Records of the Western Australian Museum Supplement No. 52: 1-11 (1995).

The duration of copulation in spiders: comparative patterns

Mark A. Elgar
Department of Zoology, University of Melbourne, Parkville, Victoria 3052, Australia

Abstract - The duration of copulation in spiders varies both-within and

between species, and in the latter by several orders of magnitude. The
sources of this variation are explored in comparative analyses of the duration
of copulation and other life-history variables of 135 species of spiders from
26 families. The duration of copulation is correlated with body size within
several species, but the pattern is not consistent and more generally there is
no inter-specific covariation between these variables. The duration of
copulation within orb-weaving spiders is associated with both the location of
mating and the frequency of sexual cannibalism, suggesting that the length
of copulation is limited by the risk of predation. Finally, entelegyne spiders
copulate for longer than haplogyne spiders, a pattern that can be interpreted
in terms of male mating strategies or the complexity of their copulatory
apparatus.

INTRODUCTION after the copulatory organ has been inserted. In

It is widely recognised that there are conflicts of species in which females mate with several males,
interest between males and females in the choice of copulation may provide the male with the
mating partner and the frequency of mating (e.g. opportunity to manipulate the sperm of other
Elgar 1992). Thus, while the principal function of males that previously mated with that female. For
copulation is to transfer gametes, the act of mating example, copulating male damselflies not only
may have several additional functions, such as transfer their own sperm, but also remove the
mate assessment or ensuring sperm priority, and sperm of rival males (e.g. Waage 1979; Siva-Jothy
activities other than sperm transfer 'can take place and Tsubaki 1989). Finally, copulating males may
during copulation. These additional activities may also deposit a sperm plug that prevents the sperm
be time-consuming, and thus the time spent of rival males from fertilising the eggs of the female
copulating may not simply reflect the amount of (e.g. Lorch et al. 1993; Masumoto 1993).
time required to transfer gametes. Despite the While longer copulations may provide
potential benefits of these activities, the additional opportunities to increase mating success, there are
time spent copulating may also incur costs, such as several costs of mating that may favour shorter
reducing the amount of time available to find other durations (see Daly 1978; Lewis 1987).. For some
mates (e.g. Parker 1970a). Therefore, the species, the additional time spent copulating may
considerable variation in the duration of copulation reduce the time available to males for seeking cUld
that occurs both within and between species may courting other females (e.g. Parker 1970a), and the
be at least partly attributable to the different costs time available to females for foraging. Copulating
and benefits of various durations. pairs may also be more vulnerable to predators
There are several reasons why copulation may be because they are more obvious or less able to
longer than is necessary to transfer sperm. escape. Finally, the proximity necessary for mating
Eberhard (1985) argues that copulation for some may increase the likelihood that parasites and
species may represent an additional component of infectious diseases are transmitted between the
courtship, allowing the female to exercise some mating partners.
degree of mate choice. Supporting evidence for this Studies of the factors responsible for the
view is provided by species in which copulation variation in the duration of copulation have
takes place without insemination, such as the focussed primarily on single species, and
linyphiid spider Lepthyphantes leprosus, in which the particularly the relationship between the duration
male courts and then copulates with the female, of copulation and sperm competition (e.g. Parker
even though the male is unable to transfer sperm 1970a; Jackson 1980; Siva-Jothy and Tsubaki 1989;
(van Helsdingen 1965). It may also explain why Lorch et al. 1993). In contrast, few studies have
insemination does not always occur immediately examined the patterns of inter-specific variation in
2 M.A. EIgar

the duration of copulation (but see Parker 1970b). (Ridley 1989). Nevertheless, character differences
The aim of this study is to both describe the between species within lower taxonomic categories
variation in the duration of copulation in spiders, can, and do, arise through evolutionary processes.
and explore some of the factors that may explain Consequently, comparative studies are frequently
this variation. plagued with the problem of distinguishing
Male spiders lack primary copulatory organs, patterns that arise through convergent and/ or
and instead transfer sperm using modified parallel evolution from those that arise from shared
pedipalps. As a consequence, the transfer of ancestry (Harvey and PageI1991).
gametes in spiders involves two phases. The initial This problem has stimulated the development of
phase, called sperm induction, involves the increasingly sophisticated statistical methods for
transfer of sperm from the male gonopore to his comparative analyses (Harvey and Pagel 1991), all
palps. First the male constructs a sperm web, of which require an accurate phylogenetic
usually a small, horizontally-suspended, triangular arrangement of the data. Unfortunately, the
structure. He then presses his abdomen against the phylogenetic relationships of spiders are still
surface of this web, moving his abdomen up and unclear, and in many taxa the systematics are
down, until a drop of sperm emerges from his incomplete (Coddington 1990; Coddington and
genital opening and is deposited on the sperm web. Levi 1991). Hence, these statistical techniques are
The male dips his pedipalps into the sperm, which unlikely to be appropriate for the present study
is taken up into the palps. Females can only mate and have not been used. Instead, a simple approach
after they have completed their final moult, and has been used in which values for higher
following a successful courtship, the male taxonomic levels are the units of analysis (e.g. Elgar
copulates with the mature female by inserting his et al. 1990). In most cases, this is the family, and the
palpal organ into her genital opening. There are value calculated is the avera-ge of the constituent
differences between species in the frequency of values for the genera. The generic values are the
palpal insertions, and whether the palps are averages of their constituent species values. This is
inserted· simultaneously or consecutively. Further not an ideal procedure, and hence the patterns
details of the copulatory behaviour of spiders are revealed by this analysis should be interpreted
provided by Foelix (1982). cautiously, since it is possible that the patterns may
reflect phylogenetic affinities, rather than
convergent or parallel evolution.
METHODS AND DATA The duration of copulation and body length
Data on the duration of copulation, assembled values were logarithmically transformed in order
from the literature and unpublished data, were to normalise their distributions. Statistical analyses
obtained for 135 species of spiders in 27 families. were performed using procedures in the Systat 5.2
The number of observations of copulations used to computer package (Wilkinson 1991).
yield a species value varied from study to study,
and in some cases may represent a single
observation. It is unlikely that this is an exhaustive RESULTS AND DISCUSSION
summary of the available data. Furthermore, the The duration of copulation in the present sample
values for particular species may change with the of spider species varied by several orders of
accumulation of more observations. Additional magnitude. In some species, especially web-
information on male and female body length, and building spiders, copulation is completed in less
several behavioural characters, were also obtained than half a minute (e.g. Bristowe 1958; Robinson
for most of these species. These data, together with and Robinson 1980; Elgar and Nash 1988). In
the sources, are given in the Appendix. The contrast, copulation in the salticid spider Pseudicius
taxonomy of spiders are still poorly understood, can last up to 15 hours (Jackson 1986a). The
despite considerable progress over the last twenty distribution of the data favoured short copulations;
years (see Coddington and Levi 1991), and hence almost three-quarters of the families surveyed
the species names given in the Appendix may copulated for less than an hour, and
differ from the original sources. It also seems representatives of only three families copulated for
inevitable that some of the generic names listed in longer than two hours (see Fig. 1).
the Appendix will change.
Body size
Analysis The duration of copulation may be influenced by
In general, species within lower taxonomic body size in several ways, if the number of sperm
categories have similar characteristics as a result of transferred is positively correlated with the
their most recent shared ancestry. Hence, species duration of copulation (e.g. Austad 1982; Fahey
can rarely be assumed t.o represent independent 1992; but see Jackson 1980; Watson 1991). For
data points for the purposes of statistical analyses example, larger females may produce more eggs
Duration of copulation in spiders 3
10
have slower flight responses. Many spiders build
silk nests, which may allow them to remain
concealed and hence avoid predators (see Jackson
1986b). If the risk of predation is higher for spiders
in copula, then spiders mating in the open should
6 copulate for shorter periods than those that mate
inside the nest. The results of studies of several
cursorial spiders are consistent with this view. The
4 duration of copulation is longer for spiders mating
inside the nest than outside in the dysderid Dysdera
crocata (Jackson and Pollard 1982), and in three
2
salticids Holoplatys sp. (Jackson and Hardin 1982),
Plexippus paykulli (Jackson and McNab 1989) and
Phidippus johnsoni (Jackson 1980).
o
0-20 21-40 41-60 61-80 81-100 101-120 121-140 141-160 >160 The location of copulation may similarly
influence inter-specific variation in the duration of
Duration of copulation (min) copulation. For example, Portia is a web-building
Figure 1 Frequency distribution of the duration of salticid that never mates inside a nest, and also has
copulation across 26 families of spiders. a relatively short copulation time for a salticid
(Jackson and Hallas 1986). Unfortunately, there are
than smaller females, and thus longer durations of insufficient data to examine this possibility for
copulation may be required to transfer more cursorial spiders; although there are records of
sperm. The patterns obtained from studies of spiders living in nests, it is not always clear
several species are not consistent. The duration of whether mating takes place within these nests.
copulation is positively correlated with female size Nevertheless, a similar argument may apply to orb-
in the golden orb-weaver Nephila plumipes (Elgar, weaving spiders, which mate in different locations
unpublished) and autumn spider Metellina on the web. In general, orb-weaving spiders
segmentata (Prenter et al. 1994), but with male body copulate at either the central hub of the web, or
size in the orb-weaver Araneus diadematus (Elgar while suspended from a specially constructed
and Nash 1988). However, there were no thread that is attached to either the orb-web or the
significant correlations between the duration of supporting threads. Spiders copulating while
copulation and either male or female body size in suspended from a mating thread may be at a
the orb-weavers Gasteracantha minax or Argiope higher risk of predation than those copulating at
appensa (Elgar, unpublished). Interestingly, these the hub, because pairs on a mating thread can be
intra-specific correlations are not apparent in attacked from any direction and may be less able
comparisons across taxa. There is no correlation to take evasive action, while pairs at the hub are
across taxa between the duration of copulation and protected at least from one side by the orb-web.
either male body length (r = 0.22, ns, n = 26) or The comparative data on the duration of
female body length (r = 0.09, ns, n = 26). copulation among orb-weaving spiders provide
It is possible that the duration of copulation is support for this view: genera that mate on the hub
influenced by size dimorphism. Perhaps copulation (Argiope, Gasteracantha, Herennia, Isoxya, Mecynofa,
takes longer in strongly dimorphic species because Nephila, Nephilengys and Phonognatha) Fe:rp.ain in
the relatively smaller males will take longer to copula for significantly longer than genera that
transfer larger quantities of sperm. The residuals mate on a thread (Araneus, Cyclosa, Cyrtophora,
from the regression equation of male size on female Eriophora, Gea, Metellina, Micrathena, Uloborus and
size are one measure of size dimorphism: families Zilla) (Table 1).
with males that are much smaller than females
have negative residuals, while positive residuals
indicates little size dimorphism (see Elgar et al. Sexual cannibalism
1990). There is a significant positive correlation Sexual cannibalism may influence at least some
between male and female body size (r = 0.91, P < components of the courtship behaviour of species
0.001, n = 26), but there is no evidence that size in which it occurs (e.g. Elgar 1991, 1992; Prenter et
dimorphism (measured as residuals) and the al. 1994). Like predation, it may also influence the
duration of copulation are correlated (r = 0.34, ns, duration of copulation. For example, the risk of
n = 26). sexual cannibalism may increase with the duration
of copulation: larger male orb-weavers Araneus
Risk of predation diadematus are less likely to be cannibalised by
Animals in copula may be at a higher risk of females than smaller males, and larger males also
predation because they are more easily detected or copulate for longer periods than smaller males
 M.A. Elgar

(Elgar and Nash 1988). Alternatively, a female may in spiders: a male Linyphia litigosa may guard a
control the duration of copulation more effectively female from rival males by destroying her web so
by killing her mate. Both conditions predict that that other males cannot detect her sex pheromones
durations of copulation will be shorter among (Watson 1986); and males of many species place a
sexually cannibalistic species than non-cannibalistic sperm plug in the female's reproductive tract
species. which prevents the sperm of other males from
Sexual cannibalism occurs relatively frequently fertilising the eggs (e.g. Masumoto 1993). These
in three (Araneidae, Metidae and Pisauridae) of the behaviours all function to reduce the probability
28 families sampled (see Elgar 1992). Although the that the sperm of a rival male fertilises the female's
duration of copulation appears to be shorter in eggs, and they can lead to variation in both the
sexually cannibalistic families (Table 1), the duration of copulation and patterns of sperm
variation is sufficiently large that the difference is precedence (e.g. Lorch et al. 1993).
not significant. Comparisons of the duration of
copulation among orb-weaving spiders and among Reproductive morphology
salticids reveals that sexually cannibalistic genera Spermathecal morphology of spiders has
copulate for shorter times than those genera that interesting implications for sperm precedence
are not usually sexually cannibalistic (Table 1). patterns, assuming that little, or no, mixing of the
sperms occurs when females mate with several
Female mating frequency males (see Austad 1984; Eberhard et al. 1993). In
Multiple mating by females is a powerful general, spiders can be conveniently assigned to
selection pressure favouring a variety of male one of two groups according to their spermathecal
behaviours that ensure the eggs of the female he morphology. The spermatheca of one group, the
has mated with are not fertilised by the sperm of a entelegynes, consists of two ducts; the insemination
rival male. The mechanisms of sperm competition, duct that opens near the vaginal opening and into
the competition between the ejaculates of different which the male intromittent organ dispenses
males over fertilization of a given set of eggs seminal fluid, and the fertilisation duct from which
(Parker 1970b), can be quite varied. For example, sperms issue when the eggs are fertilised. In
males may either displace or remove sperm of contrast, the spermathecae of haplogyne spiders
previously-mating males (e.g. Siva-Jothy and are more simple, consisting of a single duct that
Tsubaki 1989), or produce large quantities of sperm joins the lumen of the spermatheca to the vagina.
if the fertilization success of a male depends upon Sperms pass through this duct on their way to the
the proportionate representation of his sperm in storage organ, and must then return by the same
the female's reproductive tract (e.g. Parker 1990; route when the eggs are fertilised (see Foelix 1982).
Stockley and Purvis 1993). Similar strategies occur For entelegyne spiders with a 'conduit'

Table 1 Sources of variation in the duration of copulation in spiders.

Sources of variation Mean SE n t1

duration
(min)

Location of mating (among orb-weavers)

Mating thread 3.7 1.9 9 2.98**
Hub 37.6 14.3 8
Sexual cannibalism (all families)
Sexually cannibalistic 15.3 7.2 3 1.03
Not sexually cannibalistic 54.8 12.9 24
Sexual cannibalism (among orb-weavers)
Sexually cannibalistic 2.8 3.5 7 2.84**
Not sexually cannibalistic 31.5 12.0 10
Sexual cannibalism (among salticids)
Sexually cannibalistic 6.5 3.5 3 1.92t
Not sexually cannibalistic 61.7 36.5 12
Spermathecal morphology (all families)
Haplogyne 21.9 5.7 9 2.06*
Entelegyne 64.6 16.5 18

1. t-values are derived from tests with log-transformed data.

t. p = 0.08; * P = 0.05; ** P = 0.01
Duration of copulation in spiders 5

spermathecal tract, the last sperm to enter the transferred is positively correlated with the
female is likely to be furthest from the fertilisation duration of copulation (e.g. Austad 1982; Fahey
duct and hence the last to be used in fertilisations. 1992), selection may favour longer durations of
Thus, the sperm of the male that is first to mate copulation in entelegyne spiders than in haplogyne
with a female is thought to fertilise most or all of spiders. A comparison of the durations of
her eggs. In contrast, the 'cul-de-sac' spermathecal copulation at the family level for these two groups
tract of haplogyne spiders favours last male sperm of spiders is consistent with this view: the duration
priority because the last sperm to enter is closest to of copulation of entelegyne spiders is significantly
the fertilisation duct, and hence the first to fertilise longer than that ofhaplogyne spiders (Table 1).
the eggs (see Austad 1984). This view was, until While the covariation between the duration of
recently, generally supported by the available copulation and spermathecal morphology may
evidence for entelegyne spiders. First-male sperm have arisen through male mating str:ategies, other
precedence occurs in three families (but see Austad interpretations are possible. For example, the
1984), including. salticids (Jackson 1980), araneids longer copulation duration of entelegyne spiders
(Vollrath 1980; Christenson and Cohn 1988), and may arise if these species more commonly place
linyphiids (Austad 1982; Martyniuk and Jaenike and remove sperm plugs. Alternatively, the
1982; Watson 1982), but not in an agelenid duration of copulation may be positively correlated
(Masumoto 1993). However, there appears to be with the complexity of the copulatory apparatus.
highly variable sperm precedence arrangement in The copulatory apparatus in both male and female
the haplogyne Physocyclus globosus that has a cul- haplogyne spiders is relatively simple compared
de-sac tract (Eberhard et al. 1993). with entelegyne spiders (see Fbelix 1982). Not only
Different sperm precedence patterns are likely to is the male palp of entelegyne spiders more
influence the mating systems of a wide range of complex, but the embolus (which is used to
organisms, including spiders (Austad 1984, see also transfer sperm) must pass through a complex
Ridley 1989). Males of entelegyne spiders (with female genital opening into the insemination duct.
conduit-type spermathecae) should preferentially Thus the longer duration of copulation of
associate with virgin rather than mated females, entelegyne spiders may simply reflect the longer
while males of haplogyne spiders (with cul-de-sac time required to couple successfully if there are no
spermathecae) should either show no preference or differences between the two groups in the
attempt to ensure that they are last to mate. Several frequency of palpal insertions during copulation.
lines of evidence support this prediction: in a broad This latter explanation is unlikely, however, to
survey of largely anecdotal data, Jackson (1986b) account for very long copulations.
catalogued 156 entelegyne species and five
haplogyne species that cohabit with immature
females. Watson (1990) demonstrated that males of Mating status of the female
the entelegyne Linyphia litigiosa prefer to cohabit The mating status of the female may also
with immature rather than mature females when influence the duration of copulation. For example,
both are available, and Eberhard et al. (1993) males may copulate longer with mated females
showed that males of two of three entelegyne than virgin females because they either transfer
species show a strong preference for immature more sperm or even displace the sperm of the
females, while males of all three haplogyne species previous male. Alternatively, males of entelegyne
studied did not show any preference. species may terminate copulation with mated
The differences in spermathecal morphology, females earlier because they are unlikely to fertilise
together with the associated trends in male mate any of the eggs (e.g. Suter 1990).
preferences, may also explain differences in the There are no consistent patterns among the few
duration of copulation. If male entelegyne spiders studies that compare the duration of copulation of
that mate with virgin females fertilise most of the virgin and mated females. Copulation among
female's clutch, then males could maximise their entelegyne spiders was longer for mated females
reproductive success by ejaculating larger than virgin females in Phonognatha graeffei
quantities of sperm. In contrast, it appears that (Tetragnathidae) (Fahey 1992); Phidippus johnsoni
male haplogyne spiders can obtain high levels of (Salticidae) (Jackson 1980); Argyrodes antipodiana
paternity only by preventing (e.g. with mating (Theridiidae) (Whitehouse 1991); Achaearanea wau
plugs) rival males from subsequently mating with (Theridiidae) (Lubin 1986); and Misumenoides
the female. In the absence of such mechanisms, formosipes (Thomisidae) (Dodson and Beck 1993).
males of these spiders may maximise their Males of the linyphiid Frontinella pyramitela
reproductive success by ejaculating smaller (Austad 1982) that copulated with non-virgin
quantities of sperm, but mating with many females females terminated copulation early and failed to
(see also Parker 1990). Under these conditions, and transfer sperm. However, there was no difference
assuming that the number of spermatozoa in the copulation duration of mated and virgin
6 M.A. EIgar

females in three araneids Gasteracantha minax, Mike Gray, Mark Harvey, Robert Jackson, Michael
Nephila plumipes and Argiope appensa (Elgar, Magrath, Angus Martin, and Laila Sadler for
unpublished). There are no data available for advice, discussions and comments on the
haplogyne spiders. manuscript; and the University of Melbourne, the
It is not clear why males of only some species Australian Research Council (A19130739), and the
apparently detect the mating status of the female WV Scott Fund for their financial support. Finally,
and adjust the duration of copulation accordingly. I wish to express my appreciation of Barbara York
From the perspective of the male, the duration of Main, whose book and discussions have been
copulation will depend on both the relationship responsible for stimulating and maintaining my
between the duration of copulation and the amount interest in spiders.
of sperm transferred, and on the pattern of sperm
precedence among these spiders. The data from
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University of Melbourne. biology of Portia africana, P. albimana, P. jimbriata, P.
Foelix, RF. (1982). Biology of Spiders. Harvard University labiata, and P. shuitzi, araneophagic, web-building
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Forster, L.M. (1992). The stereotyped behaviour of sexual webs, predatory versatility, and intraspecific
cannibalism in Latrodectus hasseIti Thorell (Araneae: interactions. New Zealand Journal of Zoology 13: 423-
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Fraser, J.B. (1987). Courtship and copulatory behaviour interactions of Holoplatys sp. indet., a New Zealand
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Harvey, P.H. and Pagel, M.D. (1991). The Comparative Jackson, RR and Macnab, A.M. (1989). Display, mating,
Method in Evolutionary Biology. Oxford University and predatory behaviour of the jumping spider
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Mallos (Araneae, Dictynidae). H. The relationship Jackson, RR and Macnab, A.M. (1991). Comparative
between courtship, mating, agression and study of the display and mating behaviour of
cannibalism in species with differing types of social lyssomanine jumping spiders (Araneae: Salticidae),
organization. Revue Arachnologique 2: 103-132. especially Asemonea tenuipes, Goleba puella, and
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Zealand Journal ofZoology 16: 1-16. (Lycosa rabida Walckenaer): a quantitative study.
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Lorch, P.D., Wilkinson, G.S. and Reillo, P.R (1993). insertion patterns. Animal Behavior 22: 94-99.
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stalk-eyed fly Cyrtodiopsis whitei (Diptera: copulation duration in Mnais pruinosa pruinosa Selys
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309-311. securing tactics and sperm precedence. Behavioral
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1-218. Manuscript received 5 April 1994; accepted 21 July 1994.
Duration of copulation in spiders 9

APPENDIX
The duration of copulation of 135 spider species from 26 families.

Taxa Body length Copulation Sexual Source

(mm) duration cannibal
Male Female (min)
HAPLOGYNES
Dipluridae
Microhexura montivaga 2.5 2.4 0.5 No Coyle (1985)
Thelechoris karschi 16.0 14.0 40.4 No Coyle and O'Shields (1990)
Dysderidae
Dysdera crocata .12.0 10.0 17.0 Rare Jackson and Pollard (1982)
Harpactea hombergi 7.0 6.0 4.0 No Bristowe (1958)
Hexathelidae
Porrhothele antipodiana 27.0 Jackson and Pollard (1990)
Hypochilidae
Hypochilus pococki 16.0 11.0 1.4 No Catley (1993)
Oonopidae
Oonops pulcher 2.0 1.5 1.5 No Bristowe (1958)
Pholcidae
Pholcus phalangioides 7.5 7.5 65.5 No Uhl (1993)
Psilochorus simoni 2.5 2.5 22.0 No Bristowe (1958)
Scytodidae
Scytodes thoracica 5.0 4.0 44.0 No Bristowe (1958)
Segestriidae
Segestria jlorentina 22.0 15.0 10.0 Bristowe (1958)
Tetragnathidae
Pachygnatha clercki 8.9 5.3 100.0 Bristowe (1958)
Pachygnatha degeeri 25.0 Bristowe (1958)
Tetragnatha extensa 8.5 6.5 15.0 No Bristowe (1958)
ENTELEGYNES
Agelenidae
Agelena limbata 193.0 Masumoto (1991)
Coelotes atropos 150.0 Bristowe (1958)
HoloZena adnexa 81.8 Fraser (1987)
Textrix denticuZata 7.0 6.5 30.0 No Bristowe (1958)
Araneidae
Araneus diadematus 12.0 8.0 0.3 Yes Elgar and Nash (1988)
Argiope aemuZa 22.0 4.5 1.5 Yes Robinson and Robinson (1980)
Argiope aetheria 14.0 5.0 0.9 Yes Robinson and Robinson (1980)
Argiope appensa 22.0 6.0 1.5 Yes Present study
Argiope argentata 16.0 4.0 0.9 Yes Robinson and Robinson -<1980)
Argiope aurocincta 0.9 Yes Robinson and Robinson (1980)
Argiope bruennichi 15.0 4.5 0.3 . Yes Bristowe (1958) "

Argiope cuspidata 0.9 Yes Robinson and Robinson (1980)

ArgiopejlavipaZpis 14.0 5.5 0.9 No Robinson and Robinson (1980)
Argiope jlorida 16.0 4.5 0.9 Yes Robinson and Robinson (1980)
Argiope ocyaZoides 9.0 4.0 0.5 Yes Robinson and Robinson (1980)
Argiope picta 19.0 5.5 0.9 Yes Robinson and Robinson (1980)
Argiope radon 18.0 6.0 0.9 Yes Robinson and Robinson (1980)
Argiope reinwardti 18.0 4.5 0.9 Yes Robinson and Robinson (1980)
Argiope savignyi 14.0 3.5 0.9 No Robinson and Robinson (1980)
Cyclosa caroZi 6.0 3.0 0.4 No Robinson and Robinson (1980)
Cyclosa insuZana 6.5 3.5 0.4 No Robinson and Robinson (1980)
Cyclosa sp. alpine # 1 0.4 No Robinson and Robinson (1980)
Cyclosa sp. Mt Kaindi # 1 0.4 No Robinson and Robinson (1980)
Cyclosa sp. Wau # 5 1.2 No Robinson and Robinson (1980)
Cyrtophora moZuccensis 19.0 3.5 33.0 Yes Berry (1987)
Cyrtophora nympha 0.9 No Robinson and Robinson (1980)
Eriophora fuliginea 30.0 27.0 0.4 Yes Robinson and Robinson (1980)
Gasteracantha brevispina 9.0 2.5 73.0 No Robinson and Robinson (1980)
Gasteracantha cancriformis 7.2 2.2 69.0 No Robinson and Robinson (1980)
10 M.A. EIgar

Taxa Body length Copulation Sexual Source

(mm) duration cannibal
Male Female (min)

Gasteracantha curvispina 40.0 No Robinson and Robinson (1980)

Gasteracantha jalcicornis 49.0 No Robinson and Robinson (1980)
Gasteracantha minax 8.5 3.0 100.0 No Present study
Gasteracantha signifera 54.0 No Robinson and Robinson (1980)
Gasteracantha sp. 13.0 No Robinson and Robinson (1980)
Gasteracantha taeniata 53.0 No Robinson and Robinson (1980)
Gasteracantha theisi 65.0 No Robinson and Robinson (1980)
Gasteracantha versicolor 74.0 No Robinson and Robinson (1980)
Gea eff 6.6 3.0 0.9 Yes Robinson and Robinson (1980)
Isoxya tabulata 75.0 No. Robinson and Robinson (1980)
Mecynogea lemniscata 33.0 No Robinson and Robinson (1980)
Micrathena sagittata 8.5 4.5 0.9 No Robinson and Robinson (1980)
Micrathena schreibersi 14.3 5.4 2.0 No Robinson and Robinson (1980)
Micrathena sexpinosa 16.2 5.6 1.2 No Robinson and Robinson (1980)
Zilla Wau# 1 0.4 Yes Robinson and Robinson (1980)
ZillaWau#2 0.4 Yes Robinson and Robinson (1980)
Clubionidae
Clubiona cambridgei 8.0 7.0 44.0 Rare Pollard and Jackson (1982)
Clubiona reclusa 7.0 6.0 60.0 No Bristowe (1958)
Clubiona terrestris 6.0 5.0 80.0 No Bristowe (1958)
Dictynidae
Dictyna calcarata 5.0 4.0 60.5 Yes Jackson (1979)
Dictyna civica 14.0 Jackson (1979)
Dictyna latens 15.0 Jackson (1979)
Dictyna uncitata 40.0 Jackson (1979)
Dictyna volucripes 3.3 2.7 87.0 No Starr (1988)
Dictyna volupis 3.8 3.1 80.0 No Jackson (1979)
Ixeuticus longiquus 80.0 Jackson (1979)
Mallos gregalis 5.0 4.0 100.0 No Jackson (1979)
Mallos trivittatus 7.0 5.0 39.0 No Jackson (1979)
Gnaphosidae
Drassodeslapidosus 14.0 11.5 31.0 No Bristowe (1958)
Herpyllus blackwalli 10.0 8.0 24.0 Rare Bristowe (1958)
Heteropodidae
Heteropoda venatoria 23.0 20.0 160.0 Rare Ross et al. (1982)
Micrommata virescens 13.5 9.0 280.0 No Bristowe (1958)
Linyphiidae
Frontinella pyramitela 3.5 3.2 30.7 Rare Austad (1982)
Lepthyphantes leprosus 3.0 2.5 49.0 No van Helsdingen (1965)
Linyphia litigiosa 4.5 3.5 240.0 No Watson (1991)
Lycosidae
Alopecosa accentuata 12.0 9.0 0.5 No Bristowe (1958)
Lycosa malitiosa 98.0 No' Costa and Sotelo (1984)
Lycosa rabida 18.5 11.5 59.0 No Rovner (1972)
Schizocosa saltatrix 166.0 No Rovner (1974)
Xerolycosa miniata 6.0 5.0 0.5 No Bristowe (1958)
Metidae
Herennia ornatissima 12.0 3.0 6.3 No Robinson and Robinson (1980)
Metellina segmentata 6.0 5.0 2.7 Yes Prenter et al. (1994)
Nephila clavipes 25.0 6.0 25.6 Yes Christenson et al. (1985)
Nephila plumipes 22.0 5.0 0.9 Yes Present study
Nephila maculata 43.0 5.0 17.0 Yes Robinson and Robinson (1980)
Nephila pilipes 40.0 4.5 20.0 No Robinson and Robinson (1980)
Nephila senegalensis 24.7 4.6 8.0 No Clausen (1987)
Nephilengys malabarensis 16.7 4.8 1.5 No Robinson and Robinson (1980)
Nesticidae
Nesticus cellulanus 5.0 4.0 7.0 No Bristowe (1958)
Pisauridae
Dolomedes jimbriatus 22.0 12.0 0.9 Yes Arnqvist (1992)
Pisaura mirabilis 13.5 11.0 58.0 Yes Bristowe (1958)
Duration of copulation in spiders 11

Taxa Body length Copulation Sexual Source

(mm) duration cannibal
Male Female (min)

Tetragnathidae
Phonognatha graeffei 8.0 5.0 111.0 Rare Fahey (1992)
Salticidae
Asemonea tenuipes 4.5 3.5 4.4 Jackson and Macnab (1991)
Cobanus mandibularis 7.1 No Jackson (1989)
Euophrys frontalis 4.0 3.0 45.0 No Bristowe (1958)
Evarcha falcata 7.0 5.0 200.0 Bristowe (1958)
Goleba puella 4.5 4.5 1.3 Jackson and Macnab (1991)
Holoplatys sp. 5.0 4.4 17.0 No Jackson and Harding (1982)
Jacksonoides queenslandica 6.0 6.0 3.1 No Jackson (1988)
Lyssomanes viridis 7.5 5.5 14.5 No Jackson and Macnab (1991)
Marpissa muscosa 8.0 8.0 20.0 Bristowe (1958)
Marpissa nivoyi 5.0 4.0 47.0 No Bristowe (1958)
Menemerus sp. 3.5 3.5 9.0 No Jackson (1986)
Mopsus mormon 12.0 12.0 44.4 No Jackson (1983)
Myrmarachne lupata 5.0 5.0 10.9 No Jackson (1982a)
Phiddipus femoratus 10.0 7.0 12.0 No Jackson (1982c)
Phiddipus johnsoni 9.0 8.0 179.0 Yes Jackon (1980)
Plexippus paykulli 0.7 Yes Jackson and Macnab (1989)
Portia fimbriata 8.5 5.5 6.0 Yes Jackson (1982b)
Portia sp. 1.7 Yes Jackson and Hallas (1986)
Pseudicius sp. #1 3.5 3.5 9.0 No Jackson (1986a)
Pseudicius sp. #2 3.5 3.5 900.0 No Jackson (1986a)
Simaetha paetula 7.0 7.0 12.7 Yes Jackson (1985)
Thorellia ensifera 5.0 5.0 5.8 No Jackson and Whitehouse (1989)
Theridiidae
Achaearanea lunata 3.0 2.5 0.2 Bristowe (1958)
Achaearanea tepidariorum 7.0 4.0 0.2 Bristowe (1958)
Achaearanea wau 4.5 1.9 0.7 No Lubin (1986)
Argyrodes antipodiana 3.0 2.5 21.3 Whitehouse (1991)
Enoplagnatha ovata 5.0 4.0 15.0 Bristowe (1958)
Episinus truncatus 4.0 4.0 5.0 Bristowe (1958)
Latrodectus hasselti 10.0 5.0 27.0 Yes Forster (1992)
Steatoda bipunctata 7.0 5.0 100.0 No Bristowe (1958)
Steatoda nobilis 12.0 9.2 25.0 Snazell and Jones (1993)
Thomisidae
Diaea dorsata 6.0 4.0 5.0 Bristowe (1958)
Diaea socialis 6.6 5.6 30.0 No T. Evans (pers comm)
Misumenoides formosipes 4.4 Dodson and Beck (1993)
Philodromus dispar 5.0 4.0 0.5 Bristowe (1958)
Philodromus fallax 6.0 5.0 4.0 Bristowe (1958)
Xysticus cristatus 7.0 5.0 90.0 No Bristowe (1958)
Xysticus lanio 7.0 5.0 110.0 Bristowe (1958)
Uloboridae
Uloborus walckenaerius 5.0 3.5 10.0 No Bristowe (1958)
Zoridae
Zora spinimana 6.0 4.5 6.0 Bristowe (1958)