ARTICLE IN PRESS

ANIMAL BEHAVIOUR, 2007, --, --e-doi:10.1016/j.anbehav.2006.10.017

Male mate choice and size-assortative pairing in

a jumping spider, Phidippus clarus
C HA D D. H O E FL E R

Organismic and Evolutionary Biology and Department of Plant, Soil and Insect Sciences,
Division of Entomology, University of Massachusetts, Amherst
(Received 18 January 2006; initial acceptance 14 May 2006;
nal acceptance 27 October 2006; published online - - -; MS. number: A10344R)

Male choice is receiving greater attention from biologists, but remains understudied compared to investigations of female choice. I studied male choice and its potential role in size-assortative pairing in the jumping spider Phidippus clarus. Adult males cohabit with and guard immature females shortly before they
mature. During this period, I monitored the sex ratio and maturation rate of females in the eld, because
these parameters can inuence conditions favouring male choice. The adult sex ratio was initially male
biased and became female biased following a relatively synchronous female maturation period. I collected
maleefemale pairs from the eld and found that they were size-assortatively paired for tibia length. I then
tested whether males could discriminate between females by size, using tibia length as a measure of size.
Both small and large males preferred to pair with large females that matured sooner compared with small
females that matured later. Males also discriminated between females when exposed only to female silken
nests, suggesting that cues associated with silk convey information about maturation and tibia length, two
characters that were highly inversely correlated. Finally, large males were more likely to defeat small males
in maleemale contests, and as the size disparity between contestants increased, the likelihood that the
larger individual would win also increased. A male preference for larger females combined with a large
male advantage in maleemale contests is likely to strongly inuence size-assortative pairing. Species
that are known to mate or pair assortatively may be valuable for studies of male choice.
2007 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Keywords: breeding synchrony; ghting; jumping spider; maleemale contests; mate choice; mate guarding; Phidippus
clarus; size-assortative pairing

Darwin (1871) put forth the idea that males are competitive
and females are choosy in mating behaviour. Our understanding of sex roles was later reinforced by Batemans
(1948) classic study on Drosophila melanogaster. However,
the DarwineBateman paradigm of competitive males and
choosy females may be too simplistic. First, in species where
male parental investment is greater than that of females,
typical sex roles are reversed (Trivers 1972; Andersson
1994). Second, although single sperm are relatively less expensive to produce than single eggs, spermatozoa are delivered not individually but by the millions, and the ejaculate
also contains potentially costly accessory-gland secretions
(Dewsbury 1982). This evidence, coupled with mate

Correspondence and present address: C. D. Hoeer, Department of

Zoology, Miami University, Oxford, OH 45056, U.S.A. (email:
hoeecd@muohio.edu).
0003e 3472/07/$30.00/0

searching, courtship, nuptial gifts, mate guarding, mating,

sperm competition and risks of predation, necessarily increases the costs of male reproduction (Dewsbury 1982;
Andersson 1994; Bonduriansky 2001). Third, if a male cannot mate with all reproductive females in the population,
he should benet by choosing females capable of producing more offspring (Altmann 1997). In that case, we
should expect males to choose females based on features
that indicate potential reproductive success, such as body
size (Gwynne 1991; Andersson 1994; Owens & Thompson
1994).
There is not a strict dichotomy between choosiness and
competition: even the more competitive sex may also
show mate choice. This selectivity has been shown in
numerous systems in which males appear to make no
material contribution to their offspring, including crustaceans (Jormalainen et al. 1994), insects (van Dongen et al.
1998; Harari et al. 1999), sh (Sargent et al. 1986),

1
2007 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Please cite this article in press as: Chad D. Hoeer, Male mate choice and size-assortative pairing in a jumping spider, Phidippus clarus, Anim. Behav. (2007),
doi:10.1016/j.anbehav.2006.10.017

ARTICLE IN PRESS
2

ANIMAL BEHAVIOUR, --, -

amphibians (Arak 1983; Arntzen 1999), reptiles (Shine

et al. 2001), birds (Saether et al. 2001; Griggio et al.
2005) and mammals (Smuts 1987; Preston et al. 2005).
Nevertheless, relatively few have studied male mate choice
compared to the substantial research effort devoted to
female mate choice (Andersson 1994; Bonduriansky
2001). Consequently, there is a widely held view that
the intricacy of female choice far exceeds that of male
choice, but it is unknown whether this is an accurate perception of sexual differences or a side effect of the paucity
of studies focused on males (Shine et al. 2003). We therefore need a better understanding of male choice, including
the circumstances and mechanisms by which males might
be selective.
Male choice has also been implicated as a mechanism
for size-assortative mating or pairing (Ridley 1983), a very
common pattern (Crespi 1989a, b). Size-assortative mating (i.e. large males mating with large females, small males
with small females) has piqued the interest of biologists
because of its inuence in sexual selection and the evolution of mating systems and its striking effects on the genetic variation and demographics of populations (Wade
& Arnold 1980; Crespi 1989a; Parker & Partridge 1998).
Male choice can help to generate this pattern if (1) there
is a combined occurrence of male choice for large females
with a large-male advantage in intrasexual contests
(Ridley 1983), (2) the sexes mutually favour larger mates
(Burley 1983) or (3) male choice is size dependent; that
is, large males prefer large females because they are highly
fecund and defensible, and small males prefer small females because the likelihood of mating with large females
is remote. In some cases, these mechanisms may be mutually reinforcing (explanations 1 and 2 above) or mutually
incompatible (explanations 1 and 3 or 2 and 3 above). Understanding how male choice inuences size-assortative
mating can better our understanding of the evolution of
sex roles.
Phidippus clarus (Araneae: Salticidae) is a ubiquitous, sexually dimorphic jumping spider that inhabits old elds
throughout North America. Jumping spider behaviour
has been extensively studied. Jackson and colleagues alone
have published over 150 studies covering myriad topics including prey capture (Taylor et al. 1998), interactions between predators and prey (Jackson et al. 1998), ghting
ability (Clark et al. 1999), aggressive mimicry (Jackson &
Carter 2001), nectar feeding (Jackson et al. 2001), chemosensory responses (Jackson et al. 2002) and araneophagy
(Harland & Jackson 2004). Phidippus clarus, like other jumping spiders, is a diurnal and highly visual organism (Land
1972) with corresponding vision-based behaviour. For example, vision is critical for predation (e.g. Jackson 2000;
Li et al. 2003), courtship and mate choice (e.g. Clark &
Morjan 2001), and during aggressive encounters (e.g. Wells
1988; Taylor et al. 2001). Phidippus clarus also builds silken
nests that are used, for example, when mating, ovipositing
and guarding young (Hoeer & Jakob 2006), and it is
known to reuse the nests of Hibana gracilis (Araneae: Anyphaenidae; Johnson 1995). Phidippus clarus has the most
restricted reproductive cycle of the three predominant congeners (P. audax, P. clarus, P. princeps; Roach 1988), mating in
early to mid-July and ovipositing in August.

Phidippus clarus is an interesting species in which to expect male choice and size-assortative mating. My preliminary observations indicated that adult males visit and
guard penultimate female conspecics (i.e. the instar
before reproductive adult) over a period of several weeks
before female maturation. Precopulatory guarding systems
such as this are common and can be indicative of
male choice, because males may show differential sexual
responses towards females varying in mate quality
(Bonduriansky 2001).
Inherent in all precopulatory guarding systems is the
active defence of a prospective mate. Adult P. clarus males
ght for the access to guarding positions, and males in
guarding positions are often near females (e.g. below the
nest, in an adjoining silken nest). Multiple males do not
appear to guard the same female as has been documented
in other spiders that show precopulatory guarding (e.g.
Dodson & Beck 1993). A large-male advantage in contests
over mates has been widely documented (Andersson
1994); therefore, smaller males may be forced to defend
smaller females. Thus, it seemed reasonable to assume
that the P. clarus precopulatory guarding system satised
conditions favouring both male choice and size-assortative
pairing. Lastly, this precopulatory guarding system also
seemed to satisfy conditions for partial sex-role reversal
(Gwynne 1991; Bonduriansky 2001). Species with partial
sex-role reversal show variations on the DarwineBateman
pattern, such as male mate choice with maleemale competition and/or female mate choice. The frequency of
occurrence of partial sex-role reversal is unknown.
My aims in this study were (1) to monitor the number of
males and females over the course of the P. clarus reproductive cycle, because sex ratio can establish favourable conditions for male choice to occur, (2) to document and
quantify size-assortative pairing in the eld, because male
mate choice is a mechanism that can inuence this pattern,
(3) to explore how male choice may be operating and (4)
to examine the inuence of body size on success in
maleemale contests, because a large-male advantage coupled with male mate choice can produce size-assortative
pairing.

METHODS AND RESULTS

Study Sites
All eld work was conducted from 2001 to 2004 on or
near the campus of the University of Massachusetts,
Amherst, U.S.A., in old elds that contained a variety of
grasses, shrubs and owering plants, predominantly milkweed, Asclepias syriaca, goldenrod, Solidago spp., sweet
clover, Melilotus spp., and asters, Aster spp. In 2005,
maleemale contests were staged at the Ecology Research
Center of Miami University, Oxford, Ohio, U.S.A.

Experiment 1: Sex Ratio in the Field

Methods
Mating systems are inuenced largely by factors such as
sex ratio, so I monitored two population parameters:

Please cite this article in press as: Chad D. Hoeer, Male mate choice and size-assortative pairing in a jumping spider, Phidippus clarus, Anim. Behav. (2007),
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HOEFLER: MALE CHOICE IN JUMPING SPIDERS

(1) the number of males and females and (2) the

maturation rate of females. On 7e8 June 2001, I set up
a 30  30-m grid of 90-cm-tall surveyors ags (Ben
Meadows, Janesville, Wisconsin, U.S.A.) spaced 2 m apart.
Phidippus clarus readily constructs nests inside pieces of
tubing (Popson 1999; Hoeer & Jakob 2006). I made articial nest sites of exible clear plastic plumbers tubing
(about 3.8 cm long  1.5 cm diameter) and painted them
black with spray paint (Krylon at black, Sherwin-Williams, Cleveland, Ohio, U.S.A.). I tied tubes to agpoles
with string so that the tubes hung horizontally about
40 cm above the ground. I then allowed a 3-week colonization period for individuals to nd nest tubes and construct silken nests within them (see also Hoeer & Jakob
2006).
During the rst 3 weeks in both July and August 2001, I
checked nest tubes for occupancy daily between 1900 and
2000 hours. I recorded the sex of the individual(s) present.
For females, I noted whether they were penultimate
instars or adults, which can be distinguished by marked
differences in coloration patterns and size. To test the null
hypothesis that the ratio of males to females did not differ
signicantly from 1:1 for each day, I conducted goodnessof-t tests.

Results
As represented by the population of spiders in articial
nest tubes, the adult P. clarus population sex ratio was signicantly male biased for the rst 2 days of observations
(goodness-of-t test: day 1: c21 54:5, N 66, P < 0.001;
day 2: c21 46:1, N 68, P < 0.001; Fig. 1a). On day 3,
the sex ratio did not signicantly differ from 1:1
(c21 3:11, N 93, P 0.078). On day 4, the sex ratio
was signicantly female biased (c21 6:69, N 109,
P < 0.01). For the remaining dates in July and August
2001, the population remained signicantly female biased
(all PS < 0.001; Fig. 1a, b). I found no males during August.
Three females were adult on the rst day of observations
(3.7%). By day 4, 73.9% of females were mature; by day
5, 96.2% of females were mature. After day 10, 100% of
observed females were mature for July and August 2001
(Fig. 1c). Thus, the operational sex ratio, OSR (i.e. the ratio
of fertilizable females to sexually active males; Emlen &
Oring 1977) changed rapidly from completely male biased
to completely female biased.

Experiment 2: Size-assortative Pairing

Methods
In early June 2002, I placed surveyor ags with articial
nest tubes in two adjacent elds as described above.
I allowed male and female P. clarus 3 weeks to locate
nest tubes and form pairs. On 3 July 2002, I collected every pair of adult male and penultimate female spiders that
had colonized the nest tubes from both elds. In the
laboratory, I anaesthetized individuals of each pair using
carbon dioxide gas and recorded the following measurements: tibia length (right anterior), cephalothorax (head)
width, abdomen length and abdomen width. Linear
measurements were made (0.01 mm) using dial calipers

under a dissecting microscope. I did not record mass,

which changes rapidly with evaporative uid loss in other
spiders (Pollard et al. 1995). I tested for the correlation between all permutations of male and female body measurements for all pairs using linear regression. I also regressed
cephalothorax width (which does not change for the duration of an instar) with abdomen width (which is a exible character) for both males and females. I regressed the
residual values of this analysis for males and against that
of females to examine whether body condition contributes to pairing patterns (Jakob et al. 1996).

Results
Size-assortative pairing was characteristic of pairs under
eld conditions (N 39). Male and female tibia lengths
(a measure of absolute size) were positively correlated
(linear regression: R2 0.294, P 0.0004; Fig. 2). No other
permutation of body measurements was signicantly correlated. Likewise, individuals were not assortatively paired
by body condition (R2 0.01, P 0.55).

Experiment 3: Male Choice: Females

and Female Silk Present
Methods
I staged simultaneous choice tests where adult males of
a particular size class were presented with two penultimate
females of different sizes. During the rst 2 weeks of June
2003, I collected penultimate-instar males and reared
them to adulthood in the laboratory. Experience with
members of the opposite sex can inuence mate choice;
for example, subadult experience affects adult choice in
wolf spiders (Hebets 2003). Therefore, I collected penultimate-instar males because they do not associate with
females. Males were maintained individually in plastic
cages (18  13  11 cm) with articial nest tubes on
a 13:11 h light:dark cycle at approximately 26 C. I provided spiders with approximately ve early instar crickets
(Acheta domesticus, Top Hat Cricket Farm, Kalamazoo,
Michigan, U.S.A.) per week, and provided water ad libitum
in test tubes plugged with cotton. In mid-June 2003, I collected penultimate-instar females and maintained them as
described above with one exception. Because cues indicating closeness to female maturity could potentially confound the interpretation of the results, I continued to
maintain females until they matured. I recorded the number of days until the ultimate moult for each female and
treated it as an uncontrolled variable in the analysis. Cages
housing males and females were kept out of view of one
another, and blinders constructed of white construction
paper (Art Street, Riverside Paper Company, Mount Pleasant, Iowa, U.S.A.) were placed between cages housing
members of the same sex to prevent individuals from
interacting visually. I recorded tibia length and categorized all individuals as small, medium or large. I made
these categorizations based on tibia size alone, because I
conrmed size-assortative pairing for tibia length only.
Within each sex, small individuals were those with
tibia lengths in the lowest quartile range (X  SE:
males 2.41  0.024 mm; females 1.06  0.014 mm),

Please cite this article in press as: Chad D. Hoeer, Male mate choice and size-assortative pairing in a jumping spider, Phidippus clarus, Anim. Behav. (2007),
doi:10.1016/j.anbehav.2006.10.017

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ANIMAL BEHAVIOUR, --, -

120
(b)

(a)
105
Number of individuals

90
75
60
45
30
15
0
0

10 12 14 16 18 20 22
July 2001

8 10 12 14 16 18 20 22
August 2001

120
(c)
105
Number of individuals

90
75
60
45
30
15
0

10

July 2001
Figure 1. Numbers of adult male (B) and female (C) jumping spiders observed in artificial nest tubes in the field in (a) July 2001 and (b)
August 2001. (c) Numbers of adult (,) and penultimate (-) female instars observed in July 2001.

medium individuals had tibia lengths within the middle

two quartile ranges (males 3.05  0.049 mm, females
1.55  0.031 mm) and large individuals had tibia lengths
in the upper quartile range (males 3.69  0.021 mm,
females 2.04  0.022 mm). I used only small and large
individuals, to maximize my ability to detect size-dependent male choice if it occurred.
Small and large adult males were randomly assigned to
a pair of penultimate females, one large and one small. To
simulate outdoor conditions, simultaneous choice tests
were conducted in an old eld on 1e4 July 2003. Each test
male was placed in a clear, circular plastic cage (10 cm
diameter, 6.5 cm height) with two holes (1.5 cm diameter)
cut on opposite sides of the cage walls. Test females were
placed in clear plastic cages (17  12.5  6 cm) with one
hole (1.5 cm diameter) cut on one side of the cage. Each
females nest tube was taped to the ceiling/lid of the
cage, because P. clarus show negative geotaxis (personal
observation). One end of a 30-cm piece of clear plumbers
tubing (1.5 cm diameter) was tted into the hole of each

female cage so that the end of the tubing was ush with
the cage wall. The opposite end was tted similarly into
one of the holes of the male cage. Thus, the male cage
sat between the female cages (Fig. 3). None of the cages
had previously held spiders. Spiders were introduced to
their respective cages after the apparatus had been set
up. All individuals could potentially have moved to any
cage in this set-up; however, in a previous study, females
moved less than males and often remained in their silken
nests, but adult males moved frequently and over great
distances (Hoeer & Jakob 2006). On day 1, half of the
test males were placed in the experimental set-up with
a pair of females between 1000 and 1130 hours. After
24 h, I returned to the eld and recorded the location of
the male and the size class of the selected female. I considered areas of no choice to include the centremost (i.e.
male) cage and the connected tubing, because the male
could not interact with either female. I treated the presence of a male in the cage of a female as a choice. Female
cages and their connected tubing were then switched from

Please cite this article in press as: Chad D. Hoeer, Male mate choice and size-assortative pairing in a jumping spider, Phidippus clarus, Anim. Behav. (2007),
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16
Small female
14

Large female

12

3.5
Male tibia (mm)

(a)

10
8
3
6
4
2.5

1.4

1.6
1.8
Female tibia (mm)

2.2

Frequency

2
1.2

2
0
16
Female that moulted
second
Female that moulted
first

14

Figure 2. Correlation between adult male tibia length and penultimate female tibia length (Pearson correlation: r 3 7 0.542,
P 0.0004).

(b)

12
10

end to end, males were returned to their cages, and I allowed another 24-h choice period to control for potential
side biases. On days 3 and 4, I repeated these steps with
the remaining half of the spiders. I analysed the dichotomous data via goodness-of-t tests. After females moulted
to adulthood in the laboratory, I reanalysed the male
choice data via goodness-of-t tests using the categories
moulted rst and moulted second for each pair of females. I also tested for a correlation between female tibia
length and moulting date.

8
6
4
2
0

Results

(c)

1.5

0.5
6

Nest tube

Male enclosure
Female enclosure
Figure 3. Experimental set-up for simultaneous choice tests conducted in the field.

Large male

2.5

Female tibia (mm)

Results supported the hypothesis that male choice

affects assortative pairing. Both large and small males
chose large females over small females (goodness-of-t
test: large males, test 1: c21 7:12, N 17, P < 0.01; large
males, test 2: c21 4:76, N 17, P < 0.05; small males,
test 1: c21 4:00, N 16, P < 0.05; small males, test 2:
c21 6:25, N 16, P < 0.025; Fig. 4a). Reanalysed data
showed that both large and small males chose females
that moulted rst over females that moulted second in
both tests (large males, test 1: c21 7:12, N 17,
P < 0.01; large males, test 2: c21 4:76, N 17, P < 0.05;
small males, test 1: c21 4:00, N 16, P < 0.05: small
males, test 2: c21 6:25, N 16, P < 0.025; Fig. 4b).
Female body size and moulting date were signicantly

Small male

8
9
10
11
12
Date of maturity (July 2003)

13

14

Figure 4. (a) Number of small and large adult males choosing small
or large penultimate females in 2003. (b) Reanalysed data from
Fig. 4a, grouping females by when they moulted to adulthood.
(c) Correlation between small and large penultimate female tibia
length and date of maturity in July 2003.

Please cite this article in press as: Chad D. Hoeer, Male mate choice and size-assortative pairing in a jumping spider, Phidippus clarus, Anim. Behav. (2007),
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ANIMAL BEHAVIOUR, --, -

inversely correlated (Pearson correlation: r64 0.916,

P < 0.0001; Fig. 4c).

Experiment 4: Male Choice: Only Female

Silk Present
Methods
In experiment 3, choice could have been based on
either female tibia size or a putative cue indicating
closeness to maturity. To test the potential inuence of
cues associated with silk in P. clarus, I conducted simultaneous choice tests as described for experiment 3 with the
following difference. Instead of placing females in their respective test cages, I used the nest tube only, which came
from the cages that had housed females in the laboratory.
All nest tubes contained silken nests. I was concerned that
potential chemical cues from silk may dissipate over time,
so I checked for male choice after 12 and 24 h. Moreover, I
did not test for side biases because males did not show
a side bias in experiment 3. Lastly, I included spiders
from all size classes in the tests, because I was unable to
collect as many spiders as in the previous year. Small,
medium and large males were randomly assigned to
a pair of females consisting of one relatively large female
and one relatively small female (e.g. a medium and small
female pair or a large and medium female pair).

Results
Results were consistent with the hypothesis that male
choice inuences size-assortative pairing. After 12 h, 24
males made a choice and ve males did not. Males, and
the corresponding female pairs, that did not make a choice
after 12 h were excluded from further tests. After 12 h,
males chose nest tubes of females that moulted rst over
those that moulted second (goodness-of-t test:
c21 4:17, N 24, P < 0.05; Fig. 5a). Males did not discriminate between females on the basis of tibia length
(c21 2:67, N 24, P 0.1; Fig. 5b), but statistical power
was low (53.6%). After 24 h, 19 males made a choice
and ve males did not. Males that made a choice did
not show a signicant preference based on either proximity to moult (c21 2:58, N 19, P 0.11; Fig. 5c) or size
(c21 1:32, N 19, P 0.11; Fig. 5d). Female body size
and moulting date were signicantly inversely correlated
(Pearson correlation: r46 0.885, P < 0.0001; Fig. 5e).

Experiment 5: MaleeMale Contests

Methods
Adult males did not show size-dependent pairing preferences; thus, this mechanism of size-assortative pairing
can be eliminated. However, I tested the inuence of male
size on ghting outcome, because a large-male advantage
in contests, coupled with the nding that males preferred
large females, could produce size-assortative pairing. In
late June 2005, I staged maleemale contests in an old eld
at the Ecology Research Center of Miami University. All
contests were conducted in the eld, and males were
collected from nest tubes as described earlier. Males were

randomly paired and placed in clear plastic snap-cap vials

(5.5  2.5 cm diameter). Black electric tape was placed
around each vial to prevent males from visual contact
before trials began.
To conduct a trial, I placed one end of a dowel rod
(90 cm long  0.8 cm diameter) into the ground. I slid
a deli cup lid (12 cm diameter) with a slit cut in the middle
onto the dowel, and used the lid as a stage for the contests.
I secured the stage by placing a small amount of polymer
clay below the slit where the dowel was inserted. Some
deli cup lids were reused during the experiment, and these
were thoroughly cleaned using 75% ethanol. Vials with
male spiders were inverted so that the lid faced the
bottom, which caused individuals to ascend the upsidedown vial. After a 5-min acclimation period, I removed
lids, inverted the vials open end up, and placed the vials
about 5 cm apart on the stage. Males interacted, and a trial
ended when one spider (loser) ed from the other (winner). Individuals were collected and preserved in 75% ethanol to measure tibia lengths. I measured tibia lengths in
the laboratory using a dissecting microscope tted with an
electronic lar micrometer. To evaluate the inuence of
size and the probability that larger individuals win contests against smaller rivals, I used logistic regression
models using the size difference of rivals as the predictor
of the categorical response larger male wins (Hardy &
Field 1998; Taylor & Jackson 2003).

Results
Of 27 staged contests, 21 yielded usable data. Data from
six contests could not be used because I was unable to
capture one or both of the individuals after the contest
ended. Logistic regression revealed a positive effect of size
difference between rivals on the probability that the larger
individual would win (N 21, b 9.18, R2 0.36,
G2 9.02, P 0.0027; Fig. 6).

DISCUSSION
Evidence from both eld and laboratory studies suggests
that male mate choice is important in the jumping spider
P. clarus, a species with partial sex-role reversal. In the
eld, I documented a dramatic swing from a male-biased
to a female-biased adult sex ratio in only a few days.
This situation sets the stage for sexual selection: at the
start of the season, males have the opportunity to be
choosy, because penultimate females, but not adult females, are common. Males should also be competitive,
because females mature relatively synchronously, which
can constrain male reproductive success if females mate
only once. I also documented size-assortative pairing in
the eld. I reject the hypothesis that size-assortative pairing arises because male mate choice is size dependent: in
eld experiments, males of all sizes preferred larger
females that matured sooner over smaller females that
matured later. Cues associated with silk appeared to be important to males, because these cues conveyed information regarding female maturation and size. It is likely
that male choice, in conjunction with a large-male

Please cite this article in press as: Chad D. Hoeer, Male mate choice and size-assortative pairing in a jumping spider, Phidippus clarus, Anim. Behav. (2007),
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HOEFLER: MALE CHOICE IN JUMPING SPIDERS

18

18
(b)

16

16

14

14

12

12
Frequency

Frequency

(a)

10
8

10
8

Female moult second

Female moult first

14

Larger female

14
(d)

12

12

10

10
Frequency

Frequency

(c)

8
6

8
6

Smaller female

Female moult first

Female moult second

Larger female

Smaller female

2.5
(e)

Female tibia (mm)

1.5

0.5
8

10 11 12 13 14 15 16
Date of maturity ( July 2004)

17

18

Figure 5. Number of males exposed to female silk only for 12 h that chose (a) either penultimate females that moulted to maturity sooner or
those that matured later and (b) either larger or relatively smaller penultimate females. Number of males exposed to female silk only for 24 h
that chose (c) either females that moulted to maturity sooner or those that matured later and (d) either relatively larger or relatively smaller
penultimate females. (e) Correlation between penultimate female tibia length and date of maturity in July 2004.

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Probability larger male wins

0.75

0.5

0.25

0
0

0.25

0.5

0.75

1.25

Tibia length difference

Figure 6. Relationship between the size difference between adult
P. clarus males and the probability that the larger male would win.
The line represents the predicted probabilities derived from a logistic
regression. Points above the line indicate wins by the larger male;
points below the line indicate wins by the smaller male.

advantage in maleemale contests, leads to size-assortative

pairing. I discuss each of these ndings in more detail
below.

Sex Ratio in the Field

On the rst 2 days of observations, adult sex ratios were
male biased for P. clarus individuals that were monitored at
nest tubes, but then shifted to a female bias in July and
August 2001, with no males found in August 2001. This
change can be explained by the abrupt decline of males
from the population, which may have occurred for several
reasons. I occasionally (N 18) observed sexual cannibalism (i.e. females consuming males) throughout the month
of July, but only after females matured. The decline of
males also coincided with a fairly synchronous maturation of females; some males may have emigrated to other
elds or were differently attracted to nest tubes after
females matured. The life span of males may have also
been affected by female assessment and/or courtship: in
a laboratory experiment, males with courtship and/or
mating experience had signicantly lower survivorship
than males without courtship and mating experience,
and no males with courtship experience survived into
August (unpublished data). Similarly, in an earlier study,
no adult males were collected (via sweep netting) in
August (Hoeer & Jakob 2006). These results are consistent with the eld observation that a substantial number
of males were found dead (seemingly uninjured) within
articial nest tubes unoccupied by females. Sex-ratio calculations were based on individuals monitored at nest
tubes only. Although nest tube monitoring is not a perfect
method for reasons listed above, it is likely to provide
a fair sense of the numbers of males and females. In the
laboratory, both females and males readily accept nest
tubes in enclosures (personal observation). Similarly,
nest tubes are promptly used by P. clarus in the eld
(Hoeer & Jakob 2006), suggesting that they are valuable

to both males and females. Plett (1962) reported that salticids will often ght for access to ideal nest site locations.
However, only about 50% of tubes were used by spiders
during the monitoring of this population; thus, they
were not a limiting resource. Also, nest tubes were checked
at dusk, when jumping spiders are known to return to
them (Popson 1999; Hoeer & Jakob 2006), so it seems
likely that the sex ratio observed in the nest tubes was reasonably representative.
Sexual selection theory predicts that, when sex ratios
are not highly female biased and if males can detect the
moult stage of the female, male mate guarding can evolve
(Grafen & Ridley 1983; Ridley 1983). Although the P. clarus sex ratio became female biased, because this species is
protandrous, the OSR was perfectly male biased during the
precopulatory guarding phase. When OSRs are female
biased, male milkweed beetles, Tetraopes tetraophthalmus,
become increasingly choosy (Lawrence 1986). When
OSRs are unbiased, assortative mating is expected to be
possible and even very intense (Harari et al.1999). The
OSR for P. clarus appears to switch rapidly from male
biased to female biased during and following female maturity. However, calculating the OSR exactly is difcult once
females begin to mature, because it is unknown whether
females mate with one or more than one male. If females
mate once, then fertilized females should be excluded
from OSR estimates. However, given the rapid decline in
males in the population, the general pattern of a switch
to a female-biased OSR holds.
The rapid decline of the male population shortly after
female maturation and the behaviours of males (i.e.
precopulatory guarding) suggest that virgin females are
valuable and that the prospect for multiple mating is
remote. Male salticids typically leave apparent mating
plugs in the females genital openings after mating (Taylor
& Jackson 1999), and these may impede the intromission
of later mates (Jackson 1981). Even if the apparent mating
plug fails to prevent insemination by second or future
mates, the rst mate may still have an advantage. Salticid
spermatheca are designed in a manner that appears to
favour priority to rst-male sperm (Austad 1984; Eberhard
et al. 1993). Each of these factors suggests considerable reproductive rewards for rst mates that might explain the
prevalence of cohabitation in these spiders and why males
would be both choosy and competitive.

Size-assortative Pairing
Phidippus clarus were assortatively paired in the eld,
based on tibia length. Comparable correlations have
been documented in natural populations of other organisms (Fairbairn 1988), including spiders (Rubenstein
1987; Masumoto 1999). What process might give rise to
this pattern? Crespi (1989b) ascribed size-assortative mating to mate choice, the interaction of male choice and
maleemale competition, constraints on pairing, habitat
selection, differential female availability by size and seasonal variation in size. I eliminated the possibility that
size-dependent male choice is solely responsible for generating this pattern. Similarly, pairing constraints were not

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evident, as males of all sizes selected large females in male

choice experiments. It also seems unlikely that differential
male or female availability by size could explain the results. Although the maturation rate of males was not measured, all males observed at the beginning and throughout
the term of this investigation were adult. Hoeer & Jakob
(2006) found that female P. clarus were more site faithful
than males: of 102 marked individuals, approximately
50% of females and 11% of males were site faithful.
Only one male returned to the same site on more than
two evenings, and that individual was paired with the
same female each evening (C. D. Hoeer & E. M. Jakob,
unpublished data). Male-guarding time with a particular
immature female, therefore, was relatively short, and
often ended before the female matured. Thus, it is very unlikely that the size-assortative pattern was due to larger,
early maturing males and females pairing and smaller,
later-maturing individuals pairing. Males move frequently
and sometimes over long distances (Hoeer & Jakob
2006), perhaps to check the maturation status of multiple
females, only guarding those very close to maturity. Some
males may have also been displaced by larger rivals.
Size-assortative pairing could have resulted from a combination of a large-male advantage in maleemale competition, which I documented, and which is common in
spiders (e.g. Christenson & Goist 1979; Austad 1983;
Watson 1990; Dodson & Beck 1993; Hack et al. 1997;
Whitehouse 1997), including jumping spider contests
(e.g. Wells 1988; Jackson & Cooper 1991), and with female
choice, which I did not test, or male choice. I examined
male choice experimentally and discuss these results in
the next section. Although I did not quantify malee
male encounter rates, maleemale contests in the eld
were frequent during the cohabitation phase.

Male Choice
Empirical studies suggest that males use a variety of
mechanisms to assess female reproductive quality including visual and tactile assessment (Bonduriansky 2001).
Male olfactory assessment of female sex pheromones is
well known in insects (e.g. Wasserman & Zweig 1991;
Wcislo 1992; van Dongen et al. 1998; Polak et al. 1998)
and spiders (e.g. Prouvost et al. 1999; Searcy et al. 1999;
Papke et al. 2001; Tichy et al. 2001; Gaskett et al. 2004;
Roberts & Uetz 2004) and is hypothesized to be the
most widespread form of mate assessment, although combinations of mechanisms are likely to be the rule rather
than the exception (Bonduriansky 2001). It is unclear
whether male P. clarus use both tactile/mechanical and
pheromonal cues, but it is clear that cues associated with
silk alone are informative. This is consistent with previous
studies of spiders. Roberts & Uetz (2005) discovered that
male wolf spiders, Schizocosa ocreata, are able to evaluate
the reproductive status and receptivity of female conspecics based on cues associated with silk. Male Pardosa milvina wolf spiders use silk and excreta cues for mate choice
and can distinguish between virgin and mated females
(Rypstra et al. 2003). Jackson (1986) demonstrated that
male P. johnsoni jumping spiders are able to discriminate

between penultimate females based on cues associated

with their silken nests. Also, some male jumping spiders
have chemosensor-rich pedipalps, which they use to
touch the females body and silk before initiating cohabitation (Taylor & Jackson 1999).
Visual cues may also allow males to assess females. In
controlled eld experiments, small and large adult male
P. clarus preferred large penultimate females to small females. Mating with females with long tibias may have
tness benets: in a 2003 eld study, Hoeer (unpublished
data) found a strong correlation between female tibia
length and the number of spiderlings that emerged from
egg sacs (F1,35 156.23, N 37, R2 0.817, P < 0.0001).
Female size and time to sexual maturity were signicantly
inversely correlated, and their potential independent roles
cannot be teased apart at present. Males of all sizes (small,
medium, large) also preferred the silk of penultimate females that moulted sooner over those that moulted later.
Although female size and date of sexual maturity were,
again, signicantly inversely correlated, males did not discriminate between females on the basis of tibia length at
the P 0.05 level, but statistical power was low (53.6%).
It is reasonable to assume that the smaller size disparity
between females (compared to the 2003 study) whose
silk was used in the 2004 experiment may have made
discrimination more difcult. However, the results were
marginally nonsignicant, and tibia length and date of
maturity appeared to provide the same information, given
their strong correlation (r 0.885). Note, however, that
all simultaneous choice tests were conducted in the absence of maleemale competition, which may bias choice
in nature. An interesting next step would be to examine
how ghting experience may affect male choice.
Adult male arthropods most commonly show preferences for females that are young or virgin and/or large,
heavy, or fat. Preferences such as these often maximize
the number of progeny sired by the fertilizing male:
young or virgin females may be preferred because of rstmale sperm priority patterns or decreased sperm competition in general, large invertebrate females are usually
more fecund (Clutton-Brock 1988), and heavy or fat
females are often gravid (Bonduriansky & Brooks 1998).
However, other male preferences have also been documented: male orb-web spiders, Argiope keyserlingi, prefer
females with narrow abdomens to those with broad
abdomens (Herberstein et al. 2002). Herberstein et al.
(2002) argue that there are two reasons for this preference: (1) males avoid sexual cannibalism, a behaviour
characterized by inseminated females only, and (2) males
have higher fertilization success. Phidippus clarus males
may be choosing females for similar reasons: to minimize
risks of cannibalism by pairing with penultimate females
(on 18 occasions, I observed adult females cannibalizing
males) and to maximize fertilization success by pairing
with females that are, in contrast to A. keyserlingi, large.
Males may also prefer to pair with females closer to
adulthood, because this may increase opportunities for
polygynous mating by decreasing the time invested in
a single female. As previously described, entelegyne spiders, such as P. clarus (and A. keyserlingi), have female
conduit spermathecae. This morphology often favours

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10

ANIMAL BEHAVIOUR, --, -

the sperm of rst mates (Eberhard et al. 1993) and may

also select for male preferences for virgin females. Female
size and mating status would thus appear to strongly
shape P. clarus male mating preferences.

Conclusions
Male choice greatly contributes to the mating system of
P. clarus, a system that has the same characteristics as
those of many other species. Yet, apart from a few species
showing complete sex-role reversal, male mate choice
remains understudied. This may be due, in part, to the
widespread belief that female choice is more common
than male choice, a perception that is probably the result
of the skewed approach to the study of mate choice:
female choice has received greater research effort. Theory
suggests that selection for male choice increases with variation in female reproductive quality and costs associated
with mating, two attributes common to many, if not all,
mating systems. Therefore, male choice may be more
prevalent and important than once believed. Thus, there
remains an urgent need to consider the role of male choice
in studies of mating systems and sexual selection, and
species with size-assortative pairing or mating may be
important for such research.
Acknowledgments
I thank Christa Skow and Kate McHugh for eld assistance. Christa Skow, Jeremy Houser and Liv Baker provided helpful advice during the development of this
project. Ben Normark, Jeff Podos, Ron Prokopy and Paul
Sievert offered helpful suggestions on all aspects of this
project and manuscript. Ann Rypstra, Jen Riem, Jason
Schmidt, George Uetz, Shawn Wilder, Kerri Wrinn and
two anonymous referees provided helpful comments on
the manuscript. Finally, I am deeply indebted to Elizabeth
Jakob for the unwavering support and the autonomy to
develop and pursue this research. This study was supported by an American Arachnological Society student
research grant to C.D.H., and by an National Science
Foundation (NSF) Short-term Grant for Exploratory Research (SGER) grant, a University of Massachusetts Faculty
Research Grant and a grant from the Cooperative State
Research Extension, Education Service, U. S. Department
of Agriculture, Massachusetts Agricultural Experimental
Station (No. 00829) to E.M.J. Some of the work included
in this manuscript was conducted with the support of NSF
grants DBI 0216776 and DBI 0216947.
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Please cite this article in press as: Chad D. Hoeer, Male mate choice and size-assortative pairing in a jumping spider, Phidippus clarus, Anim. Behav. (2007),
doi:10.1016/j.anbehav.2006.10.017