Male pipefish prefer dominant over attractive females
Identifieur interne : 004182 ( Istex/Corpus ); précédent : 004181; suivant : 004183Male pipefish prefer dominant over attractive females
Auteurs : Anders Berglund ; Gunilla RosenqvistSource :
- Behavioral Ecology [ 1045-2249 ] ; 2001-07.
English descriptors
- KwdEn :
- Anim behav berglund, Aquarium, Attraction display, Attraction displays, Attractive displays, Behav, Behav ecol sociobiol berglund, Behavioral ecology, Berglund, Bernet, Brood pouch, Brown females, Brown males, Choice index, Competition duration, Competitive ability, Competitive displays, Competitive interactions, Cowbird song, Dancing duration, Direct observations, Dominant females, Dominant males, Ecol, Female choice, Female ornaments, Female pair, Green females, High quality females, Indirect mate choice, Intense competition, Linear regression line, Male choice, Male competition, Male mate choice, Males mating, Maria sandvik, Mate choice, Mate choice experiment, Mating competition, Opaque divider, Ornament, Ornament display, Ornament display duration, Other females, Risky behaviors, Rosenqvist, Same result, Same signal, Sexual selection, Shorter time, Sociobiol, Syngnathus typhle, Temporary ornament, Total time, Typhle, Wilcoxon test, mate choice, mate competition, ornament, pipefish, sexual selection, signal honesty.
- Teeft :
- Anim behav berglund, Aquarium, Attraction display, Attraction displays, Attractive displays, Behav, Behav ecol sociobiol berglund, Behavioral ecology, Berglund, Bernet, Brood pouch, Brown females, Brown males, Choice index, Competition duration, Competitive ability, Competitive displays, Competitive interactions, Cowbird song, Dancing duration, Direct observations, Dominant females, Dominant males, Ecol, Female choice, Female ornaments, Female pair, Green females, High quality females, Indirect mate choice, Intense competition, Linear regression line, Male choice, Male competition, Male mate choice, Males mating, Maria sandvik, Mate choice, Mate choice experiment, Mating competition, Opaque divider, Ornament, Ornament display, Ornament display duration, Other females, Risky behaviors, Rosenqvist, Same result, Same signal, Sexual selection, Shorter time, Sociobiol, Syngnathus typhle, Temporary ornament, Total time, Typhle, Wilcoxon test.
Abstract
Animals may obtain information guiding their choice between potential partners from observing competitive interactions and displays between them, or from displays directed at the choosing individual. In the sex-role reversed pipefish Syngnathus typhle females display a temporary ornament (a color pattern) to other females as well as to males. We have previously shown that display of female ornaments per se is attractive to males. Here we show that information from competitive displays can override such direct attraction displays as signals in the partner choice process. In a mate choice experiment, an enclosed male could choose between two females. On the first experimental day, females could interact freely, while on the second day they were isolated from each other. When female-female competition was allowed, the ornament display was directed more to the other female than to the male: Time competing, rather than time courting the male, correlated with ornament display duration. However, ornament display under competition and ornament display in the absence of competition did not correlate significantly. In fact, females competing more intensively on day one displayed the ornament less on day two. Furthermore, the ornament display during the first, but not the second, day predicted male mate choice on the second day. Thus, males remembered previous information from competitive displays and used it rather than immediate information from displays in the absence of female-female competition. We suggest that competitive displays more reliably signal female quality as compared to noncompetitive ones, and that males benefit from mating with dominant females.
Url:
DOI: 10.1093/beheco/12.4.402
Links to Exploration step
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<front><div type="abstract" xml:lang="en">Animals may obtain information guiding their choice between potential partners from observing competitive interactions and displays between them, or from displays directed at the choosing individual. In the sex-role reversed pipefish Syngnathus typhle females display a temporary ornament (a color pattern) to other females as well as to males. We have previously shown that display of female ornaments per se is attractive to males. Here we show that information from competitive displays can override such direct attraction displays as signals in the partner choice process. In a mate choice experiment, an enclosed male could choose between two females. On the first experimental day, females could interact freely, while on the second day they were isolated from each other. When female-female competition was allowed, the ornament display was directed more to the other female than to the male: Time competing, rather than time courting the male, correlated with ornament display duration. However, ornament display under competition and ornament display in the absence of competition did not correlate significantly. In fact, females competing more intensively on day one displayed the ornament less on day two. Furthermore, the ornament display during the first, but not the second, day predicted male mate choice on the second day. Thus, males remembered previous information from competitive displays and used it rather than immediate information from displays in the absence of female-female competition. We suggest that competitive displays more reliably signal female quality as compared to noncompetitive ones, and that males benefit from mating with dominant females.</div>
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<monogr><title level="j">Behavioral Ecology</title>
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<abstract xml:lang="en"><p>Animals may obtain information guiding their choice between potential partners from observing competitive interactions and displays between them, or from displays directed at the choosing individual. In the sex-role reversed pipefish Syngnathus typhle females display a temporary ornament (a color pattern) to other females as well as to males. We have previously shown that display of female ornaments per se is attractive to males. Here we show that information from competitive displays can override such direct attraction displays as signals in the partner choice process. In a mate choice experiment, an enclosed male could choose between two females. On the first experimental day, females could interact freely, while on the second day they were isolated from each other. When female-female competition was allowed, the ornament display was directed more to the other female than to the male: Time competing, rather than time courting the male, correlated with ornament display duration. However, ornament display under competition and ornament display in the absence of competition did not correlate significantly. In fact, females competing more intensively on day one displayed the ornament less on day two. Furthermore, the ornament display during the first, but not the second, day predicted male mate choice on the second day. Thus, males remembered previous information from competitive displays and used it rather than immediate information from displays in the absence of female-female competition. We suggest that competitive displays more reliably signal female quality as compared to noncompetitive ones, and that males benefit from mating with dominant females.</p>
</abstract>
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<item><term>mate choice</term>
</item>
<item><term>mate competition</term>
</item>
<item><term>ornament</term>
</item>
<item><term>pipefish</term>
</item>
<item><term>sexual selection</term>
</item>
<item><term>signal honesty</term>
</item>
<item><term>Syngnathus typhle</term>
</item>
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<istex:document><article xml:lang="en" article-type="research-article">
<front>
<journal-meta>
<journal-id journal-id-type="hwp">beheco</journal-id>
<journal-id journal-id-type="nlm-ta">Behav Ecol</journal-id>
<journal-id journal-id-type="publisher-id">beheco</journal-id>
<journal-title>Behavioral Ecology</journal-title>
<abbrev-journal-title abbrev-type="publisher">Behavioral Ecology</abbrev-journal-title>
<issn pub-type="ppub">1045-2249</issn>
<issn pub-type="epub">1465-7279</issn>
<publisher>
<publisher-name>Oxford University Press</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.1093/beheco/12.4.402</article-id>
<article-id pub-id-type="other">0120402</article-id>
<article-id pub-id-type="pii">1465-7279</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Article</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Male pipefish prefer dominant over attractive females</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Berglund</surname>
<given-names>Anders</given-names>
</name>
<xref rid="AFF1">
<sup>a</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Rosenqvist</surname>
<given-names>Gunilla</given-names>
</name>
<xref rid="AFF2">
<sup>b</sup>
</xref>
</contrib>
<aff id="AFF1"><label><sup>a</sup>
</label>
Department of Animal Ecology, Uppsala University,
Norbyvägen 18d, S-752 36 Uppsala, Sweden</aff>
<aff id="AFF2"><label><sup>b</sup>
</label>
Department of Zoology, Norwegian University of Science and Technology, N-7491
Trondheim, Norway</aff>
</contrib-group>
<author-notes>
<corresp>
Address correspondence to A. Berglund. E-mail:
<ext-link xlink:href="anders.berglund@ebc.uu.se" ext-link-type="email">anders.berglund@ebc.uu.se</ext-link>
.</corresp>
</author-notes>
<pub-date pub-type="ppub">
<month>7</month>
<year>2001</year>
</pub-date>
<volume>12</volume>
<issue>4</issue>
<fpage>402</fpage>
<lpage>406</lpage>
<history>
<date date-type="accepted">
<day>7</day>
<month>9</month>
<year>2000</year>
</date>
<date date-type="received">
<day>14</day>
<month>4</month>
<year>2000</year>
</date>
<date date-type="rev-recd">
<day>1</day>
<month>9</month>
<year>2000</year>
</date>
</history>
<permissions>
<copyright-statement>International Society of Behavioral
Ecology</copyright-statement>
<copyright-year>2001</copyright-year>
</permissions>
<abstract xml:lang="en">
<p>Animals may obtain information guiding their choice between potential
partners from observing competitive interactions and displays between them, or
from displays directed at the choosing individual. In the sex-role reversed
pipefish <italic>Syngnathus typhle</italic>
females display a temporary ornament (a
color pattern) to other females as well as to males. We have previously shown
that display of female ornaments per se is attractive to males. Here we show
that information from competitive displays can override such direct attraction
displays as signals in the partner choice process. In a mate choice
experiment, an enclosed male could choose between two females. On the first
experimental day, females could interact freely, while on the second day they
were isolated from each other. When female-female competition was allowed, the
ornament display was directed more to the other female than to the male: Time
competing, rather than time courting the male, correlated with ornament
display duration. However, ornament display under competition and ornament
display in the absence of competition did not correlate significantly. In
fact, females competing more intensively on day one displayed the ornament
less on day two. Furthermore, the ornament display during the first, but not
the second, day predicted male mate choice on the second day. Thus, males
remembered previous information from competitive displays and used it rather
than immediate information from displays in the absence of female-female
competition. We suggest that competitive displays more reliably signal female
quality as compared to noncompetitive ones, and that males benefit from mating
with dominant females.</p>
</abstract>
<kwd-group kwd-group-type="KWD" xml:lang="en">
<kwd>mate choice</kwd>
<kwd>mate competition</kwd>
<kwd>ornament</kwd>
<kwd>pipefish</kwd>
<kwd>sexual selection</kwd>
<kwd>signal honesty</kwd>
<kwd>
<italic>Syngnathus typhle</italic>
</kwd>
</kwd-group>
<custom-meta-wrap>
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<meta-value>402</meta-value>
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<meta-value>Article</meta-value>
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</article-meta>
</front>
<body>
<sec>
<title></title>
<p>A sexually selected signal usually serves both as an armament (a weapon or
a status badge) and as an ornament (a mate attractant;
<xref rid="REF6">Berglund et al., 1996</xref>
).
However, this may not be invariably true for all sexually selected signals
(<xref rid="REF20">Forsgren, 1997</xref>
;
<xref rid="REF26">Moore and Moore, 1999</xref>
;
<xref rid="REF27">Qvarnström
and Forsgren, 1998</xref>
), because messages like “high fighting
ability” and “tender parental care” may be difficult to
communicate convincingly through the same signal (Zahavi A, personal
communication). Nevertheless, in a majority of cases the same signal has been
found to function in both contexts
(<xref rid="REF6">Berglund et al., 1996</xref>
; Mateos
and Carranza, <xref rid="REF24">1997</xref>
,
<xref rid="REF25">1999</xref>
). Furthermore, signals of
competitive ability can be expected to be attractive in themselves. Given that
such information is used in mate choice, will a direct observation of
competitive interactions/displays be more important to a choosing individual
than observations of some ornamental display meant to attract the chooser?
Since not necessarily all individuals signal their quality honestly, we expect
direct observations of competitive displays to yield more reliable information
about the condition of the potential partner than do purely attractive
displays. This is because condition is immediately put to trial in a contest
situation (through potentially costly fights), whereas the punishment for
discovery of a faked signal of attraction is much less severe (a lost mating
opportunity; see e.g., <xref rid="REF6">Berglund et al.,
1996</xref>
; <xref rid="REF32">West and King,
1980</xref>
; <xref rid="REF33">West et al.,
1981</xref>
). Because a low frequency of dishonest signals of attraction
may be evolutionarily stable (<xref rid="REF23">Kokko,
1997</xref>
; <xref rid="REF29">Viljugrein,
1997</xref>
), a preference for sources of information that have passed a
test may be common in animals. This is further corroborated by the frequent
observation that competitive encounters in many species seem to be staged or
incited by the choosing sex (Bisazza et al.,
<xref rid="REF14">1989a</xref>
,<xref rid="REF15">b</xref>
;
<xref rid="REF16">Borgia, 1981</xref>
;
<xref rid="REF17">Cox and LeBoeuf, 1977</xref>
;
<xref rid="REF18">Farr and Travis, 1986</xref>
;
<xref rid="REF28">Thornhill, 1988</xref>
).</p>
<p>In this article we show that in a sex-role reversed species, the pipefish
<italic>Syngnathus typhle</italic>
L., a male's mate choice is influenced more by a
female's competitive display than by her attraction display. We also show that
males are able to remember and utilize earlier information about competitive
ability in preference to more recent information from attraction display in
their choice of mates, indicating the overriding value of information from
competitive interactions for mate choice in this species.</p>
<sec>
<title>The pipefish</title>
<p>A lengthy and ritualized mutual dance precedes copulation in all pipefish.
In <italic>S. typhle</italic>
, females transfer eggs to a brood pouch under the male's
tail, where he then fertilizes them. Subsequently, embryos are nourished and
oxygenated in the pouch until birth several weeks later (Berglund et al.,
<xref rid="REF11">1986a</xref>
,<xref rid="REF12">b</xref>
).
Males consequently enjoy a paternity confidence of exactly 100%, as confirmed
by a recent microsatellite analysis of maternity and paternity in this species
(<xref rid="REF22">Jones et al., 1999</xref>
). From
many laboratory experiments and field observations we already know that <italic>S.
typhle</italic>
is sex-role reversed: males are typically more choosy and females
compete more intensely than males for access to mates (Berglund,
<xref rid="REF4">1999</xref>
,
<xref rid="REF5">2000</xref>
; Berglund and Rosenqvist,
<xref rid="REF7">1990</xref>
,
<xref rid="REF8">1993</xref>
; Berglund et al.,
<xref rid="REF11">1986a</xref>
,<xref rid="REF12">b</xref>
;
Vincent et al., <xref rid="REF30">1994</xref>
,
<xref rid="REF31">1995</xref>
). Fecundity increases
with body size in both males and females, and given a choice both prefer to
mate with larger partners (<xref rid="REF11">Berglund et
al., 1986a</xref>
).</p>
</sec>
<sec>
<title>The ornament</title>
<p>Female <italic>S. typhle</italic>
display a temporary ornament when interacting
either with other females or with males. The ornament is a striped contrasting
dark color pattern resembling a row of capital B's along the female's side
(<xref rid="REF19">Fiedler, 1954</xref>
). Ornament
appearance is rapid, occurring within a minute, and the display may also
vanish just as fast. Both males and females are normally extremely cryptic
because their color, shape, and movements look like eelgrass (<italic>Zostera
marina</italic>
L.; Vincent et al.,
<xref rid="REF30">1994</xref>
,
<xref rid="REF31">1995</xref>
). Therefore, the ornament
display decreases crypsis in females, which consequently are reluctant to
display under perceived predation threat
(<xref rid="REF13">Bernet et al., 1998</xref>
).</p>
<p>We have previously shown that females who spontaneously display the
ornament for a longer time enjoy a higher mating success than females giving
briefer display (<xref rid="REF13">Bernet et al.,
1998</xref>
). Furthermore, when kept with a male, females are more likely
to display the ornament under female-female competition than in the absence of
competition (<xref rid="REF13">Bernet et al.,
1998</xref>
). Moreover, females displaying more are also more fecund than
similar-sized females displaying less
(<xref rid="REF10">Berglund et al., 1997</xref>
). In
addition, males have been found to prefer ornamented females over
nonornamented ones: an experiment where females were manipulated (painted) to
differ in ornamentation but not in behavior (females were sedated and
mechanically moved up and down by a motor) confirmed that males pay attention
to the ornament even when female behavior was kept constant
(<xref rid="REF9">Berglund and Rosenqvist,
2001</xref>
).</p>
</sec>
<sec>
<title>Pipefish mating competition</title>
<p>Overt aggression is less apparent in these slow-moving, joint-jawed, and
toothless fishes. Female <italic>S. typhle</italic>
compete for males indirectly
through dominance hierarchies, and large females may interfere with and
substantially decrease reproduction in small ones
(<xref rid="REF1">Berglund, 1991</xref>
). Competition
may, however, also be overt. In nature male <italic>S. typhle</italic>
actively search
for, choose among, and reject some females, while females vigorously display,
often in temporary groups in a lek-like fashion. Males typically swim
cryptically within the eelgrass in search of females. Females display by
swimming up and down and in and out of the eelgrass, with their ornaments in
full bloom. Once such a displaying group of females is encountered by a male,
he may start dancing and mating with one of them, usually a large individual
(Vincent et al., <xref rid="REF30">1994</xref>
,
<xref rid="REF31">1995</xref>
). Females actively
compete among themselves for matings during such group displays, and may try
to herd other females away from the male
(<xref rid="REF31">Vincent et al., 1995</xref>
). When
this happens, mating is interrupted, and indeed attempts to copulate can be
thwarted for a long time by interference from other females (Berglund A and
Rosenqvist G, own field and aquaria observations, unpublished data).</p>
<p>Displaying, dancing, and mating are risky behaviors in the otherwise
reticent life of pipefish (<xref rid="REF2">Berglund,
1993</xref>
; <xref rid="REF13">Bernet et al.,
1998</xref>
; <xref rid="REF21">Fuller and Berglund,
1996</xref>
). Therefore, it may well pay males to select dominant
females, which are able to discourage other females from courtship
interference, as this will reduce the time spent on potentially risky
behaviors. Moreover, males do not have to worry about the potential conflict
between female signals of dominance and signals of caring ability, as females
contribute no care, only eggs. Moreover, we have never observed female
aggression towards males. Thus, our prediction is that males should treasure
dominance and competitive ability highly in females, and that direct
observations by males of such abilities in females may be more important than
attraction displays in the male's mate choice process. Indeed, any female can
display her ornament, but females having done so under actual female-female
competition have demonstrated their power and competitive ability in a
potentially costly situation. Therefore, males mating with such evidently
dominant females can expect a smooth courtship and copulation process, gaining
direct benefits for themselves in terms of reduced risk, as well as possible
direct and/or genetic benefits for their offspring from mating with high
quality females.</p>
</sec>
</sec>
<sec>
<title>METHODS</title>
<p>We caught the pipefish in shallow eelgrass meadows in the Gullmar Fjord on
the Swedish west coast during the latter half of May 1997, before the breeding
season commenced. We kept sexually mature males and females in separate
aquaria to assure that individuals would be reproductively active. Stock
aquaria contained plastic plants and continuously renewed sea water (natural
temperature, salinity, and light regime). Fishes were fed brine shrimps
(<italic>Artemia</italic>
), small, wild-caught crustaceans, and frozen mysids <italic>ad
lib</italic>
.</p>
<p>To explore the importance of competitive versus purely attractive displays
we ran a mate choice experiment 28 May-20 June. Each of 24 trials was run for
2 days, with one male choosing between two females. The females came from
different stock aquaria. The male was separated from the females by a
transparent plastic divider. During the first day females could interact
freely with one another, but during the second day they were separated from
one another by an opaque divider (<xref rid="FIG1">Figure
1</xref>
). Separated females could not see or smell each other, but the
male could see and smell both females both days as water flowed from the
female compartment(s) into the male's and then out of the aquarium. Thus,
males could observe and dance with females on the first day, and females could
compete, dance and display their ornament on that day. On day one, we placed
the females in the rear compartment and the male in the front compartment
(<xref rid="FIG1">Figure 1</xref>
), and videotaped them
for 10 h. On this day, we measured the time each fish spent dancing, how long
females displayed their ornaments and whether females competed or not. A male
and a female were defined as dancing when they simultaneously bobbed up and
down in close proximity while facing one another. We considered females to be
competing when they pursued each other round in the aquarium for more than a
minute.</p>
<p>
<fig id="FIG1" position="float">
<label>
<bold>Figure 1</bold>
</label>
<caption>
<p>The experimental design on day two. A male with his brood pouch is seen in
the foreground, and two enclosed females in the back. On day one the opaque
partitioning between the females was absent.</p>
</caption>
<graphic xlink:href="bhec-12-04-06-f01"></graphic>
</fig>
</p>
<p>On the second day, we measured male choice of a female in the absence of
female-female competition by placing the opaque divider between the females
and filming for another 10 h. Females were allocated to either chamber
randomly. After that males and females were released back into the wild. No
mortality occurred during trials. We measured the duration of female resting,
swimming, dancing, and ornament display, and the duration of male resting,
swimming, and dancing before either female. The female with which the male
spent the longest time was considered as the chosen one (a longer time with
the female has earlier been shown to accurately predict mate choice;
<xref rid="REF3">Berglund, 1994</xref>
).</p>
<p>We filmed two trials simultaneously in 125 1 aquaria (45 × 50 ×
60 [height] cm). The aquaria were continuously provided with surface sea
water. Temperature and light conditions followed natural conditions. We
planted fresh eelgrass in beach sand in the aquaria for shelter. Fish were not
fed during trials. We selected females of similar standard length (within 10
mm of each other) and of the same contrast score (the vaguely striped basic
pattern from which the ornament is formed, as estimated by eye on a five-grade
scale; see <xref rid="REF13">Bernet et al., 1998</xref>
for details), but individually recognizable by color (estimated by eye). New
males and females were used in each trial. Males were smaller than females
(males 177 ± 21 mm, <italic>n</italic>
= 24, females 226 ± 27 mm,
<italic>n</italic>
= 48, <italic>t</italic>
<sub>70</sub>
= 8.00, <italic>p</italic>
<.001), as in
the natural situation (in our 1997 field samples males were on average 161
±
28 mm, females 191 ± 38 mm).</p>
<p>Videotaping was done with Hitachi Hi-8 WM-H80E video cameras connected to
Panasonic AG-6730 time-lapse super-VHS video recorders equipped with AG-IA670
time code generators/computer interfaces, using the 24 h time-lapse mode
(which employs 1/8 normal speed). Intense light (a 100W spot and a 20W
luminescent lamp placed 1/2 m above each aquarium) was used during filming, in
addition to natural light from windows. Videotapes were analyzed using the
Observer 3.0 VTA software, and persons scoring videos were unaware of the
objective of the study. The Observer software kept track of duration and
latency of behaviors with a precision of less than a second, which is an
accuracy far more than enough for these slow-moving fishes.</p>
<p>Statistical probabilities reported from these experiments are two-tailed.
Nonparametric tests were used whenever the assumptions of parametric tests
were not met. Experiments were performed under a license from the Swedish
ethical board.</p>
</sec>
<sec>
<title>RESULTS</title>
<p>We classified the two females in a pair as “the one displaying the
ornament for a longer time” and “the one displaying for a shorter
time,” respectively, during day one. We found no significant differences
between these two female categories in standard body length (long displaying
females were on average 228 ± SD 29 mm, short displaying 221 ±
26 mm, paired <italic>t</italic>
test, <italic>t</italic>
<sub>19</sub>
= 1.42, <italic>p</italic>
=.2)
or contrast (mean 3.10 ± 2.00 for both groups of females, Wilcoxon
matched-pairs test, <italic>T</italic>
= 6, <italic>n</italic>
= 20 pairs, <italic>p</italic>
= 1). In
four trials both females either constantly displayed the ornament or did not
display it at all; these females were discounted. When females instead were
categorized as being “chosen” or “rejected,” that is,
which female the male spent most or least time with on day 2, again female
lengths did not differ (chosen females 229 ± 27 mm, rejected 224
±
26 mm, <italic>t</italic>
<sub>23</sub>
= 1.25, <italic>p</italic>
=.2), nor did
contrast (3.08 ± 0.4 in both groups, <italic>T</italic>
= 85, <italic>n</italic>
= 24
pairs, <italic>p</italic>
= 1). Thus, our experiment included closely matching pairs
of females.</p>
<p>Female color did not significantly influence male choice (females broadly
categorized as brown or green, 17 brown and seven green females were chosen,
binomial <italic>p</italic>
=.152) or which female displayed the ornament for the
longest duration on day one (14 brown versus six green, binomial <italic>p</italic>
=.116). Pairing was not color-assortative: green males chose two green and six
brown females, and brown males chose four green and 10 brown females (Fisher's
Exact test <italic>p</italic>
= 1). Similarly, females did not display their ornament
in a color-assortative fashion: green females displayed more to two green and
five brown males, and brown females more to five green and six brown males
(Fisher's Exact test <italic>p</italic>
=.6).</p>
<p>Females displaying the ornament for a longer time on day one were preferred
by males on the second day, compared to females displaying for a shorter time
(<xref rid="FIG2">Figure 2</xref>
). Preference is here
estimated as the total time a male spent in front of a female, but if instead
the dancing duration with either female on day two is used as a choice index
the same result emerges (males danced with more ornamented females for 15.9
±
15.4 min and with less ornamented for 6.2 ± 9.8 min;
<italic>T</italic>
= 26, <italic>n</italic>
= 20 pairs, <italic>p</italic>
<.01). No choice for any
particular female was evident on day one (<italic>p</italic>
>.3 for dancing
duration), but on that day dances were both infrequent and of short durations.
Females displaying their ornaments for longer on day two were, however, not
preferred by males as compared to females displaying for shorter on day two
(291.4 ± 144.0 versus 306.0 ± 143.4 min total time before the
respective females; <italic>T</italic>
= 104, <italic>n</italic>
= 20 pairs, <italic>p</italic>
= 1).
If dancing duration day two is used instead of total time as a choice index,
the same result emerges (<italic>p</italic>
=.5).</p>
<p>
<fig id="FIG2" position="float">
<label>
<bold>Figure 2</bold>
</label>
<caption>
<p>The female displaying the ornament for longer on day one (with
female-female competition) was chosen by the male on day two (with females
separated), as compared to the female displaying for shorter (Wilcoxon
matched-pairs test, <italic>T</italic>
= 32, <italic>p</italic>
<.01, <italic>n</italic>
= 24).
Mate choice is estimated in terms of the total time spent by a male in front
of either female on day two. Black squares are medians, boxes quartiles and
bars represent minimum and maximum values. Inserts show mate choice aquaria
from above on day one and two.</p>
</caption>
<graphic xlink:href="bhec-12-04-06-f02"></graphic>
</fig>
</p>
<p>Similarly, females chosen on day two had displayed their ornaments for
longer on day one (chosen females for 233.2 ± 176.7 min, rejected for
130.3 ± 151.5 min; <italic>T</italic>
= 40, <italic>n</italic>
= 24 pairs, <italic>p</italic>
=.015) but not on day two (123.7 ± 171.8 and 99.7 ± 128.4 min,
respectively; <italic>T</italic>
= 85, <italic>n</italic>
= 24 pairs, <italic>p</italic>
=.5).</p>
<p>On day one, females could either display the ornament to each other, in
which case the duration of ornament display would correlate with the duration
of competitive interactions, or females could display the ornament to males,
in which case ornament display duration would correlate with the duration of
intersexual interactions (i.e., dancing duration). It turned out that ornament
display correlated significantly with competition duration, but not with
dancing duration (<xref rid="FIG3">Figure 3</xref>
),
evidently because most females were busy competing and did not dance at all.
As the competition duration by necessity was the same for both females in a
pair, competition duration was regressed on the average ornamentation duration
for each female pair. Similarly, the average dancing duration of each female
pair was regressed on competition duration to avoid pseudoreplication. This
result also emerges from a multiple correlation (regression summary
<italic>F</italic>
<sub>2,21</sub>
= 9.61, <italic>p</italic>
=.001) for the effects of
competition and dancing duration on ornament display duration:
β
<sub>competition</sub>
= 0.544 (<italic>p</italic>
=.01) and
β
<sub>dance</sub>
= 0.215 (<italic>p</italic>
=.28). Clearly competition more
accurately predicted ornamentation than did dancing, and, moreover, this
predominantly competition-induced ornamentation significantly predicted male
mate choice the next day.</p>
<p>
<fig id="FIG3" position="float">
<label>
<bold>Figure 3</bold>
</label>
<caption>
<p>The duration of female ornament display on day one (with female-female
competition) correlated with (a) the duration of female-female competition
that day (Spearman <italic>R<sub>s</sub>
</italic>
=.60, <italic>n</italic>
= 24, <italic>p</italic>
<.002, linear regression line showed in graph), but not with (b) the
dancing duration with the male (Spearman <italic>R<sub>s</sub>
</italic>
= -.05,
<italic>n</italic>
= 24, <italic>p</italic>
=.8). The regression in (a) is significant even if
the extreme data point in the upper right corner is removed (Spearman
<italic>R<sub>s</sub>
</italic>
=.55, <italic>n</italic>
= 23, <italic>p</italic>
<.01).</p>
</caption>
<graphic xlink:href="bhec-12-04-06-f03"></graphic>
</fig>
</p>
<p>The more intensely females engaged in competition on day one, the less
likely they were to display the ornament on day two
(<xref rid="FIG4">Figure 4</xref>
). Pseudoreplication
is compensated for by using average ornament display duration within each pair
on day two. Interestingly, this correlation is driven by the chosen females:
when analyzed separately, competition duration on day one for chosen females
correlated significantly with ornament display day two (<italic>R<sub>s</sub>
</italic>
= -.432, <italic>n</italic>
= 24, <italic>p</italic>
=.035), but not so for rejected females
(<italic>R<sub>s</sub>
</italic>
= -.302, <italic>n</italic>
= 24, <italic>p</italic>
=.15).</p>
<p>
<fig id="FIG4" position="float">
<label>
<bold>Figure 4</bold>
</label>
<caption>
<p>The more time females spent competing on day one, the less they displayed
their ornament on the subsequent day when female-female interactions were
prevented (Spearman <italic>R<sub>s</sub>
</italic>
= -.42, <italic>n</italic>
= 24, <italic>p</italic>
<.05, linear regression line shown in graph).</p>
</caption>
<graphic xlink:href="bhec-12-04-06-f04"></graphic>
</fig>
</p>
</sec>
<sec>
<title>DISCUSSION</title>
<p>When female-female competition was allowed, females displayed the ornament
to one another more persistently than they did to the male. The female
displaying most intensely is likely to be the dominant one within a pair.
Clearly, a more intense ornament display (or other information from watching
competitive interactions) made this female attractive to the male, as males
used this information on the following day when females no longer were allowed
to compete: The male spent most time before the female that had displayed for
longer in the inter-female competitive situation. Indeed, this past
information of ornament or competitive displays completely overrode more
recently acquired information, and significantly influenced male mate choice.
Furthermore, ornament display under competition did not correlate
significantly with ornament display in the absence of competition.
Consequently, on the day of the male's choice, ornament display contained
little or no information regarding female dominance. Thus, male pipefish seem
to remember and make use of information regarding partner dominance,
presumably according to the reliability of that information. An alternative
explanation is that males are conservative and stick to a decision once made.
However, `conservatism' as a reason to why males should ignore recent display
information and instead prefer older information seems unlikely. Moreover, in
nature these fish do not form pair bonds or exhibit territoriality
(<xref rid="REF30">Vincent et al., 1994</xref>
),
further making the “conservatism” explanation unlikely. Note that
the effect of mating competition on the choosing sex is not merely one of
constraining the options for choice (indirect mate choice, see
<xref rid="REF34">Wiley and Poston, 1996</xref>
) but
active mate choice.</p>
<p>We ran a mirror-image experiment in 1996, with females separated the first
day and competing the second (<xref rid="REF9">Berglund and
Rosenqvist, 2001</xref>
). Another difference from the experiment reported
here was that males had full access to females on the second day, such that
copulations were possible and occurred (this could not be allowed in the
present experiment, as males would then get pregnant and lose interest in
females the second day). In the mirror-image experiment, with females
separated during the first day, ornament display day one significantly
predicted ornament display on the second day, when female-female interactions
were allowed. Ornament display on either day also significantly predicted male
mate choice. This makes a possible alternative explanation to our results
unlikely, namely the explanation that time itself, rather than the absence of
female-female competition on day two, caused the results. Both males and
females chose and behaved similarly on both days in the mirror-image
experiment, so time effects, such as experience with the experimental set-up,
are unlikely to have confounded the results presented here. Thus, when females
were separated on the first day, their ornament display was the same on the
second day with competition, but when competition preceded separation this was
no longer true. Does intense competition make females less able, or less
willing, to subsequently display their attractiveness? The finding that
females competing more intensely on day one displayed their ornaments to a
lesser extent on day two (<xref rid="FIG4">Figure
4</xref>
) may suggest that intense competition is tiring or otherwise
discourages females from performing the display on the following day. Indeed,
long-term reproductive inhibition has been demonstrated in this species
before: Larger, and presumably dominant, females may decrease reproductive
activity in smaller females (<xref rid="REF1">Berglund,
1991</xref>
). Alternatively, a female having proven her potential during
female-female encounters may not need to display her attractiveness further,
as males will rank such information more highly than pure displays of
attraction. Presently, we cannot tell whether females become dispirited or
arrogant after an episode of intense competition, but the finding that only
chosen, not rejected, females refrained from displaying their attractiveness
on day two suggests arrogance.</p>
<p>Thus, ornament display under female-female competition may be a more
reliable and important signal than intersexual ornament display. Males mating
with dominant females may gain direct benefits for themselves in terms of
reduced risk during courtship, as well as possible direct and/or genetic
benefits for their offspring from mating with high quality females. Therefore,
male pipefish prefer beautifully ornamented females, but, above all,
victorious ones.</p>
</sec>
</body>
<back>
<ack>
<p>The work was funded by the Swedish Natural Research Council (grant to A.B.)
and by the Norwegian Research Council (grant to G.R.). We thank Klubban
Biological Station and Kristineberg Marine Research Station for research
facilities. Thanks to Arnt Narve Bordal, Maria Sandvik, and Hanna
Schiöler-Wasslavik for field assistance and
video analysis, and to Ingrid Ahnesjö, Ian
Fleming, Elisabet Forsgren, Maria Sandvik, and Staffan Ulfstrand for valuable
comments on an earlier draft.</p>
</ack>
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</article>
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<mods version="3.6"><titleInfo lang="en"><title>Male pipefish prefer dominant over attractive females</title>
</titleInfo>
<titleInfo type="alternative" lang="en" contentType="CDATA"><title>Male pipefish prefer dominant over attractive females</title>
</titleInfo>
<name type="personal"><namePart type="given">Anders</namePart>
<namePart type="family">Berglund</namePart>
<affiliation>Department of Animal Ecology, Uppsala University, Norbyvägen 18d, S-752 36 Uppsala, Sweden</affiliation>
<role><roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal"><namePart type="given">Gunilla</namePart>
<namePart type="family">Rosenqvist</namePart>
<affiliation>Department of Zoology, Norwegian University of Science and Technology, N-7491 Trondheim, Norway</affiliation>
<role><roleTerm type="text">author</roleTerm>
</role>
</name>
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<genre type="research-article" displayLabel="research-article" authority="ISTEX" authorityURI="https://content-type.data.istex.fr" valueURI="https://content-type.data.istex.fr/ark:/67375/XTP-1JC4F85T-7">research-article</genre>
<originInfo><publisher>Oxford University Press</publisher>
<dateIssued encoding="w3cdtf">2001-07</dateIssued>
<copyrightDate encoding="w3cdtf">2001</copyrightDate>
</originInfo>
<language><languageTerm type="code" authority="iso639-2b">eng</languageTerm>
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</language>
<abstract lang="en">Animals may obtain information guiding their choice between potential partners from observing competitive interactions and displays between them, or from displays directed at the choosing individual. In the sex-role reversed pipefish Syngnathus typhle females display a temporary ornament (a color pattern) to other females as well as to males. We have previously shown that display of female ornaments per se is attractive to males. Here we show that information from competitive displays can override such direct attraction displays as signals in the partner choice process. In a mate choice experiment, an enclosed male could choose between two females. On the first experimental day, females could interact freely, while on the second day they were isolated from each other. When female-female competition was allowed, the ornament display was directed more to the other female than to the male: Time competing, rather than time courting the male, correlated with ornament display duration. However, ornament display under competition and ornament display in the absence of competition did not correlate significantly. In fact, females competing more intensively on day one displayed the ornament less on day two. Furthermore, the ornament display during the first, but not the second, day predicted male mate choice on the second day. Thus, males remembered previous information from competitive displays and used it rather than immediate information from displays in the absence of female-female competition. We suggest that competitive displays more reliably signal female quality as compared to noncompetitive ones, and that males benefit from mating with dominant females.</abstract>
<note type="author-notes">Address correspondence to A. Berglund. E-mail: anders.berglund@ebc.uu.se .</note>
<subject lang="en"><genre>KWD</genre>
<topic>mate choice</topic>
<topic>mate competition</topic>
<topic>ornament</topic>
<topic>pipefish</topic>
<topic>sexual selection</topic>
<topic>signal honesty</topic>
<topic>Syngnathus typhle</topic>
</subject>
<relatedItem type="host"><titleInfo><title>Behavioral Ecology</title>
</titleInfo>
<titleInfo type="abbreviated"><title>Behavioral Ecology</title>
</titleInfo>
<genre type="journal" authority="ISTEX" authorityURI="https://publication-type.data.istex.fr" valueURI="https://publication-type.data.istex.fr/ark:/67375/JMC-0GLKJH51-B">journal</genre>
<identifier type="ISSN">1045-2249</identifier>
<identifier type="eISSN">1465-7279</identifier>
<identifier type="PublisherID">beheco</identifier>
<identifier type="PublisherID-hwp">beheco</identifier>
<identifier type="PublisherID-nlm-ta">Behav Ecol</identifier>
<part><date>2001</date>
<detail type="volume"><caption>vol.</caption>
<number>12</number>
</detail>
<detail type="issue"><caption>no.</caption>
<number>4</number>
</detail>
<extent unit="pages"><start>402</start>
<end>406</end>
</extent>
</part>
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<identifier type="istex">848CE6AC64DDA7ADE118030ED403544E3F7F7F6C</identifier>
<identifier type="DOI">10.1093/beheco/12.4.402</identifier>
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<identifier type="local">0120402</identifier>
<accessCondition type="use and reproduction" contentType="copyright">International Society of Behavioral Ecology</accessCondition>
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