N. T. (1978). Systematic interrelationship of mating, vaginal plug
position, and sperm transport in the rat. Physiology & Behavior, 20, 303–309. https://doi.
org/10.1016/0031-9384(78)90224-X.
Matthews, M., & Adler, N. T. (1979). Relative efficiency of sperm transport and number of
sperm ejaculated in the female rat. Biology of Reproduction, 20, 540–544. https://doi.org/
10.1095/biolreprod20.3.540.
Maury, E. A. (1968). Aportes al conocimiento de los escorpiones de la República Argentina.
I. Observaciones biológicas sobre Urophonius brachycentrus (Thorell, 1877) (Bothriuridae).
Physis, 27(75), 407–418.
Maury, E. A. (1980). Dos nuevos Pseudocleobis de la Patagonia (Arachnida, Solifugae,
Ammotrechidae). Physis Sección C, 39, 41–43.
Maury, E. A. (1984). Las familias de solı́fugos americanos y su distribución geográfica
(Arachnida, Solifugae). Physis Sección C, 42, 73–80.
Maya-Morales, J., Jimenez, M. L., Murugan, G., & Palacios-Cardiel, C. (2017). Four new
genera of funnel-web spiders (Araneae: Agelenidae) from the Baja California Peninsula in
Mexico. The Journal of Arachnology, 45(1), 30–66. https://doi.org/10.1636/JoA-S-16-
024.1.
McCrone, J. D., & Levi, H. W. (1964). North American widow spiders of the Latrodectus
curacaviensis group (Araneae: Theridiidae). Psyche, 71(1), 12–27. https://doi.org/
10.1155/1964/86469.
McLean, C. J., Garwood, R. J., & Brassey, C. A. (2018). Sexual dimorphism in the Arachnid
orders. PeerJ, 6, e5751. https://doi.org/10.7717/peerj.5751.
Metter, D. E. (1964). A morphological and ecological comparison of two populations of the
tailed frog, Ascaphus truei Stejneger. Copeia, 181–195.
Miller, G. L., & Miller, P. R. (1987). Life cycle and courtship behavior of the burrowing
wolf spider Geolycosa turricola (Treat) (Araneae, Lycosidae). The Journal of Arachnology,
385–394. https://www.jstor.org/stable/3705854.
Miyatake, T., Katayama, K., Takeda, Y., Nakashima, A., Sugita, A., & Mizumoto, M.
(2004). Is death–feigning adaptive? Heritable variation in fitness difference of
death–feigning behaviour. Proceedings of the Royal Society of London. Series B: Biological
Sciences, 271(1554), 2293–2296. https://doi.org/10.1098/rspb.2004.2858.
Montgomery, T. H. (1903). Studies on the habits of spiders, particularly those of the mating
period. Proceedings of the Academy of Natural Sciences of Philadelphia, 59–149.
Moore, A. J., & Wilson, P. (1993). The evolution of sexually dimorphic earwig forceps:
Social interactions among adults of the toothed earwig, Vostox apicedentatus. Behavioral
Ecology, 4, 40–48. https://doi.org/10.1093/beheco/4.1.40.
Mora, G. (1990). Parental care in a neotropical harvestman, Zygopachylus albomarginis
(Arachnida: Gonyleptidae). Animal Behaviour, 39, 582–593. https://doi.org/10.1016/
S0003-3472(05)80425-7.
Muma, M. H. (1966a). Mating behavior in the solpugid genus Eremobates Banks. Animal
Behaviour, 14, 346–350. https://doi.org/10.1016/S0003-3472(66)80095-7.
ARTICLE IN PRESS
Muma, M. H. (1966b). Egg deposition and incubation for Eremobates durangonus with notes
on the eggs of other species of Eremobatidae (Arachnida: Solpugida). Florida Entomologist,
49, 23–31. https://doi.org/10.2307/3493311.
Muma, M. H., & Muma, K. E. (1988). The arachnid order Solpugida in the United States
(Supplement 2, A Biological Review) (pp. 1–35). Silver City, New Mexico: Privately
published for the authors by Southwest Offset.
Nappi, A. J. (1965). Notes on the courtship and mating habits of the wolf spiders Lycosa helluo
Walckenaer. The American Midland Naturalist, 74(2), 368–373.
Noble, G. K., & Putnam, P. G. (1931). Observations on the life history of Ascaphus truei
Stejneger. Copeia, 1931(3), 97–101. https://doi.org/10.2307/1437329.
Olguı́n-Perez, L., Francke, O. F., & Carbajal-Saucedo, A. (2021). Evidence of piercing and
sexual differences in venom composition in a sexual stinging scorpion (Scorpiones:
Euscorpiidae). The Journal of Arachnology, 49(1), 98–107. https://doi.org/10.1636/
JoA-S-19-056.
Olivero, P. A. (2014). Evolución en escorpiones: Un enfoque multidisciplinario utilizando a
Bothriurus bonariensis como modelo. (PhD dissertation, Córdoba, Argentina: Universidad
Nacional de Córdoba.
Olivero, P. A., Mattoni, C. I., & Peretti, A. V. (2017). Differences in mating behavior
between two allopatric populations of a neotropical scorpion. Zoology, 123, 71–78.
https://doi.org/10.1016/j.zool.2017.06.009.
Olivero, P. A., Vrech, D. E., Oviedo-Diego, M. A., Mattoni, C. I., & Peretti, A. V. (2019).
Courtship performance as function of body condition in an ‘ancient’ form of sperm trans-
fer. Animal Biology, 69(1), 33–46. https://doi.org/10.1163/15707563-00001041.
Osterloh, A. (1922). Beitr€age zur Kenntnis des Kopulationsapparates einiger Spinnen.
Zeitschrift f€
ur Wissenschaftliche Zoologie, 119, 326–421.
Outeda-Jorge, S. (2010). Corte e aspectos da biologia reprodutiva do escorpião brasileiro Tityus
bahiensis (Scorpiones: Buthidae) (PhD dissertation). São Paulo, Brazil: Universidade de
São Paulo.
Oviedo-Diego, M. (2022). Biologıá Reproductiva y Evolución en arácnidos: estudio
multidisciplinario en dos especies modelo de escorpiones (Scorpiones, Bothriuridae, Urophonius)
(PhD thesis). Argentina: Universidad Nacional de Córdoba.
Oviedo-Diego, M., Costa-Schmidt, L. E., Mattoni, C. I., & Peretti, A. V. (2021). Interaction
between sexual communication functions leads to reproductive interference in two
syntopic scorpion species. Animal Behaviour, 181, 83–93. https://doi.org/10.1016/
j.anbehav.2021.08.029.
Pandram, B., & Sharma, V. K. (2018). Mating strategies of solifugae Galeodes olivier 1791
(Arachnida: Galeodidae) in laboratory conditions. International Journal of Zoology and
Applied Biosciences, 3, 141–144.
Parisot, C. (1962). E tude de quelques opilions de Lorraine. Vie Millieu, 13, 179–197.
Parker, G. A. (1970). Sperm competition and its evolutionary consequences in the insects.
Biological Reviews, 45, 525–567. https://doi.org/10.1111/j.1469-185X.1970.tb01176.x.
Parker, G. A. (1984). Sperm competition and the evolution of animal mating strategies. In
R. L. Smith (Ed.), Sperm competition and the evolution of animal mating systems (pp. 1–60).
New York: Academic Press.
Parker, G. A. (1998). Sperm competition and the evolution of ejaculates: Toward a theory
base. In T. R. Birkhead, & A. P. Møller (Eds.), Sperm competition and sexual selection
(pp. 3–54). New York: Academic Press.
Partan, S. R. (2013). Ten unanswered questions in multimodal communication. Behavioral
Ecology and Sociobiology, 67(9), 1523–1539. https://doi.org/10.1007/s00265-013-1565-y.
Pereira, W., Elpino-Campos, A., Del-Claro, K., & Machado, G. (2004). Behavioral reper-
tory of the neotropical harvestman Ilhaia cuspidata (Opiliones, Gonyleptidae). The Journal
of Arachnology, 32(1), 22–30. https://doi.org/10.1636/S02-35.
ARTICLE IN PRESS
Peretti, A. V. (1993). Estudio de la biologıá reproductiva en escorpiones argentinos (Arachnida.

Scorpiones): un enfoque etológico. (PhD dissertation, Córdoba, Argentina: Universidad
Nacional de Córdoba.
Peretti, A. V. (1995). Análisis de la etapa inicial del cortejo en Bothriurus bonariensis
(C. L. Koch) (Scorpiones, Bothriuridae) y su relación con el reconocimiento sexual.
Revue Arachnologique, 11(4), 35–45.
Peretti, A. V. (2001). Patrones de resistencia femenina y respuesta del macho durante el
apareamiento en escorpiones Bothriuridae y Buthidae: que hipótesis puede explicarlos
mejor. Revista de Etologıá (Brasil), 3, 25–45.
Peretti, A. V. (2003). Functional morphology of spermatophores and female genitalia in
bothriurid scorpions: Genital courtship, coercion and other possible mechanisms.
Journal of Zoology, 261(2), 135–153. https://doi.org/10.1017/S095283690300400X.
Peretti, A. V. (2010). An ancient indirect sex model. In Leonard, & Córdoba-Aguilar (Eds.),
The evolution of primary sexual characters in animals (pp. 218–248). New York: Oxford
University Press.
Peretti, A. V. (2014). Sexual selection in neotropical species: Rules and exceptions. In R. H.
Macedo, & G. Machado (Eds.), Sexual selection: Perspectives and models from the neotropics
(pp. 33–52). Cambridge, Massachusetts: Academic Press. https://doi.org/10.1016/
B978-0-12-416028-6.00002-5.
Peretti, A. V., Aacosta, L. E., & Benton, T. G. (1999). Sexual cannibalism in scorpions: Fact
or fiction? Biological Journal of the Linnean Society, 68(4), 485–496.
Peretti, A. V., Acosta, L. E., & Martinez, M. A. (2000). Comportamiento de apareamiento en
tres especies de Bothriurus del grupo prospicuus: estudio comparado y su relación. Revue
Arachnologique, 13(5), 73–91.
Peretti, A. V., & Córdoba-Aguilar, A. (2007). On the value of fine-scaled behavioural
observations for studies of sexual coercion. Ethology Ecology and Evolution, 19(1),
77–86. https://doi.org/10.1080/08927014.2007.9522583.
Peretti, A. V., Vrech, D. E., & Hebets, E. A. (2021). Solifuge (camel spider) reproductive
biology: An untapped taxon for exploring sexual selection. The Journal of Arachnology,
49(3), 299–316. https://doi.org/10.1636/JoA-S-20-037.
Peretti, A. V., & Willemart, R. H. (2007). Sexual coercion does not exclude luring behavior
in the climbing camel-spider Oltacola chacoensis (Arachnida, Solifugae, Ammotrechidae).
Journal of Ethology, 25(1), 29–39. https://doi.org/10.1007/s10164-006-0201-y.
Perez-González, A., & Werneck, R. M. (2018). A fresh look over the genital morphology
of Triaenonychoides (Opiliones: Laniatores: Triaenonychidae) unravelling for the first
time the functional morphology of male genitalia. Zoologischer Anzeiger, 272, 81–92.
https://doi.org/10.1016/j.jcz.2017.12.010.
Petrunkevitch, A. (1911). Sense of sight, courtship, and mating in Dugesiella hentzi (Girard),
a theraphosid spider from Texas. Zoologische Jahrb€ ucher Abteilung f€
ur Systematik, 31, 355–376.
Piacentini, L. N., Scioscia, C. L., Carbajal, M. N., Ott, R., Brescovit, A. D., & Ramı́rez,
M. J. (2017). A revision of the wolf spider genus Diapontia Keyserling, and the relationships
of the subfamily Sosippinae. Araneae: Lycosidae.
Polis, G. A., & Farley, R. D. (1979). Behavior and ecology of mating in the cannibalistic scor-
pion, Paruroctonus mesaensis Stahnke (Scorpionida: Vaejovidae). Journal of Arachnology, 33–46.
Polis, G. A., & Sissom, W. (1990). Life history. In G. A. Polis (Ed.), The biology of scorpions
(pp. 161–223). Stanford: Stanford University Press.
Poy, D., Ramı́rez, M. J., Michalik, P., & Piacentini, L. N. (2019). Copulatory mechanics in
the wolf spider Agalenocosa pirity reveals a hidden diversity of locking systems in Lycosidae
(Araneae). Journal of Morphology, 281(2), 250–257.
Preston-Mafham, K. G. (1999). Notes on bridal veil construction in Oxyopes schenkeli Lessert,
1927 (Araneae: Oxyopidae) in Uganda. Bulletin of the British Arachnological Society, 11,
150–152.
ARTICLE IN PRESS
Preston-Mafham, R., & Preston-Mafham, K. (1993). The encyclopedia of land invertebrate

behaviour. Cambridge, Massachusetts: MIT Press.
Proctor, H. C. (1998). Indirect sperm transfer in arthropods: Behavioral and evolutionary
trends. Annual Review of Entomology, 43, 153. https://doi.org/10.1146/annurev.ento.
43.1.153.
Pruitt, J. N., Berning, A. W., Cusack, B., Shearer, T. A., McGuirk, M., Coleman, A., et al.
(2014). Precopulatory sexual cannibalism causes increase egg case production, hatching
success, and female attractiveness to males. Ethology, 120(5), 453–462. https://doi.org/
10.1111/eth.12216.
Punzo, F. (1997). Dispersion, temporal patterns of activity, and the phenology of feeding
and mating behavior in Eremobates palpisetulosus (Solifugae, Eremobatidae). Bulletin of
the British Arachnological Society, 10, 303–307.
Punzo, F. (1998a). The Biology of Camel-Spiders (Arachnida, Solifugae). Boston: Kluwer
Academic Publishers.
Punzo, F. (1998b). Natural history and life cycle of the solifuge Eremobates marathoni Muma &
Brookhart (Solifugae, Eremobatidae). Bulletin of the British Arachnological Society, 11,
111–118.
Punzo, F. (2006). Life history, ecology, and behavior of the wolf spider, Arctosa littoralis
(Araneae: Lycosidae) in Florida. Florida Scientist, 99–115. https://www.jstor.org/
stable/24322345.
Ramı́rez, M. J. (2003). The spider subfamily Amaurobioidinae (Araneae, Anyphaenidae):
A phylogenetic revision at the generic level. Bulletin of the American Museum of Natural
History, 2003(277), 1–262. https://doi.org/10.1206/0003-0090(2003)277<0001:
TSSAAA>2.0.CO;2.
Ramı́rez, M. J., & Michalik, P. (2014). Calculating structural complexity in phylogenies using
ancestral ontologies. Cladistics, 30(6), 635–649. https://doi.org/10.1111/cla.12075.
Requena, G. S., & Machado, G. (2014). Mating behavior of a neotropical arachnid with
exclusive paternal care. Acta ethologica, 17(1), 23–30. https://doi.org/10.1007/s10211-
013-0152-6.
Riechert, S. E., Singer, F. D., & Jones, T. C. (2001). High gene flow levels lead to gamete
wastage in a desert spider system. Genetica, 112(113), 297–319.
Riechert, S. E., & Tracy, C. R. (1975). Thermal balance and prey availability: Bases for a
model relating web-site characteristics to spider reproductive success. Ecology, 56(2),
265–284. https://doi.org/10.2307/1934960.
Roberts, J. A., & Uetz, G. W. (2005). Information content of female chemical signals in the
wolf spider, Schizocosa ocreata: Male discrimination of reproductive state and receptivity.
Animal Behaviour, 70, 217–223.
Romero-Lebrón, E., Oviedo-Diego, M. A., Elias, D., Vrech, D. E., & Peretti, A. V. (2019).
Effect of the mating plug on female chemical attractiveness and mating acceptance in a
scorpion. Ethology, 125(4), 184–194. https://doi.org/10.1111/eth.12842.
Ross, L. K. (2009). Notes and observations on courtship and mating in Tityus (Atreus)
magnimanus Pocock, 1897 (Scorpiones: Buthidae). Journal of Venomous Animals and Toxins
Including Tropical Diseases, 15, 43–53. https://doi.org/10.1590/S1678-91992009000100005.
Rosin, R., & Shulov, A. (1963). Studies on the scorpiont Nebo hierochonticus. Proceedings of the
Zoological Society of London, 140(4), 547–575. https://doi.org/10.1111/j.1469-7998.
1963.tb01989.x.
Rovner, J. S. (1971). Mechanisms controlling copulatory behavior in wolf spiders (Araneae:
Lycosidae). Psyche, 78(3), 150–165. https://doi.org/10.1155/1971/60846.
Rowsell, J., & Cushing, P. E. (2020). Mating behaviour of Eremobates pallipes (Say, 1823)
(Arachnida: Solifugae: Eremobatidae). Arachnology, 18(4), 399–408. https://doi.org/
10.13156/arac.2020.18.4.399.
ARTICLE IN PRESS
Rypstra, A., & Wilder, S. (2007). Male control of copulation duration in a wolf
spider (Araneae, Lycosidae). Behaviour, 144(4), 471–484. https://doi.org/10.1163/
156853907780756012.
Sadana, G. L. (1972). Mechanics of copulation in Lycosa chaperi Simon (Araneida: Lycosidae).
Bulletin of the British Arachnological Society, 2, 87–89.
Schneider, J., Uhl, G., & Herberstein, M. E. (2015). Cryptic female choice within the genus
Argiope: A comparative approach. In A. Peretti, & A. Aisenberg (Eds.), Cryptic female
choice in arthropods (pp. 55–75). Cham: Springer.
Schoenberg, D. (2021). Males of the wolf spider Rabidosa rabida use two mechanisms to stun females
during copulation (PhD dissertation). Murray, United States: State University.
Schwerdt, L., Pompozzi, G., Copperi, S., & Ferretti, N. E. (2014). Cortejo y cópula de Lycosa
poliostoma y Hogna gumia, dos especies de arañas lobo (Araneae, Lycosidae) de Argentina
(p. 273). III Congreso Uruguayo de Zoologı́a.
Scott, C. E., Andersson, A. G., & Andrade, M. C. B. (2018). A review of the mechanisms and
functional roles of male silk use in spider courtship and mating. The Journal of Arachnology,
46, 173–206. https://doi.org/10.1636/JoA-S-17-093.1.
Scott, C., Vibert, S., & Gries, G. (2012). Evidence that web reduction by western black
widow males functions in sexual communication. The Canadian Entomologist, 144(5),
672–678. https://doi.org/10.4039/tce.2012.56.
Sentenská, L., Šedo, O., & Pekár, S. (2020). Biting and binding: An exclusive coercive mat-
ing strategy of males in a philodromid spider. Animal Behaviour, 168, 59–68. https://doi.
org/10.1016/j.anbehav.2020.08.001.
Shapiro, H. A. (1936). The biological basis of sexual behaviour in amphibia: I. The exper-
imental induction of the mating reflex (coupling) in Xenopus Laevis (The South African
clawed toad) by means of pregnancy urine and of anterior pituitary extracts, with the
production of fertilised ova. Journal of Experimental Biology, 13(1), 48–56. https://doi.
org/10.1242/jeb.13.1.48.
Shulov, A. (1940). On the biology of two Latrodectus spiders in Palestine. Proceedings of the
Linnean Society of London, 152(3), 309–328. https://doi.org/10.1111/j.1095-8312.
1940.tb00267.x.
Singer, F., Morris, A., Becker, E., Riechert, S., Xu, H., Hale, J., et al. (2000). Analysis
of courtship success in the funnel-web spider Agelenopsis aperta. Behaviour, 137(1),
93–117. https://doi.org/10.1163/156853900501890.
Singer, F., & Riechert, S. E. (1995). Mating system and mating success of the desert spider
Agelenopsis aperta. Behavioral Ecology and Sociobiology, 36(5), 313–322. https://doi.org/
10.1007/BF00167792.
Snow, L. S., Abdel-Mesih, A., & Andrade, M. C. (2006). Broken copulatory organs
are low-cost adaptations to sperm competition in redback spiders. Ethology, 112(4),
379–389. https://doi.org/10.1111/j.1439-0310.2006.01163.x.
Snow, L. S., & Andrade, M. C. (2004). Pattern of sperm transfer in redback spiders:
Implications for sperm competition and male sacrifice. Behavioral Ecology, 15(5),
785–792. https://doi.org/10.1093/beheco/arh080.
Stanley, E., Francescoli, G., & Toscano-Gadea, C. A. (2016). Mating behavior of the
solitary neotropical harvestman Pachyloides thorellii (Arachnida: Opiliones). The Journal
of Arachnology, 44(2), 210–217. https://doi.org/10.1636/J15-33.
Stephenson, B., & Verrell, P. (2003). Courtship and mating of the tailed frog (Ascaphus truei).
Journal of Zoology, 259(1), 15–22. https://doi.org/10.1017/S095283690200331X.
Tallarovic, S. K., Melville, J. M., & Brownell, P. H. (2000). Courtship and mating in the
giant hairy desert scorpion, Hadrurus arizonensis (Scorpionida, Iuridae). Journal of Insect
Behavior, 13(6), 827–838. https://doi.org/10.1023/A:1007858400402.
Toscano-Gadea, C. A. (2010). Sexual behavior of Bothriurus buecherli (Scorpiones:
Bothriuridae) and comparison with the B. prospicuus group. The. Journal of Arachnology,
38(2), 360–363. https://doi.org/10.1636/Hi09-31.1.
ARTICLE IN PRESS
Toscano-Gadea, C. A., & González, M. (2019). The sexual behaviour of the neotropical wolf
spider Lycosa inornata (Araneae: Lycosidae), with comments on maternal behaviour.
Arachnology, 18(3), 270–275. https://doi.org/10.13156/arac.2019.18.3.270.
Toscano-Gadea, C. A., & González, M. (2020). ¿El aroma del amor...engaña? Comportamiento
sexual de Lycosa poliostoma y su respuesta ante feromonas de otro licósido sincrónico y simpátrido.
VI Congreso Latinoamericano de Aracnologı́a (VICLA), virtual: 253.
Tuni, C., Schneider, J., Uhl, G., & Herberstein, M. E. (2020). Sperm competition when
transfer is dangerous. Philosophical Transactions of the Royal Society B, 375(1813),
20200073. https://doi.org/10.1098/rstb.2020.0073.
Uhl, G., & Elias, D. (2011). Communication. In M. E. Herberstein (Ed.), Spider behaviour:
Flexibility and versatility (pp. 127–189). Cambridge, UK: Cambridge University Press.
Uhl, G., Nessler, S. H., & Schneider, J. M. (2010). Securing paternity in spiders? A review on
occurrence and effects of mating plugs and male genital mutilation. Genetica, 138(1),
75–104.
Vibert, S., Scott, C., & Gries, G. (2016). Vibration transmission through sheet webs of hobo
spiders (Eratigena agrestis) and tangle webs of western black widow spiders (Latrodectus
hesperus). Journal of Comparative Physiology A, 202(11), 749–758. https://doi.org/10.
1007/s00359-016-1113-0.
Vrech, D. E., Oviedo-Diego, M. A., Olivero, P. A., & Peretti, A. V. (2019). Successive mat-
ings produce opposite patterns on ejaculate volume and spermatozoa number in an
ancient arthropod model with indirect sperm transfer. Canadian Journal of Zoology,
97(7), 579–587. https://doi.org/10.1139/cjz-2018-0179.
Wernz, J. G., & Storm, R. M. (1969). Pre-hatching stages of the tailed frog, Ascaphus truei
Stejneger. Herpetologica, 86–93. https://www.jstor.org/stable/3890950.
Weygoldt, P. (1969). The biology of pseudoscorpions. Cambridge, Massachusetts: Harvard
University Press.
Weygoldt, P. (1990). Arthropoda -Chelicerata: Sperm transfer. In K. G. Adiyodi, & R. G.
Adiyodi (Eds.), Reproductive biology of invertebrates. Vol. IV, part B. fertilization, development,
and parental care (pp. 77–119). New Dehli, Bombay, Calcutta: Oxford & IBH
Publishing Co.
Weygoldt, P. (2000). Whip spiders (Amblypygi): Biology, morphology and systematics. Stenstrup,
Denmark: Apollo Books.
Weygoldt, P. (2002). Sperm transfer and spermatophore morphology of the whip spiders
Sarax buxtoni, S. brachydactylus (Charinidae), Charon cf. grayi, and Stygophrynus brevispina
nov. spec. (Charontidae) (Chelicerata, Amblypygi). Zoologischer Anzeiger-A Journal of
Comparative Zoology, 241(2), 131–148.
Wharton, R. A. (1987). Biology of the diurnal Metasolpuga picta (Kraepelin) (Solifugae,
Solpugidae) compared with that of noctumal species. Journal of Arachnology, 14,
363–383. https://www.jstor.org/stable/3705676.
Wheeler, W. C., Coddington, J. A., Crowley, L. M., Dimitrov, D., Goloboff, P. A.,
Griswold, C. E., et al. (2016). The spider tree of life: Phylogeny of Araneae based on
target-gene analyses from an extensive taxon sampling. Cladistics, 33(6), 574–616.
Whitehouse, M. E., & Jackson, R. R. (1994). Intraspecific interactions of Argyrodes
antipodiana, a kleptoparasitic spider from New Zealand. New Zealand Journal of
Zoology, 21(3), 253–268.
Wilder, S. M., & Rypstra, A. L. (2004). Chemical cues from an introduced predator
(Mantodea, Mantidae) reduce the movement and foraging of a native wolf spider
(Araneae, Lycosidae) in the laboratory. Environmental Entomology, 33(4), 1032–1036.
https://doi.org/10.1603/0046-225X-33.4.1032.
Wilder, S. M., Rypstra, A. L., & Elgar, M. A. (2009). The importance of ecological and phy-
logenetic conditions for the occurrence and frequency of sexual cannibalism. Annual
Review of Ecology, Evolution, and Systematics, 40(2). https://doi.org/10.1146/annurev.
ecolsys.110308.120238.
ARTICLE IN PRESS
Willemart, R. H., Farine, J. P., Peretti, A. V., & Gnaspini, P. (2006). Behavioral roles of the
sexually dimorphic structures in the male harvestman, Phalangium opilio (Opiliones,
Phalangiidae). Canadian Journal of Zoology, 84(12), 1763–1774. https://doi.org/
10.1139/z06-173.
Willemart, R. H., Osses, F., Chelini, M. C., Macı́as-Ordóñez, R., & Machado, G. (2009).
Sexually dimorphic legs in a neotropical harvestman (Arachnida, Opiliones): Ornament
or weapon? Behavioural Processes, 80(1), 51–59. https://doi.org/10.1016/j.beproc.2008.
09.006.
Xiao, Y. H., Zunic-Kosi, A., Zhang, L. W., Prentice, T. R., McElfresh, J. S., Chinta, S. P.,
et al. (2015). Male adaptations to minimize sexual cannibalism during reproduction
in the funnel-web spider Hololena curta. Insect Sci., 22(6), 840–852. https://doi.org/
10.1111/1744-7917.12243.
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Waking beauties: Mating quiescence in arachnid

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Waking beauties: Mating

Advances in the Study of Behavior Copyright # 2023 Elsevier Inc. 1

2 Franco Cargnelutti et al.

9.4 Copulation substrate constraints 81

Waking beauties: Mating quiescence in arachnid females 3

as occurs in scorpions (Scorpiones), pseudoscorpions (Pseudoscorpiones),

4 Franco Cargnelutti et al.

Waking beauties: Mating quiescence in arachnid females 5

so-called “tonic immobility” during first episodes of amplexus (Noble &

6 Franco Cargnelutti et al.

quiescence duration has been associated with increased numbers of sperm

Waking beauties: Mating quiescence in arachnid females 7

Araneomorphae: Agelenidae (RTA)

8 Franco Cargnelutti et al.

Alopecosa solitaria Yes No III Wandering – Entelegyne

Waking beauties: Mating quiescence in arachnid females 9

Tigrosa helluo Yes No III Wandering – Entelegyne

10 Franco Cargnelutti et al.

Latrodectus geometricus No F > M II Web builder TS Entelegyne

Waking beauties: Mating quiescence in arachnid females 11

Eupalaestrus No No I Sendentary B Haplogyne

However, as we will show, female immobility may not necessarily include

Bothriurus No Absent Pre Grasping, Reorientation, None Totally docile, NA NA SP / Intersexual 2, 4, 6

50 Franco Cargnelutti et al.

Waking beauties: Mating quiescence in arachnid females 51

BOX 1 Contexts that may have favored the evolution

3. Terminologies, history, and other concerns

52 Franco Cargnelutti et al.

this female behavior by Heymons (1902) were “passive state,” “will-less

Waking beauties: Mating quiescence in arachnid females 53

54 Franco Cargnelutti et al.

4. General results of the survey

Table 6 Definitions of the operational levels of female mating quiescence.

Waking beauties: Mating quiescence in arachnid females 55

In the following sections, we compile different aspects of the biology of each

5. Quiescence and mating phase

56 Franco Cargnelutti et al.

In harvestmen, female quiescence has been described at the pre-

6. Arachnids and quiescence levels

Waking beauties: Mating quiescence in arachnid females 57

Female quiescence in solifuges has been reported in 13 species distributed

7. Behavioral and morphological traits associated

58 Franco Cargnelutti et al.

7.1 Quiescence and female body position

Waking beauties: Mating quiescence in arachnid females 59

been reported in representatives of all families (except Eremobatidae), with

60 Franco Cargnelutti et al.

Finally, in the pseudoscorpion Dactylochelifer latreillii latreillii the female

7.2 Quiescence and female movements

Waking beauties: Mating quiescence in arachnid females 61

7.3 Quiescence and male behaviors

62 Franco Cargnelutti et al.

injection (Sentenská et al., 2020) or purely a mechanical stimulus, as has been

Waking beauties: Mating quiescence in arachnid females 63

Galeodes sulfuripes, mainly because of the description of how females become

64 Franco Cargnelutti et al.

7.4 Quiescence, aggression, and cannibalism

Waking beauties: Mating quiescence in arachnid females 65