International Journal of Ichthyology - Filer

aqua
International Journal of Ichthyology
Vol. 14 (3), 10 July 2008
Aquapress
ISSN
0945-9871
aqua
International Journal of Ichthyology
Vol. 14 (3), 10 July 2008
Contents:
Gerald R. Allen, Joshua Drew and Les Kaufman: Amphiprion barberi, a new species of anemonefish
(Pomacentridae) from Fiji, Tonga, and Samoa......................................................................................................... 105-114
Edward O. Murdy: Paratrypauchen, a new genus for Trypauchen microcephalus Bleeker, 1860,
(Perciformes: Gobiidae: Amblyopinae) with a redescription of Ctenotrypauchen chinensis Steindachner,
1867, and a key to ‘Trypauchen’ group genera ......................................................................................................... 115-128
Gerald R. Allen, Joshua Drew and Paul Barber: Cirrhilabrus beauperryi, a new wrasse (Pisces: Labridae)
from Melanesia ........................................................................................................................................................ 129-140
Kapil Mandrekar and Ronald G. Oldfield: Prior residency and social experience in contests between
similar-sized juvenile black Midas cichlids, Amphilophus astorquii ........................................................................... 141-148
John E. Randall1, John L. Earle and Luiz A. Rocha: Xyrichtys pastellus, a new razorfish from the southwest
Pacific, with discussion of the related X. sciistius and X. woodi ................................................................................ 149-158
Matthew T. Craig, John E. Randall and Mark Stein: The Fourspot Butterflyfish (Chaetodon quadrimaculatus)
from the Philippines and the Solomon Islands, first records for the East Indies and Melanesia ............................... 159-164
Papers appearing in this journal are indexed in: Zoological Record;
BioLIS – Biologische Literatur Information Senckenberg;
www.aqua-aquapress.com; www.aquapress-bleher.com; www.aquageo.com; www.Joachim-Frische.com
Cover photo: Amphiprion barberi, underwater photograph of subadult, about 50.0 mm SL, Namena Island, Fiji, 5 m depth.
Photo by G. R. Allen.
A view of the crater walls from the shore of Lake Apoyo. See page 143-150. Photo by R. G. Oldfield
aqua, International Journal of Ichthyology
Prior residency and social experience in contests between similar-sized
juvenile black Midas cichlids, Amphilophus astorquii
Kapil Mandrekar1 and Ronald G. Oldfield2
1) Department of Biology, Albion College, 611 East Porter St., Albion, MI 49224, USA.
Current Address: Department of Biological Sciences, Arkansas State University,
P.O. Box 599, State University, AR 72467-0599, USA
2) Museum of Zoology, University of Michigan, 1109 Geddes Ave., Ann Arbor, MI 48109, USA.
Current address: Department of Biology, Case Western Reserve University, 10900 Euclid Ave.,
Cleveland, OH 44106, USA. E-mail: [email protected]
Received: 14 October 2007 – Accepted: 17 April 2008
Abstract
The possible roles of chemical cues and a 3-D structure in
eliciting a prior residency effect in juvenile black Midas cichlids, Amphilophus astorquii, were tested under laboratory
conditions. The effect of recent social interaction on the
outcome of contests was also tested. Fish with a clay pot in
their pre-test tanks defeated opponents without such prior
experience when test tanks contained an identical pot. Experience in a small group of conspecifics had no significant
effect on contest outcome, although fish that held lower size
ranks tended to lose contests. The subjects’ chemical cues
did not result in a prior residency effect. Interestingly, juvenile Amphilophus cichlids are not typically aggressive in their
natural environment. The current results demonstrate that,
despite this, they are sufficiently plastic to behave aggressively when resources are made to be defensible by artificially
reducing attack distance and number of competitors. In
addition, the ability to modify aggressive behavior according
to prior experience demonstrates a further degree of plasticity. This suggests that complex behavioral plasticity relating
to aggression may generally be of such high adaptive value
that it might occur in species for which it might seem to typically have little utility.
Zusammenfassung
Unter Laborbedingungen wurde bei jungen Schwarzen
Midas-Cichliden, Amphilophus astorquii, untersucht, wieweit
chemische Merkmale und räumliche Strukturen sich als
frühere Wohnraum-Erfahrung auswirken. Außerdem wurde
die Auswirkung früherer sozialer Interaktionen auf das Konkurrenzverhalten untersucht. Fische mit einem Tongefäß in
ihrem Vortest-Aquarium besiegten Gegner ohne eine solche
Erfahrung, wenn sich im Testaquarium ein entsprechendes
Gefäß befand. Erfahrungen in einer kleinen Gruppe von Artgenossen hatten keine signifikante Auswirkung auf das Konkurrenzverhalten, wenn auch Fische mit einem niedrigeren
Größen-Rang der Tendenz nach eher unterlagen. Die chemischen Merkmale spielten keine Rolle beim Effekt der früheren Wohnraumerfahrung. Interessant ist, dass junge Am141
philophus-Cichliden in ihrer natürlichen Umgebung nicht
typischerweise aggressiv sind. Trotzdem legen die neuen
Ergebnisse den Schluss nahe, dass ihr Verhalten ausreichend
plastisch ist, um dann aggressiv zu werden, wenn künstliche
Eingriffe die Angriffsdistanz und die Zahl der Konkurrenten
verringert haben und damit eine Verteidigung nötig machen.
Die Fähigkeit, das Angriffsverhalten entsprechend früheren
Erfahrungen abzuwandeln, ist ein weiterer Beleg für die Plastizität des Verhaltens. Das legt den Schluss nahe, dass eine
starke Plastizität beim Agressionsverhalten allgemein von so
hohem Anpassungswert ist, dass sie auch bei Arten auftritt,
bei denen sie anscheinend wenig Nutzen bringt.
Résumé
Les rôle possibles de signaux chimiques et une structure 3D
pour obtenir un effet de séjour préalable chez les juvéniles
d’Amphilophus astorquii ont été testés en laboratoire. L’effet
d’interactions sociales récentes sur la survenue de combats a
aussi été testé. Les poissons qui avaient un pot de terre cuite
dans leurs bacs de prétest battaient les rivaux qui n’avaient pas
connu cette situation quand les aquariums du test lui-même
contenaient un pot identique. L’expérience menée sur un
groupe de congénères ne montrait pas de résultat probant sur
l’issue du combat, même si les poissons d’un rang hiérarchique inférieur avaient tendance à perdre. Les signaux chimiques n’avaient pas d’effet de séjour préalable chez les sujets.
Chose intéressantes : les juvéniles d’Amphilophus ne sont pas
spécialement agressifs dans leur environnement naturel. Les
résultats les plus fréquents ont démontré que, malgré ce fait,
il sont suffisamment adaptés pour un comportement agressif
quand les ressources deviennent défendables par une réduction artificielle de la distance d’attaque et par le nombre de
compétiteurs. En outre, la possibilité de modifier le comportement agressif en fonction d’expériences antérieures
démontre un autre degré d’adaptabilité. Ceci suggère qu’une
plasticité comportementale complexe, en matière d’agressivité, peut être généralement d’une valeur adaptative telle
qu’elle peut apparaître chez des espèces pour lesquelles il
semble que cette faculté soit typiquement peu utile.
aqua vol. 14 no. 3 - 10 July 2008
Prior residency and social experience in contests between similar-sized juvenile black Midas cichlids, Amphilophus astorquii
Sommario
Il possibile ruolo di segnali chimici e di una struttura 3-D
nell’innescare un effetto di priorità territoriale nei giovani del
ciclide Mida nero, Amphilophus astorquii, sono stati saggiati
in condizioni di laboratorio. è stato anche valutato l’effetto di
un’interazione sociale recente sull’esito di competizioni. Pesci
posti in una vasca con un vaso d’argilla prima del test respingono coloro che non hanno avuto questa esperienza quando
l’esperimento è condotto in vasche contenenti un identico
vaso. L’esperienza in un piccolo gruppo di conspecifici non
aveva significativi effetti sull’esito della lotta, sebbene gli
esemplari di taglia inferiore tendevano a perdere la contesa.
Le tracce chimiche dei soggetti non comportavano un effetto
di priorità territoriale. Stranamente, i ciclidi giovani di Amphilophus non sono di norma aggressivi nel loro ambiente naturale. I risultati presentati qui dimostrano che, malgrado ciò,
essi sono sufficientemente plastici da comportarsi in modo
aggressivo quando le risorse rappresentano un qualcosa da difendere in presenza di una distanza d’attacco artificialmente
ridotta e di un certo numero di competitori. Inoltre, la capacità di modificare il comportamento aggressivo in base ad
esperienze acquisite dimostra un ulteriore grado di plasticità.
Ciò suggerisce che la complessa plasticità comportamentale
relativa all’aggressività può essere di elevato valore adattativo
è può essere bagaglio anche di specie per cui appare in genere
di scarsa utilità.
INTRODUCTION
Aggressive behavior in animals typically occurs in
defense of resources (Huntingford & Turner
1987). The outcome of aggressive contests between
two animals may be influenced by asymmetries
between contestants (Maynard-Smith & Parker
1976). Body size or weapons (claws, horns, teeth,
etc.) can be good predictors of contest outcomes
(Huntingford & Turner 1987). Prior experience
with the local physical or social environment may
also alter the probability that an individual will
defeat an opponent. A prior-residence effect – an
agonistic advantage to animals fighting in familiar
locales – has been well documented in fishes (e.g.,
Turner 1994, Chellapa et al. 1999, Johnsson et al.
1999). Prior social experience in a subordinate role
generally results in a decreased probability of
attaining dominance in subsequent contests, while
prior experience in a dominant role usually has little or no effect (Francis 1983). Interestingly, fishes
that are not especially aggressive in their natural
environments may nevertheless behave aggressively
when space or number of competitors are limited
(Grant 1993), as they may be under captive conditions. It is unknown whether such fishes modify
aggression according to prior experiences as do
some fishes that behave aggressively under natural
aqua vol. 14 no. 3 - 10 July 2008
conditions. The aim of the current study was to
determine if prior experience is important in influencing agonistic behavior in juveniles of a fish
species in which juveniles are known not to be
especially aggressive under natural conditions.
The behavior of Midas cichlids, Amphilophus citrinellus (Günther, 1864), has been well-studied
(Barlow 2000). Feeding territories are extremely
rare in freshwater fishes (Barlow 1993, Grant
1997), and under natural conditions Amphilophus
cichlids behave aggressively only while defending
breeding territories (Barlow 1976). Furthermore,
juvenile Amphilophus cf. citrinellus have been
observed in Lake Apoyo, Nicaragua not to be
attached to specific sites and performing only modest levels of aggression (Oldfield et al. 2006). However, in captivity, the enclosure limits the distance
required to deliver an attack, making it easier for
an individual to dominate a small number of competitors, and increased aggression is observed (Oldfield 2007). The effects of prior experience have
not been tested directly in Amphilophus cichlids,
but increased aggression after a period of residency
has been observed in adults in the laboratory, suggesting that prior residency may provide an advantage in combat (Barlow et al. 1986).
Amphilophus cichlids are capable of recognizing
some conspecific chemical cues (McKaye & Barlow
1976, Barnett 1977), therefore it was hypothesized
that chemical cues may be important in producing
an aggressive advantage gained through prior residency. An individual’s own chemical signature may
be present in its aquarium, and it was expected that
subjects tested in their own aquaria would defeat
opponents taken from foreign aquaria. Small juvenile Amphilophus cichlids remain close to 3-D structure (rocks) in their natural habitat (Oldfield et al.
2006); consequently, we inquired whether the presence of a possible shelter would result in a prior residence effect (Nijman & Heuts 2000, Johnsson &
Forser 2002). It was also expected that increased
motivation to defend a residence would decrease
latency to initiate aggression and that contest asymmetries associated with prior residency would result
in shorter fight durations than in contests involving
equally naïve fish.
The effects of prior social experience were investigated by pitting subjects that had recently been
held in the presence of conspecifics against opponents recently held in isolation. Recently subordinate subjects were expected to lose more often than
isolated fish, while recently dominant fish were
142
Kapil Mandrekar and Ronald G. Oldfield
Table I. Body size in subjects. Mass (g) was not significantly different between winners and losers within each
treatment (paired t-test).
winners
losers
mean±SD mean±SD
Treatment #
n
t
p
Control
Prior residency
1 (Chemical)
2 (Clay pot)
Social experience
3 (Dominant)
4 (Subordinate)
10
5.8±2.0
5.8±2.1
1.406 0.193
10
12
5.7±1.6
8.2±4.2
5.7±1.7
8.1±4.1
0.264 0.798
2.017 0.069
12
12
9.7±2.4
6.0±3.0
9.6±2.3
6.0±3.0
0.209 0.838
0.290 0.777
expected to win contests equally as often as isolates,
as this pattern is commonly observed in other fish
species (Francis 1983). It was expected that subordinate social experience would decrease a subject’s
motivation to fight, resulting in longer latencies to
initiate combat and shorter fight durations than
would be expected in control contests that
involved two individuals without recent social
experience.
MATERIALS AND METHODS
Two broods of Amphilophus cichlids (Fig. 1) of a
form recently described as A. astorquii (Stauffer et
al. 2008), notable for their solid black breeding
color, were produced at the University of Michigan
Museum of Zoology (free-swimming 30 October
2005, 23 November 2005) from one male and one
female collected in Lake Apoyo, Nicaragua (Fig. 2).
A general description of the laboratory has been
provided previously (Oldfield 2005). Each brood
contained approximately 175 individuals, and was
housed in a single 416 l acrylic aquarium (L×W×H
= [168 × 88 × 27 cm] + [47 × 46 × 11 cm]). Fish
were fed once per day on commercial dry fish food,
supplemented once per week with frozen brine
shrimp (Artemia sp.) or frozen blood worms (Chironomus sp.). All aquaria were filled with aged tap
water (26 °C, pH 7.6 ± 0.2).
A single experiment, consisting of one control
group and four treatments, was conducted during
summer 2006. Fish were netted out and sedated in
an MS-222 solution, weighed, measured, and
branded with silver wire that had been cooled with
dry ice (for individual identification). One brand
was placed on the body of each fish, either anterior
to or posterior to the dorsal fin, with care being
taken not to disrupt the lateral line. Fish were
paired as closely as possible to an individual of the
143
same body mass and standard length (SL). Even
from a pool of 350 fish, exact matching was not
possible for some pairs. However, size differences
in these pairs were miniscule (mean ± SD differences between pair-members: 0.1 ± 0.1 g,
0.9 ± 0.8 mm; see Table I). Each pair-member was
isolated in a standard 38 l (51 × 26 × 31 cm) glass
aquarium containing 5 cm of natural gravel substrate covering an air-powered sponge filter. Standard length of subjects ranged from 40.0 to 80.0
(60.4 ± 9.1) mm. Cardboard sheets prevented
visual contact between fish in adjacent aquaria.
Fish were held in these pre-test aquaria for 2-3
days, except for one pair that was held for 4 days.
Trials began when two fish were netted from their
pre-test tanks and simultaneously introduced into
a test tank, each of which was set up identically to
the pre-test tanks, except as noted below. Contest
start time, time at which aggressive behavior began
(displays or attacks – see Baerends & Baerends van
Roon 1950), and time at defeat, were recorded.
Defeat occurred when one fish performed three
consecutive aggressive acts without an aggressive
response by its opponent. The winner was immediately netted out and identified. Each fish was
tested once. The water was replaced with fresh
water in pre-test and test aquaria before fish were
added.
In the control group all fish were held under identical pre-test conditions. With no experiential differences among combatants, fight outcomes were
expected to be random and confirm that background conditions of subject selection and maintenance did not influence aggressive behavior. One
member of each pair (n = 10) was randomly and
nominally designated as a subject, and the other as
its opponent. Both fish in each pair were netted
from their pre-test tanks and simultaneously added
to a third identical and novel test aquarium.
Treatment 1 was designed to determine whether
familiarity with a subject’s own chemical signature
would result in a prior residence effect. One member of each pair (n = 10) was assigned the role of
subject and the other designated as its opponent.
The two fish were netted from their tanks as in the
control, but instead of being transferred into a
third neutral tank both fish were simultaneously
released back into the tank from which the subject
had been taken.
Treatment 2 was performed in order to determine
if familiarity with a potential shelter would result
in a prior residence effect. The pre-test tank of each
aqua vol. 14 no. 3 - 10 July 2008
Prior residency and social experience in contests between similar-sized juvenile black Midas cichlids, Amphilophus astorquii
subject contained a terracotta pot positioned in the
middle of the tank, with its opening facing the
tank’s left wall, while the pre-test tank of each subject’s opponent did not possess a pot. The fish were
netted out of their pre-test tanks and placed into a
test tank (n = 12) that contained a pot identical to
the pot in the subject’s tank.
Treatments 3 and 4 were performed to determine
the effects of recent social experience on contest
outcome. The subject of each pair was held in a
pre-test tank with five or six other fish while its
opponent was held in isolation. All fish in each
social group had been weighed and measured so
that the relative size of each group-held subject was
known. In treatment 3, each subject was the largest
member of its group (n = 12). Body size is a good
indicator of dominance in Midas cichlids (Barlow
1983), so these subjects were considered to have
had recent experience in a dominant social position. In treatment 4, each subject was smaller than
the largest fish in its group (n = 12), and was considered to have had recent social experience in a
subordinate role. No effort was made to position
subjects at any particular subordinate rank. After
the pre-test period, each subject and its opponent
were placed simultaneously into a third aquarium
identical to the pre-test tanks.
For each treatment, the number of contests in
which a subject defeated its opponent was analyzed
for significance by considering the chance that it
would occur at random by using a binomial test.
Latency to begin aggression and contest duration
were calculated for each trial. For each treatment,
these values were compared to the control with the
non-parametric multiple comparison method
described by Zar (1999: 226). All statistical tests
were two-tailed and alpha was set at 0.05. Residence duration did not affect outcome of contests
within any of the three treatments in which it varied (2-tailed Fisher exact tests: control, p = 1.000;
treatment 1, p = 0.500; treatment 4, p = 0.491),
and neither did body size. There was no significant
difference in mass between winners and losers in
any treatment (paired t-test, Table I).
RESULTS
Results of all of the treatments are summarized in
Table II. In the control group, six of ten trials were
won by the arbitrarily designated subjects, confirming that fight outcome was random in the
absence of a difference in prior experience. In treatment 1, possible chemical factors did not result in
a prior residency effect. There was no difference in
latency to initiate aggression or in contest duration
Fig. 1. Juvenile Amphilophus astorquii. Photo by R. G. Oldfield
aqua vol. 14 no. 3 - 10 July 2008
144
Kapil Mandrekar and Ronald G. Oldfield
between the chemical effect trials and the controls.
In treatment 2, experience with a ceramic pot
caused subjects to defeat opponents held without a
pot. Latency to begin aggression in test tanks containing a pot was longer than in controls. Duration
of contests was not different than that observed in
controls. In treatment 3, social experience as the
largest member of a group did not affect the probability that an individual would win a contest.
Latency to the onset of aggression was not different
than in controls, and neither was contest duration.
In treatment 4, the number of fish with subordinate experience that won contests was not significantly lower than expected by chance. Latency to
initiate aggression was not different than that
observed in control tests, and neither was duration.
Each of the three of the subordinate winners had
held the rank as the third largest fish in its respective group. A post hoc analysis restricted to the
eight groups in which the test fish was smaller than
most of its group-mates (size rank 4, 5, or 6 in a
group of 6) showed that all eight subjects were
defeated by isolates (binomial test: p = 0.008).
DISCUSSION
The current results demonstrate that recent experience can affect the outcome of aggressive contests
in juvenile black Midas cichlids. Experience with a
flower pot caused subjects to win contests. This is
consistent with studies on other species that found
3-D structure or gravel to affect contest outcome
(Johnsson et al. 2000, Nijman & Heuts 2000,
Johnsson & Forser 2002).
In cichlids, including Amphilophus (McKaye &
Barlow 1976, Barnett 1977), parent and offspring
recognition (Myberg 1975) and mate choice (Plenderleith et al. 2005) may occur through chemical
signals. Therefore, it was expected that chemical
signals might play a role in establishing a prior residence effect. Figler & Einhorn (1983) suggested
that this occurs in convict cichlids, Amatitlania
nigrofasciatus (Günther, 1867). However, the current null finding for A. astorquii is consistent with
Bronstein (1985), who found that chemical cues
did not influence the outcome of contests involving Betta splendens Regan, 1910.
Recent social experience did not significantly affect
an individual’s ability to overtake competitors,
although experience as a relatively small member of
a group suggested reduced dominance potential, a
pattern consistent with several studies (McDonald et
al. 1968, Francis 1983, Beaugrand & Zayan 1985,
Beacham & Newman 1987). The ability of prior
dominance relationships to affect future social
Fig. 2. Male Amphilophus astorquii that was captured in Lake Apoyo, Nicaragua, and later fathered two broods of offspring
that were the subjects of this study. Photo by R. G. Oldfield
145
aqua vol. 14 no. 3 - 10 July 2008
Prior residency and social experience in contests between similar-sized juvenile black Midas cichlids, Amphilophus astorquii
Table II. Numbers of trials in which the subject defeated its naïve opponent, and its significance as determined by the binomial test. Also included are summary statistics for the latencies to initiate aggression and the contest durations for each treatment. For these measures, significant differences between each experimental group and the control group were tested with a
non-parametric multiple comparisons procedure (see text). NA = not applicable.
Treatment #
Control
Prior residency
1 (Chemical)
2 (Clay pot)
Social experience
3 (Dominant)
4 (Subordinate)
n
# won by
subject
p
latency, min.
(mean±SD)
p
duration, min
(mean±SD)
p
10
6
0.754
1.71±0.82
NA
20.59±15.80
NA
10
12
6
12
0.754
<0.001
4.05±6.27
12.98±15.81
>0.50
0.02>p>0.01
41.89±20.34
7.97±8.07
0.50
0.20>p>0.10
12
12
5
3
0.774
0.146
4.14±2.42
5.41±5.35
0.20>p>0.10
0.20>p>0.10
20.75±20.10
24.91±30.01
>0.50
>0.50
encounters may be an example of a more general
ability to behave according to recent social experience. Barlow et al. (1975) observed changes in social
behavior in Midas cichlids as they became more
familiar with their social environment. At the begin-
ning of an experiment, aggression directed toward
fish that shared the color of a large dominant fish
was lower than expected. By the third day the effect
of color-sharing had diminished.
There was extreme variability within treatments
Fig. 3. A view looking into Lake Apoyo, showing its clear water, typically rocky substrate, and steep gradient. Photo by R.
G. Oldfield.
aqua vol. 14 no. 3 - 10 July 2008
146
Kapil Mandrekar and Ronald G. Oldfield
in latency to initiate aggression and in contest
duration. The only significant difference observed
between the control and an experimental treatment
involved the ceramic pot. In treatment 2 the
increased latency may be explained by the structure
blocking visual contact, which may have initially
resulted in fewer opportunities for aggression
(Kalleberg 1958, Bronstein 1983). Although the
durations of contests involving a pot were not significantly different than in controls, the mean
duration was much lower than in any other treatment, suggesting that residency might result in an
asymmetry that could decrease contest duration.
The increased chance of winning an aggressive
contest after experience with a ceramic pot suggests
that subjects may have identified an identical pot
as a resource to be defended. However, the possibility remains that some other aspect of experience
with the pot increased the subjects’ agonistic success. Subjects experienced with a pot might have
defeated naïve fish even if tested in the absence of
a pot. Also, the sudden introduction of a novel
flower pot might have reduced aggression in potnaïve fish. Nevertheless, the 3-D structure seemed
to provide a shelter that subjects had identified as a
resource (Riechert 1979), whereas intruders did
not recognize the value of the structure and were in
a position in which they could not fight and evaluate it simultaneously (Johnsson et al. 1999).
Experiential factors that might increase the chance
of defeating an opponent include investment in
reproduction, knowledge of food, knowledge of
hiding places, and established territorial boundaries with neighbors (Maynard Smith & Parker
1976). The subjects used here were not reproductive, they had similar knowledge of food acquisition, and they had no neighbors, leaving knowledge of the shelter as a logical explanation for the
enhanced aggressive behavior observed. In any
case, the evidence demonstrates that animals that
are not remarkably aggressive under natural conditions may have the ability not only to behave
aggressively under certain conditions, but to modify aggression according to prior experiences, as do
some animals that are typically aggressive in
nature. This suggests that complex behavioral plasticity relating to aggression may generally be of
such high adaptive value that it might occur in
species for which it might seem to typically have
little utility.
147
ACKNOWLEDGEMENTS
Jeffrey McCrary and Lorenzo López Pérez helped
in Nicaragua to collect the wild fish that spawned
the subjects used in the current experiment. Paul
Bronstein provided advice in all aspects of the project including experimental design, data analysis,
and manuscript development. The manuscript was
improved by comments made by Gerald Smith,
Peter Esselman, and William Fink. Funding was
provided by the Museum of Zoology and the
Department of Ecology and Evolutionary Biology of
the University of Michigan, and the American Cichlid Association’s Guy Jordan Research Award. Animal husbandry and experimentation was carried out
according to U.S. animal use laws and was approved
by the University of Michigan’s University Committee on Use and Care of Animals, approval #8362.
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