Marine and Freshwater Behaviour and Physiology

Vol. 43, No. 3, May 2010, 157–167

Relationships between relative size of sexual traits and male

mating success in the Mediterranean killifish Aphanius fasciatus
(Nardo, 1827)
Stefano Malavasi*, Vyron Georgalas, Francesco Cavraro and Patrizia Torricelli

Department of Environmental Sciences, University ‘Ca’ Foscari’ of Venice, Campo della

Celestia, Castello 2737/b, 30122 Venice, Italy
(Received 19 April 2009; final version received 21 March 2010)

Some relationships between the relative size of secondary sexual traits and
male mating success were investigated in the killifish Aphanius fasciatus
(Nardo, 1827). The relative ability of a male to compete with another male
for access to a ripe female was related to the relative size of male sexual
traits by means of a behavioural experiment under controlled laboratory
conditions. Furthermore, the covariation in the expression of sexual traits
was analysed from a sample of field-collected males. Results of the
behavioural experiment showed that male dominance and success were
positively related to the relative height of the dorsal fin, but not to the
pattern of vertical bars along body sides. Within the analysed male
population, males with larger fins tended to have thinner bars, whereas
there was no statistically significant correlation with bar number or with
bar asymmetry. The results were discussed in terms of advantages conferred
by larger dorsal fins in killifish breeding systems, and developmental
constraints defining a multiple set of traits correlated to male quality were
also hypothesised.
Keywords: secondary sexual traits; courtship; male–male competition;
Mediterranean killifish; Aphanius fasciatus

Introduction
Sexual selection forces explain a large part of sexual dimorphism in animal species
determining the development of conspicuous traits (secondary sexual traits, SSTs).
These traits confer significant advantages in terms of reproductive success at the level
of both intrasexual and intersexual selection (Andersson 1994). As males can fertilise
offspring at a higher rate than females, intrasexual selection is thought to be stronger
among males (Andersson 1994).
There are two critical issues for understanding sexual selection: first, the two
components of sexual selection may act in concert, independently or in opposition
(Berglund et al. 1996; McGhee et al. 2007); and secondly, both ‘ornaments and
armaments’ are based upon multiple cues, as reviewed by Ptacek (2000). In fishes,
SSTs include morphological structures, sexual displays, colouration and pheromonal

*Corresponding author. Email: mala@unive.it

ISSN 1023–6244 print/ISSN 1029–0362 online

ß 2010 Taylor & Francis
DOI: 10.1080/10236244.2010.480837
http://www.informaworld.com
158 S. Malavasi et al.

signals that are usually combined in a multivariate set of cues (Ptacek 2000). In the
male swordtail (genus Xiphophorus), for example, the elongated modification of the
caudal fin and the pigment pattern of vertical bars are combined to be attractive
ornaments for advertising to females and deterring signals for the other males
(Morris et al. 1995, 2001, 2005, 2007; Morris and Casey 1998; Benson and Basolo
2006). Other important sexual signals that have been shown to be related to
dominance in male–male contests and/or to enhance female choice are: fin size
(Kokita and Mizota 2002; Suk and Choe 2002), specialised morphological structures
such as tubercules in cyprinids (Kortet et al. 2004; Vainikka et al. 2005), breeding
and aggressive sounds (Myrberg and Lugli 2006) and patterns of colouration
(Kodric-Brown 1990; Milinski and Bakker 1990).
The interaction between male–male competition and female choice recently has
been investigated by McGhee et al. (2007), using a killifish (the fundulid Lucania
goodei) breeding system as a study model. Although the male interference was
crucial, especially at an early phase determining the initial mating pool, the female
decision exerted control in terms of the latency of spawns, number of visits and
released eggs (McGhee et al. 2007). Killifish mating systems therefore appear to be
good models for investigating this issue, as the two components of sexual selection
seem to interact with each other in determining male mating success, as previously
suggested by Fuller (2001). Furthermore, male killifish often show a set of ornaments
that may potentially act as sexual traits, such as colourful and conspicuous modified
fins and a series of vertical bars on the body sides. In this study, we investigated the
relationships between the relative size of male sexual traits, male–male competition
and male mating success in the killifish Aphanius fasciatus.
Aphanius fasciatus (Nardo, 1827) is a highly dimorphic (Gandolfi et al. 1991)
endemic cyprinodont fish occurring in coastal brackish water habitats (Maltagliati
1999). Males have vertical bars and large and modified caudal, anal and dorsal fins,
which show a brilliant colouration (Gandolfi et al. 1991). Females, which are usually
larger than males, show less developed and less colourful caudal, anal and dorsal
fins. The breeding system is based on intense competition among the males for access
to single females that are led by the male to lay eggs on aquatic vegetation (Gandolfi
et al. 1991). Although both sexes show a series of vertical bars along the body, their
colouration and conspicuousness are sexually dimorphic (Gandolfi et al. 1991).
Males have a series of 7–15 large white bars on a dark grey-blue background,
whereas females show narrow black bars on a light-grey background. Although bars
are evident in both sexes throughout the whole year, the bars become better defined
and more brilliant during the breeding season, especially in males (Malavasi,
Georgalas, Cavraro and Torricelli, personal observation). Furthermore, the number
of vertical bars constitutes a typical trait that is subjected to fluctuating asymmetry
(FA), that is, random deviation from perfect bilateral symmetry. The level of
asymmetry in the number of vertical bars has been shown, in some cases, to be a
sexually selected trait in teleost fish (Morris and Casey 1998; Schluter et al. 1998).
For this reason, in the present study we controlled the degree of asymmetry in the
number of male vertical bars as well.
We used the Mediterranean killifish breeding system as a model to address two
main questions:
(1) Is the relative size of sexual traits related to the relative ability of a male to
monopolise the female in dyadic competitive encounters?
 Marine and Freshwater Behaviour and Physiology 159

(2) Are the potential sexual traits related to each other, in terms of relative size,
within the male population?
We addressed the first question by performing an aquarium experiment under
controlled conditions, whereas we analysed samples of field collected males to
address the second question.

Materials and methods

Experimental fish and housing conditions
Fish were captured in April 2007 with a small beach seine from salt marshes located
in the Northern basin of the Venice lagoon (45 26.7900 N; 12 24.9350 E) and then
transported back to the laboratory. A sample of 77 adult males was sacrificed with
an overdose of anaesthetic (2-phenoxyethanol) for SSTs measurements. The rest of
the captured fish were kept at least 7 days prior to experimentation. Fish were
housed in communal holding tanks of 175 L provided with artificial salt water
(Ocean fish, Prodac-Italy). Salinity was kept at 25 psu which was the salinity
measured in situ at the sampling site. Temperature varied between 18 C and 20 C
and the photoperiod regime corresponded to the natural cycle for the latitude and
time of the year. The side-walls of each tank were covered with thick, dark-
blue-coloured paper to prevent fish from excessive stress and/or conditioning.
Fish were fed Chironomus sp. larvae ad libitum twice a day.

Experimental conditions and apparatus

Experiments were performed from the end of April until the beginning of June 2007,
during daylight (9:00–18:00) in 96 L (60 cm  40 cm  40 cm) glass aquaria equipped
with aerators and external filters; the aquaria were filled to a depth of 35 cm with
artificial seawater and the salinity, water temperature and light regimes were kept
identical to those of the housing tanks. Sheets of thick, dark-blue-coloured paper,
similar to those used in the housing tanks, covered the rear and lateral walls of each
experimental aquarium to minimise visual disturbance and provide an adequate
visual contrast with the fish silhouette to facilitate video-tape analysis. The bottom
of each experimental aquarium was covered with 3 cm of sand. Furthermore,
in order to offer an appropriate substratum for egg laying (Gandolfi et al. 1991),
we equipped each aquarium with a floating green mop hanging from a small
quadrant made of expanded polystyrene (5 cm  5 cm) floating on the water surface.
This structure simulated the filamentous algae (Vaucheria spp) present in the salt
marsh habitats where the fish were captured, providing a natural-like site for egg
attachment.

Experimental procedures
A group consisting of two males showing nuptial colouration and a ripe female
was captured from the holding tanks and introduced into the experimental tank.
The males selected for each experimental trial were of similar size in terms of
standard length (SL difference between 0% and 10%) and had a difference of two
vertical bars on the left body side which allowed individual recognition during
160 S. Malavasi et al.

behavioural sequence analysis. The video camera (Canon camcorder MV400) was
positioned behind a U-shape box so that disturbance to the animals was minimised
during the experimental trials. Once fish had resumed normal activity (usually
15–30 min after their introduction to the experimental tank) the video camera was
turned on and the experimental trial was started.
Each experimental trial lasted 90 min. If no interactions occurred during the first
30 min of video-recording, fish were replaced with a new group. The video-camera
tracked the female, which was followed during the whole period of video recording.
This allowed the recording of all the interactions of each or both of the two males
with the ripe female. On the whole, eight experimental trials for a total of 16 males
and eight females were conducted. At the end of each experimental trial, male fish
were sacrificed using an excess of anaesthetic (2-phenoxyethanol) and consecutively
photographed in each body side (see below for a detailed description of the
procedures and the measurements taken).

Sexual traits measurement

Photographs of both the experimental fish (n ¼ 16) and the male population sample
(n ¼ 77) were taken by means of a fixed digital camera (Pentax Optio 43WR).
Entomological pins were used to keep anal, dorsal and caudal fins open. Each fish
was photographed on both sides of the body. All photographs were transferred to a
personal computer and analysed using Image Pro-PlusÕ 3.0. Operator accuracy was
obtained by measuring one photograph taken with the same digital camera
containing 100 objects of known dimensions casually scattered and was found to
be 0.01 cm. For each fish the following measures were obtained (Figure 1): NVB:
number of vertical bars; AFA: anal fin area (cm2); DFA: dorsal fin area (cm2); CFA:
caudal fin area (cm2); CFL: caudal fin length (cm); CFW: caudal fin width (cm);

Figure 1. Drawing representing an A. fasciatus male with all the measured sexual traits,
obtained by means of the Image analyser. CFL: caudal fin length (cm); CFW: caudal fin width
(cm); DFH: dorsal fin height (cm); VBS: vertical bar span of the ‘x’th bar (cm); VBW: vertical
bar width of the ‘x’th bar (cm). : AFA: anal fin area (cm2); : DFA: dorsal fin area
(cm2); : CFA: caudal fin area (cm2).
 Marine and Freshwater Behaviour and Physiology 161

DFH: dorsal fin height (cm) measured vertically on the imaginary line uniting the
origins of the dorsal fin (see Figure 1, where length h1 ¼ length h2); VBSx: vertical
bar span (cm) of the ‘x’th bar (numbered in increasing order from caudal fin to
head); VBWx: vertical bar width of the ‘x’th bar (numbered in increasing order
from caudal fin to head). As bar width and span vary in relation to the point of
measurement (bars tend to be thinner in the dorsal area) (Figure 1), these measures
were taken on the imaginary line passing through the eye centre and the middle
point of the caudal peduncle (see Figure 1, where length a ¼ length b). VBSx and
VBWx were averaged for each part of the body for each fish becoming VBS: average
vertical bar span (cm) and VBW: average vertical bar width, respectively. For
statistical analysis only VBS and VBW were considered. AFA, DFA, CFA, CFL,
CFW and DFH were all measured only on the left side of the body while NVB,
VBSx and VBWx were measured on both sides.

Behavioural variables and data analysis

A preliminary inspection of the videotapes allowed the identification of five
courtship and mating acts (Table 1), some consistent with previous work by Grech
and Schembri (1993). The frequency of courtship acts was calculated as the ratio
between the number of acts exhibited by each male and the total duration of
courtship (number of a given act per minute). Courtship and mating acts were
alternated by intramale aggression, consisting mainly of circling displays and fin
flare behaviours (sensu Fuller 2001). Since during the aggressive sequences the two
males were often not individually recognisable, partly due to the rapidity of their
behaviour, these behaviours were not quantified.
Among the courtship behaviours, spawning can be considered as the best
indicator of male mating success, because each spawning act corresponded to one
event of egg laying by the female (Table 1). In three out of eight encounters,
spawning acts did not occur, whereas the other acts occurred in all the dyadic
contests. When spawning acts occurred, the difference between the two males in

Table 1. Courtship and mating acts identified by the inspection of the videotaped
behavioural sequences.

Courtship act Abbreviation Description

Approach A The male swims following the female or paralleling her

Circling C The male circles around the female in a horizontal plane
at a distance of about 10–15 cm.
Biting B The male prods the female’s belly with his snout
Nudging N The male prods the female on the dorsal part of her head
with his snout, then swimming above her and
maintaining the physical contact.
Spawning S The male assumes an S-shape and wraps its body tightly
to that of the female, both fish quivering the posterior
part of the body, when egg laying could be observed.
This behaviour may be observed both inside the
floating mop or near the glass walls of the aquarium.

Note: See Grech and Schembri (1993) for abbreviations and descriptions.
162 S. Malavasi et al.

terms of spawning frequency ranged from 0.01 to 1.5 n min1. The frequency of
spawning was positively related to the nudging frequency (Spearman’s correlation
r ¼ 0.97, p 5 0.05), which preceded spawning in all those replicates where spawning
occurred. By contrast, spawning was not significantly correlated to any of the other
identified behaviours (Spearman’s correlation, p 4 0.05). We therefore used nudging
as the best predictor of male dominance and male mating success, defining as
‘dominant’ and ‘subordinate’ the males with the higher and the lower frequency of
nudging acts, respectively. This was consistent with that found by McGhee et al.
(2007) in L. goodei, that is, interference between males is reflected in one male
(the dominant) that keeps the other one (the subordinate) from courtship and
spawning with the female.
We calculated a ‘Relative Monopolisation Index’ (RMI) as the difference
between the frequency of the total number of nudging acts performed by the
dominant male and that shown by the subordinate male, standardised on the total
frequency of acts performed by the male pair (i.e.: RMI ¼ (Fd  Fs/Fs þ Fd), where
F ¼ frequency of nudging act, d ¼ dominant male and s ¼ subordinate male).
RMI indicated the degree to which the dominant male was able to access the
female compared to the subordinate male. We also calculated a ‘Relative Trait Index’
(RTI) of each sexual trait, based on the difference between the size measures of a
given trait (Figure 1) of the ‘dominant’ male and that of the ‘subordinate’ male,
adjusted to the SL of the largest male of the pair, i.e.: RTI ¼ (TSd  TSs/SLm) for a
given sexual trait, where TS ¼ size of a given trait, d ¼ dominant male and
s ¼ subordinate male, SLm ¼ standard length of the largest male of the pair. An
Asymmetry Index in the number of vertical bars was also calculated as follows:
VBN(left)  VBN(right)/VBN(left) þ VBN(right), where VBN ¼ number of vertical
bars at left or right side of the body. The Relative Monopolisation Index (RMI) was
related to the Relative Trait Index (RTI), using Spearman’s correlation analysis. The
RMI was also related to the difference in the Asymmetry Index between the two
males. The size-adjusted sexual traits measured from the fins of the field-collected
males, obtained as the residuals of the Pearson’s correlation between SL and trait
size (n ¼ 77), were related to the vertical bar traits (vertical bar width and span),
to the number of vertical bars along the body and to the absolute value of the
Asymmetry Index, by means of a Spearman’s correlation test. All statistics were
carried out with the Statistica 6.0 software package.

Results
The frequency of the nudging acts in dominant males was, in some cases, four times
greater than that in subordinate males, with the Relative Monopolisation Index
ranging from 0.05 to 0.9 (Table 2). The Relative Monopolisation Index was
positively related to the Relative Trait Index based on the DFH (Table 2 and Figure
2). Although there were also positive relationships between the Relative Trait Index
based on the other fin measurements and the Relative Monopolisation Index, these
were not statistically significant (Table 2). Similarly, there was no statistically
significant relationship between the Relative Monopolisation Index and the Relative
Trait Index, as regards the vertical bar pattern, the body size difference (SL) and the
degree of asymmetry (Table 2). In the experimental trials where a higher Relative
Monopolisation Index was found (0.5–0.9, Figure 2), the dominant males had a
 Marine and Freshwater Behaviour and Physiology 163

Table 2. Mean frequency (acts/min) of nudging acts and mean size of sexual traits (cm), and
range in parenthesis, in dominant and subordinate competing males; mean values (and range
in parenthesis) of Relative Monopolisation Index (RMI, based on the frequency of nudging
acts) and Relative Trait Index (RTI, calculated for each sexual trait); and Spearman’s
correlation coefficients of the relationships between RMI and RTI, n ¼ 8.

Dominant male Subordinate male RMI/RTI Rs

Nudging behaviour 0.67 (0.17–1.5) 0.13 (0.04–0.25) 0.54 (0.05–0.92) –

Traits
SL 37.8 (35–46) 36.6 (33–43) 0.05 (0/0.08) 0.23
CFL 0.78 (0.64–0.9) 0.79 (0.68–1) 0.009 (0.07/0.05) 0.54
CFW 1.05 (0.8–1.3) 1.01 (0.7–1.3) 0.005 (0.03/0.02) 0.47
CFA 0.72 (0.58–1) 0.7 (0.4–0.83) 0.004 (0.06/0.06) 0.28
DFH 0.65 (0.45–1) 0.6 (0.3–0.8) 0.014 (0.02/0.05) 0.976*
DFA 0.6 (0.3–1.22) 0.55 (0.2–0.87) 0.01 (0.02/0.07) 0.357
AFA 0.58 (0.38–0.9) 0.49 (0.25–0.8) 0.025 (0.02/0.1) 0.42
VBS (left) 0.18 (0.14–0.22) 0.18 (0.14–0.21) 0.004 (0.02/0.02) 0.28
VBW (left) 0.07 (0.05–0.08) 0.07 (0.05–0.09) 0.0009 (0.003/0.008) 0.45
NVB (left) 9.6 (7–12) 9.1 (8–11) 0.1 (0.57/0.85) 0.14
AI 0.014 (0–0.05) 0.04 (0–0.14) 0.01 (0.014/0.08) 0.23

Notes: SL: standard length, CFL: caudal fin length (cm); CFW: caudal fin width (cm); CFA:
caudal fin area (cm2); DFH: dorsal fin height (cm); DFA: dorsal fin area (cm2); AFA: anal fin
area (cm2); VBS: average vertical bar span (cm); VBW: average vertical bar width; NVB:
vertical bars number; AI: Asymmetry Index.
*Significant correlation at p 5 0.05.

dorsal fin height of 1–2 mm larger than the subordinate males (Figure 2).
Nevertheless, in some of the trials, the dominant male was the smallest male of
the pair in terms of SL (Figure 2).
Within the sample of field-collected males, there were negative, statistically
significant correlations between some fin traits (specifically the area of dorsal, anal
and caudal fins, Table 3) and the vertical bar width (Table 3), whereas there was
neither statistically significant correlation between fin traits and bar span, nor with
bar number. Thus, males with larger fins within the population tend to possess
thinner bars (Table 3). The Asymmetry Index was negatively related to the size-
adjusted area of fins, but these relationships were not statistically significant
(Table 3).

Discussion
The results of our behavioural experiment suggested that male dominance is related
to the relative height of the dorsal fin, but not to the differences in the vertical bar
pattern or in the total body size. According to McGhee et al. (2007), male mating
success in a killifish breeding system is determined by the interaction between male
interference and mate choice, as the female may exert control on the latency to
spawn. Similarly, in our competitive arena, the ability of a male to monopolise the
female and to spawn with her, keeping the other male from courtship and spawning,
could be the result of the combination of male–male aggression and female control.
164 S. Malavasi et al.

Figure 2. Tridimensional representation of the relationships between the difference in fin

dorsal height (cm, y-axis), in body size (SL, cm, x-axis) and the Relative Monopolisation
Index (z-axis) between the dominant and subordinate male in the eight competitive
encounters.

Table 3. Spearman correlation coefficients of the relationships

between size-adjusted fin traits (see text for details), vertical bar
traits and asymmetry index in a sample of field collected males
(n ¼ 77).

Vertical bar pattern

Fin size NVB VBW VBS AI

CFL 0.02 0.09 0.02 0.017

CFW 0.08 0.14 0.08 0.15
CFA 0.11 0.25* 0.11 0.001
DFH 0.02 0.07 0.09 0.19
DFA 0.12 0.26* 0.04 0.13
AFA 0.17 0.36* 0.11 0.07

Notes: CFL: caudal fin length (cm); CFW: caudal fin width (cm);
CFA: caudal fin area (cm2); DFH: dorsal fin height (cm); DFA:
dorsal fin area (cm2); AFA: anal fin area (cm2); VBS: average vertical
bar span (cm); VBW: average vertical bar width; NVB: vertical bars
number; AI: Asymmetry Index
*Significant correlation at p 5 0.05.
 Marine and Freshwater Behaviour and Physiology 165

The relative difference in dorsal fin size between the two males may therefore have
some effects on both components of sexual selection. The understanding of the
relative weight of each component was beyond the aims of the present study.
The height of the dorsal fin could be related to the ‘fin flare’ behaviour that
occurred during intramale aggression in the Mediterranean killifish, consistent
with what has been observed by Fuller (2001) in a fundulid species. Larger and
more elevated dorsal fins may confer an advantage on a male when deterring and
repulsing other males, as well as constituting a preferred trait by the female, with the
net result of a higher degree of female monopolisation by that male. Fin size and
shape have been investigated in fishes as male SSTs, which may play a role both
in intrasexual and intersexual selection: for example, the sword in the genus
Xiphophorus (Benson and Basolo 2006) and the fin height in Rhinogobius (Suk and
Choe 2002).
These sexually selected traits may, in some cases, act antagonistically towards
other natural selection forces, making conspicuous males more vulnerable to
predation, as shown by Basolo and Wagner (2004) in Xiphophorus helleri. In other
cases, the fin modification may enhance locomotor performance, creating a positive
feedback between sexual and other natural selection forces (Oufiero and Garland
2007). Evidence that the sexually selected traits, such as elevated fins, may confer
swimming advantages has been, for example, recently shown in Paramonacanthus
japonica (Kokita and Mizota 2002). In the killifish A. fasciatus, males with larger
caudal fin areas have been found in habitats under higher predation pressures,
compared with habitats under lower predation pressures, where males had relatively
smaller caudal fins (Maltagliati et al. 2003). These authors suggest that larger
fins may enhance anti-predator performance, favouring rapid evasive manoeuvres.
Our results may partly confirm this hypothesis, as rapid manoeuvres are also needed
during male–male competition. In this competitive breeding system a male has to
obtain access to the female more rapidly than their rivals and to maintain it as long
as possible. Larger fins may therefore show a dual function. They make the male
more conspicuous due to the brilliant fin colouration (intense yellow in this species),
but they could also enhance the efficiency of rapid movements that appear to be
important in this kind of competitive system. By contrast, our results indicate that
no component of the vertical bar pattern correlates to male dominance and male
success. This is partly consistent with the results obtained by Morris et al. (1995) in
the swordtail X. multilineatus, indicating that the number of vertical bars is not
related to male mating success. However, our results from the correlative analyses
performed on the field-collected males suggest some relationships between fin size
and vertical bar pattern. In particular, males with larger fins tend to have thinner
bars. According to Morris et al. (2001), the bar frequency (number of bars/bar span),
rather than bar number, could be assessed by female choice in the swordtail, and this
seems to be a more relevant component of the vertical bar pattern also in the
Mediterranean killifish. Although the matter is controversial, the degree of symmetry
has been considered an honest signal of male mating success, being correlated with
the male genetic quality, developmental stability and providing a cue for mate
discrimination (Moller and Pomiankowsky 1993; Watson and Thornhill 1994). Our
results indicate a tendency towards a higher degree of vertical bar symmetry in males
with larger fins, but as these relationships were not statistically significant, further
investigations are needed on this issue.
166 S. Malavasi et al.

The combination of these results with those from our behavioural experiment
suggest that males with a relatively higher mating success in the population could be
those with relatively more conspicuous dorsal fins and thinner vertical bars along
the body.
The relationships between fin size and some components of vertical bar pattern
could be the result of allometric constraints underlying the development of this
multiple set of traits, but this hypothesis should be tested in future studies focused to
address the question. Further experimental studies are needed especially to assess
the functional role of vertical bars, a recurrent trait in killifish species, as bars may
act in the context of both natural (camouflage, for example) and sexual selection.

Acknowledgements
We thank Prof. Carol E. Johnston (Auburn University, AL, USA) for her critical review
of this article. Experiments and handling of fish followed the guidelines provided by the Italian
laws on the use of animals for experiments (Decreto legislativo n. 116, 1992).

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