171-181
Odonatologica 26(2):
June
I.
1997
Variation in non-territorial behaviour in male
Calopteryx splendens xanthostoma (Charpentier)
(Zygoptera:
Calopterygidae)
S.J.
Department of
Sheffield
Received
Male
April
have
assumed
variation
are
towards
and
that non-territorial
or
classified
had been
as
conspecific
males constitute
displaced
from
a
one
they
males and
fight
wait for territories
5,
2 alternative
individuals.
dependent
1996
mate-securing
Previous
category. This
whether
upon
studies
describes
study
territory. Consequently,
pre-territorial or post-territorial. Pre-territorial
territorial
instead
rarely fight;
Accepted September
non-territorial
non-territorial behaviour which is
in
Kingdom
demonstrate
commonly
of Sheffield,
University
United
2UQ,
1996 / Revised
either territorial
as
non-territorial male
males
29,
S10
damselflies
calopterygid
tactics, occurring
Plaistow
Animal & Plant Sciences,
or
not the
non-territorial
males
are
agonistic
to obtain territories. Post-territorial males
to become
vacant.
INTRODUCTION
Calopterygid
trating
on
damselflies have been the focus of much research, often
aspects of their reproductive
biology
(JOHNSON,
concen-
1962; PAJUNEN,
1966; HEYMER, 1973; WAAGE, 1973, 1979a, 1979b, 1987, 1988; FORSYTH &
MONTGOMERIE, 1987; MARDEN & WAAGE, 1990; MARDEN & ROLLINS,
1994; SIVA-JOTHY &TSUBAKI, 1989a, 1989b; SIVA-JOTHY etal., 1995). One
of the
most
reproductive
by
interesting
features of
behaviour
or
calopterygid mating systems
alternative
males. These have been
reported
(WAAGE,
1973; FORSYTH
HIGASHI,
1981; SIVA-JOTHY &
&
mate-securing
for
a
large
tactics
number of
MONTGOMERIE,
TSUBAKI,
is the variation in
commonly
exhibited
calopterygid species
1987; PAJUNEN,
1966;
1989a). As in other calopterygid
damselflies, male Calopteryx splendens xanthostoma demonstrate two alternative
mating tactics;
males
Previous work
on
are
either territorial
or
non-territorial.
odonates has shown that (i) males
and non-territorial tactics within
a
may
employ
lifetime (TSUBAKI & ONO,
both territorial
1986; FORSYTH
172
S.J.
& MONTGOMERIE, 1987;
increases with
the
(ii)
Plaistow
of non-territorial males
proportion
increasing population density (WAAGE, 1972;
1986; CONVEY,
1989);
(iii)
territorial males
have
generally
TSUBAKI & ONO,
higher copulation
a
rate
(CAMPANELLA& WOLF, 1974;FINCKE, 1984; HARVEY & HUBBARD, 1987;
FORSYTH & MONTGOMERIE, 1987; CONVEY, 1989); and
males will
become territorial if
with
provided
(iv)
non-territorial
territories (WAAGE,
vacant
1973;
CONVEY, 1989). All these observations suggest that territoriality is the favoured
mating
tactic and
provides
increased fitness benefits.
Non-territorial males become territorial in
vacated
owner.
territory
they
or
There has
enter
been
previously
no
that examines variation in the way
because the
surprising
energetically costly,
a
vacant
Because
longer
a
detailed
as
male waits
individual male C.
likely
are
possible
is manifest
territory
increase the
with
of
two
of
study
either occupy
they
ways:
escalated
an
with
fight
damselflies
calopterygid
risks of
lost
injury
or
reproductive
obtain
to
splendens
for
fighting
be very
to
a
territory,
death, the
costs
opportunity,
a vacant
occur
as
one
of
of
males)
1973; FORSYTH
WAAGE,
MARDEN & WAAGE,
a
territory
reproductive
ences
the
on
which will
both
adopt
able
(ii)
to
and
physically
STUDY SITE.
shallow
each
The
-
in
southern
fast
flowing
site all
water flow
to
0.4
as-
1987; MARDEN &
average, be older than
on
have
(e.g. PAJUNEN, 1966; HIGASHI,
Similarly, experience
mate-securing
what
pre-territo-
and
a
male
to
defend/
knowledge
post-territorial
of the
males. Differ-
extent
1994).
The aims of this
tactics of male C.
pre- and
study
splendens
post-territorial
males
were
(i)
to
xanthostoma
are
distinguish-
behaviourally.
study
stream,
ms
(post-territorial
calopterygids
allied with it, have often been associated with varia-
changes
France
territories
(>
and
same
MATERIAL AND
Vidourle
are
territory (pre-territorial males),
site will also differ between pre- and
examine
matured
1990) has suggested that the ability of
describe the alternative
there-
and
& MONTGOMERIE,
tion in behaviour (see ANDERSSON,
and
recently
territorial
must
calopterygids (FORSYTH & MONTGOMERIE, 1987;
declines with age.
in age, and the
is
for
waiting
a cost
and defended territories
Post-territorial males will,
1990).
rial males. Previous work
regain
are
a
have
Previous studies of
displaced.
sumed that all non-territorial males
1981;
obtain
previously occupied
and have since been
WAAGE,
to
that
fighting
territory.
xanthostoma may
males
types:
but have yet
reproductively active,
those males that have
two
waiting/
or
different. Whilst
and non-territorial mating tactics within a lifetime, non-territorial males
fore
a
territory
a
non-territorial males obtain territories. This is
associated with
costs
for vacated territories
searching
one
into, and win,
were
was
conducted between
(43°52’N, 04°03’E).
The
METHODS
June and
study
containing dispersed clumps
manipulatedso
') passing
over
them,
that
they
and
a
of
equal
1994
and
size
perch (15-20
1995,
on
of isolated
the
river
sections
of
pennicillatus (territories).
At
comprised
of Ranunculus
were
similar
August
sites
(0.5
cm
2
m
), with
an
upright stick)
equal
rate of
within
them
Non-territorial
SIVA-JOTHY et al.,
(see
at each
site
and the
callipers (Mitutoyo)
not have
few
between
of
sites
not
individuals
into
based
1994)
Four
age
the
on
categories
the nodus
showed
pery’
sites
distinguished
were
could
most
be
matured
ing
territory
a
males
reproductive
males
were
failing
to
recorded
territory
Detailed
site.
(i)
had
index
previously
males.
the
40 km radius:
(see
not used
in
film
on
records
a
that
of the
start
stiffness of the
subcostal
the
wings
a
sites
migra-
it
season
placed
we
wings
and
that
which
well
of the
leading edge
veins).
All of these
age
fully developed wing pig-
flexible
were
inflexible,
of all
the
patrol
the
a
dorso-ventrally
were
were
only
at
the
wings
worn
flexible from
only
distal
that
tip
were
and
‘pa-
contact with the
body (caused by
recorded
individuals
the
which
individuals,
post-territorial
previously
males
the
study
pre-territorial
males
the
either age category
that had
As
suggesting they
supported by
was
as
and
observations.
marked,
individually
could not be
carried
specific
a
out between
sites,
between
new
tactics
territory),
vacant
‘non-territorial
water surface.
1
been
had
only
observation
or
re-
that
age category
observed
clearly assigned
2.
defend-
to either
10.00-18.00 h EST. The
males
were
or
pre-
or
(ii)
position
arriving, previously
recorded.
DIS
over a
patrol flights’ seen.
Non-territorial
single territory,
time non-territorial
either
field
analyses.
were
of
were
Pre-
-
Territory
marked
take-overs
(active displacement
of a
fight).
of the
territory.
recorded.
site
assumption
switching
(take-over
and continuous
un-marked
This
males at
to
patrolling
patrols
patrolling
male
occurred,
to
1978) used to
vacated,
Data
or were
were
assess
fight
the
from 543
over
the
are
and
low
occurred
flights,
through
all territories
present
whether
patrolled
their
rapid,
individual
aggregation
displaced from,
collected
These
patrols always
The number of territories males
SOUTHWOOD,
was
at the
undamaged wings
males, had
marked
as
propensity
of
where males
held
a
did
within
neighbouring
at
marking. Consequently
did not have
harder
wings
individuals,
young
escalated
a
included
patrol,
cases
condition.
above
acquisition
In
hard
males at
observations
made
non-territorial
dispersion
1, had soft
newly arriving
TOV
an
from
the
which
2 males had
marking
all
defined
Daily
cm
ranged
Data collected
flights.
as
were
resulted in the
a
of
employed by
were
10-20
ageing
1990; SIVA-JOTHY & TSUBAKI,
WAAGE,
mainly by
tenerals,
class
class
and any males
either
records
and
defined
4, the oldest
and
were
involving
approximately
territories,
arrive,
as
holder
-
tactic
observations
the date of
AND POST-TERRITORIAL
MALES.
basis
were
OBSERVATIONS.
found
rarely
the stiffness of the costal end
calcium carbonate
PRE-
post-territorial categories
and
were
the site. Data from individuals
at
and
marking
digital
France).
isolated
males
of
cat-
age
of
pair
a
reproductive activity
found within
were
and their
pen,
mm) using
process
were
commence our
MARDEN &
also
males had
class
reproductive
pre-territorial
Post-territorial
a
of
identified
to
3
class
the
on
effectively
clearly
cently
(see
length. Age
had
region
DISTINGUISHING
were
0.01
(±
The
enamel
scheme.
largely by
slight wearing. Age
to the touch and
site
populations
(days) by
age
individuals, age
tip. Age
water in this
hard
did not
we
of adult males
their whole
to the
an
males resumed
cases
territories per site. All males
at 5-7
with
measured
particular
a
clearly distinguishable from
were
along
most
other
no
absolute
categories
mentation. The
youngest
flexible
Because
-
determined
turn
and
were
capture recorded.
in
at
maintained
hindwing
173
xanthostoma
therefore minimal.
following
age classes
wings (in
marked
was
on a
forewings
date of initial
to determine
possible
Calopteryx splendens
marked
individuals;
on
observation),
was
AGE CATEGORIES.
was
Male
Males
of release.
in 400 hrs
(twice
tion
overt effect
any
minutes
below).
(see
in
1995). Territory density
caught, individually
were
determined
egory
behaviour
or
not the
over was
non-territorial
the
a
the
patrol
noted, and
of non-territorial
territories,
at
that made
patrol
territory they
patrols,
made
by
89
174
S.J. Plaistow
RESULTS
TERRITORIAL BEHAVIOUR
Territorial males of
These males
were
site and
territory.
intruder
male
intruding
Fights
As
were
with C.
repelled
was
territory faithful,
surface
territory
and forwards,
perch
on
the
(often facing
the
over
s,
Escalated
the
after which,
fights
were
each
other)
increasing
volving
males
other
perching
one
and carried
a
on
distances
out
male.
an
fight developed.
after
50-100
encountering
above
cm
identifiable
or
from slow,
1988) which
site.
fight
stereotyped
several
‘see-saw’
up and
dis-
flights
ranged
territories and in-
neighbouring
spiralling chases, spanning
The duration of escalated
between bouts of fighting)
to
became escalated.
gradually speeded
with
(often overlapping
the
attempt
to
lasted for about
they proceeded through
as
the
short chases, backwards
Territory disputes generally
males), and culminated in high-speed,
perched
the
cases
occasion,
of the males would attempt
one
of the males retreated
clearly
reproductive
were
not retreat
approximately
territory. Occasionally
1966; WAAGE,
(see PAJUNEN,
the whole
of
majority
and
territory
male did
intruding
tinctive, hierarchical stages progressing
over
in the
on
male that entered
conspecific
1988),
sites.
itself or
territory
territory, following which, the opponent would repeatedly
physically dislodge
30-120
the
pursuit flight. However,
males hovered
two
on
emer-
oviposition
as
types: ‘territory disputes’ and ‘escalated fights’. Ter-
two
occurred when the
the resident male. The
perched
and chased any
brief (< 10 s)
a
females
by
in his intrusion into the
persisted
as
and
maculata (WAAGE,
after
classified
ritory disputes
used
pennicillatus,
nearby vegetation. They intercepted
their
xanthostoma defended isolated,
Calopteryx splendens
of Ranunculus
gent patches
the time
fights (including
from 360
to
8220
all females that landed on,
or
flew past, their
ranged
s
(x±se
=
2550±821.4 s).
Territorial males
attempted
to court
territory, although they rarely pursued
bounds of their
ering flight
the
just
territory. During courtship,
around the
forewings,
hindwings
son,
water
out.
were
female.
noticeably
held
relatively
manner.
this
or
in
nearby vegetation.
After
returning
to
guarding
beat
characteristic hovsustained
frequency.
to
body
hold their
(1995) provide
either occurred
the
In
using
compariin
a
males threw themselves
continuing
al.
courtship display. Copulation
close, non-contact, post-copulatory
wing
display,
et
out a
predominately
and away from the
stationary,
surface and floated with the current,
of the
was
increased in
Following
great distance outside the
males carried
Flight
HEYMER (1973) and SIVA-JOTHY
scriptions
self,
perched
which
tinctive ‘cross-like’
the
females for any
territory,
of their
more
on
the
disonto
hindwings
detailed de-
territory
it-
territorialmales showed
recent
mate.
Non-territorial behaviour
in
Calopteryx splendens
175
xanthoswma
NON-TERRITORIAL BEHAVIOUR
Non-territorial males did
bushes close
-specific; they
observation
As
not
defend
often
were
period
and in
seen
a
display
most
in
was,
pursuing
to
usually
rial
of
males
rarely
observation),
more,
the
to
took
territory
showed
and
flying
ovipositing
on
1973; FORSYTH & MONT-
s.
the whole, relatively unsuccessful and
flee these
place
xanthostoma showed
chasing
off the
copulation
no
intruding
courtship
male. Some-
attempts with the non-terri-
successful
in
getting
a
fe-
after which females
vegetation,
had been abducted. Non-territo-
they
behaviour towards these females (n= 1 in 400 h
guarding
typically
were
in bank-side
from which
retreated if
tolerated the presence of
they
involved
were
her for considerable distances. On the few occasions (n=5 in
copulation
returned
sitesame
on
400 h of observation) when non-territorial males
male in tandem,
always
within the
C.
resulted in the territorial male
commonly
times, females would attempt
sites
perch
females that
maculata (WAAGE,
towards females. This tactic
torial male
grab
to
non-territorial male
1987),
were not
copulation attempts
resource,
attempting
C.
different
in the bank-side
perched
mobile over the entire reproductive site.
some cases were
site and
perch
defended territories. As
GOMERIE,
but
resource,
occupying
non-territorial males defended no
down from
a
other male’s territories. Non-territorial males
to
further-
challenged by territory holders;
non-territorial males.
conspecific
NON-TERRITORIAL PATROL FLIGHTS
Non-territorial patrol
flights
territorial males
although
tories. Non-territorial patrols
intruding
a
(iv)
a
male attempted
territory; (iii)
vacant
fight
49
an
an
the
sponsible
a
pre-
or
for 242
The remaining
a
male
was
tactic
a
one
repelled
female of which
obtaining
non-territorialmales,
over
neighbouring
of four
outcomes,
a
terri-
(i) The
male obtained
intruding
after
flights
observed
brief
198 flights
was
or
not
by
(36.5%)
were
(63.5%)
made
103
by
(0.9%)
pursuit flight;
361
were
were
(66.5%)
resulted in
were
a
associated
successful,
resulted in
fight.
unequivocally assignable
post-territorial
(29.9%)
for
males
were re-
pre-territorial
territorial males
non-territorial males whose
known.
(3.9%)
5
(20.6%)
post territorial male. Of these,
to
21
only
territory,
a vacant
resident males, and 112
flights (70.1 %), compared
bouring territories,
securing
resulted in
patrol flights observed,
resulted in the male
pursuit flights by
either
intruding,
flights
abduction of a female; (ii) the
abduction of
In total, 345 of the 543
to
made these
generally
intruding
non-territorial
attempted
(9.0%)
brief
characteristic of
occurred.
Of the 543
with
were
occasionally
previous
males.
defending neighalternative
mate-
176
S.J.
Plaistow
Fig. 1.
The
relationship
(forewing length) and
of initial
tor of
were
new
they
Day
I
Post-territorial males
-lerritorial males
declined
(Fig.
throughout
older and had
were
the
season
Pre-territorial males had
patrol flights,
fights
with
fights during
occupying
at
21st
the
study
25th
was
the
June
un-
first
site.
1995
1995.
August
than pre-
forewing length) progressively
as
(Spearman’s
a
r=-0.332, N=135, pcO.OOOl)
r,
territory
holders.
In
vacated territories without
pre-territorial
and
males
flights they
taking
post-territorial
number of territories
patrolled
made per hour
bution of non-territorial patrols
a
POST-TERRITORIAL MALES
significantly
comparison,
more
a
post-territorial
fight (Tab. I).
7.1 times
long
males showed
over
per hour,
(Tab. I).
over
A
or
males showed
preferentially patrolling
on
a
to
no
obtain
through
males avoided
more
was
to
a
a
territories
significant
by
dif-
obtain territories,
territory. However,
significant
difference in
comparison
of the
dispersion
territories revealed that (i)
the
patrols
over
and distri-
pre-territorial
territories;
non-random allocation of non-territorial
the last
non-
esca-
the number of non-territorial pa-
random distribution of non-territorial
post-territorial
There
territorial males
as
fight during
to
territories
non-territorial patrols, and obtained significantly
post-territorial
showed
AND
significantly greater propensity
and obtained
ference in the time taken for pre- and post-
trol
observed
was
marked
males
All
Methods).
were
season
day 58,
indica-
good
1).
-territorial
with
(see
significantly longer wings
BEHAVIOURAL DIFFERENCES BETWEEN PRE-
lated
were
a
AND POST-TERRITORIAL MALES
Male size (measured
(Tab. I).
arrivals
of the
male size
because unmarked
individuals
time
whilst
PHYSICAL DIFFERENCES BETWEEN PRE-
marking provides
seasonality
marked
between
time of season. Date
territory they
had defended
males
and
(ii)
patrols,
(Tab. I).
DISCUSSION
Differences
in age and
adult male odonate
size have
mating
frequently
been
correlated with variation in
behaviour (TSUBAKI & ONO, 1987; FORSYTH &
Non-territorial
in
behaviour
Calopteryx splendent
177
xanthostuma
Table I
A
comparison of
pre-
and
post-territorial
Male
Features
male
Calopteryx splendens
type
considered
Test statistic
pre-territorial
post-territorial
36
30
29.09±0.25
30.07±0.18
Number of males observed
(mm)
Wing length
un-paired
t
=
n,
2.733±0.14
2.043±0.12
Age
of takeover
TOV": 5
D1S“:
Fight
fight?
or no
fight;
no
Time to obtain
TOV”: 14
fight:
5
no
54.2±15
a new
3
8
light:
X
X
387.0±81
Mean
number
5.96±1.3
'hr
(male
over
2.59±0.38
l.48±0.2l
‘dispersion’
mean
territorial
random,
J
of
n,
<1
dispersed,
=
0.99±0.16
non-
patrol flights (1
2.78±0.76
0.0007
0.005
=
13.867
0.001
=
n,
=
=
n,
=
=
19,
17
0.0008
n,
=
26,
29
0.12
n,
=
26,
29
0.23
MWU-tesf,
=
>1
23,
=
9.950
MWU-tesf,
patrol flights (male'hr ')
The
n,
MWU-tesf,
n,
2.32±0.45
0.003
30
MWU-tesf.
')
of non-territorial
19
=
24
test.
3
18
n,
of territories
patrolled
=
=
X
territory (mins)
Number
t-test,
n,
3
DIS ; 5
22
fight:
=
P
X' test.
1
16
3.16,
MWU-tesf,
n,
Type
xanthostoma
n,
=
25
0.028
26
=
aggregated)
The
in
difference
compared
to
a
0.99±0.16
dispersion
random
in
difference
compared
to
a
2.78±0.76
dispersion
test,
of non-territorial
to that
Results
on
expected by
are
b
TOV,
“D1S,
■’The
takeover
active
of
chance
at
p<0.05,
Whitney
a
vacant
index
and indicated
in italics.
U-test.
territory.
displacement of
dispersion
stat
a
territory
is defined
as
holder
after
variance/mean
an
escalated
fight.
(SOUTHWOOD,
1978).
184,
N
0.018
21
1-sample
=
N for test
compared
significant
"MWU-test. Mann
test,
0.795
1-sample
=
=
N
17
Wilcoxon
0.38±0.05
the last
defended,
=
stat
for test
patrol flights
territory
70.5,
=
Wilcoxon
random
distribution
Proportion
1-sample
test, stat
for test
distribution
The
Wilcoxon
292,
=
25
0.001
178
SJ. Plaistow
MONTGOMERIE, 1987; GR1BB1N & THOMPSON, 1991). Whereas age differences
truly
may
in
ence
individuals
xanthostoma
be due
simply
throughout
largest individuals, having emerged
in time of
ence
obtain
is the
as
1992). If this
Age-related changes
declining
of
learning
have
1990)
that
to
fight
and
often
&
individuals. In
having
young
to
a
lower RHP, and
Consequently, they
about
seven
study
times greater for
Pre-territorial males
haps suggesting
that
An alternative,
differ in
changes
were
they
to
are
the
males of C.
to
fight
and
wait for territories
post-territorial
s.
and/or
of
likely
to
males
learning.
more
reproductive
likely
to
enter
avoid. WAAGE (1988) has
fusion of residency’
ever,
in C.
s.
larly,
a
lack of
possible
inter-
males in
1988; MARDEN
displaced by
to
become
to
to
a
to
older,
holders.
Further-
territory
was
males.
pre-territorial
non-territorial
&
younger
are
vacant.
obtain
patrols,
per-
males.
post-territorial
Post-territorial males know
site
or
and the
to
escalated
pointed
out
be
its
not
because
reproductive
a
value of
territo-
specific
individuals with
fights
were
may
they
do
will be the
commonly
not
be the
explain why pre-territorial
no
result, pre-territorial
not
know who
that, in situations where there is
that ‘confusion of residency’
does
As
the
neighbouring
reproductive
occupants.
between two males, escalated
experience
on
inexperienced
fights
xanthostoma, pre-territorial males
fights, suggesting
one
xanthostoma
compared
fight during
territory boundaries,
information about the
to
WAAGE,
territories de-
displace territory
superior competitors compared
territories. Pre-territorial males will tend
males may be
or
on
non-exclusive, explanation is that pre- and post-territorial males
experience
position
as
high RHP, enabling
a
often
site, and may have ‘information’about conspecific rivals
ries,
to
males.
studies
post-territorial
WAAGE,
show that the time taken
more
are
differ-
a
unable
experience
Previous
1994).
males have
1987;
post-territorial
may be forced
the results of this
more,
are
holders. Several studies (TSUBAKI & ONO,
may be unable
so
in the
defend and/or re-gain
pre-territorial
s.
be
be the result of ei-
to
lower RHP. If this is true,
a
MONTGOMERIE,
comparison,
result of
MONTGOMERIE, 1987; MARDEN &
WAAGE, 1990) have reported that territorial males
have
thought
or
ANDERSSON,
displace territory
1987; FORSYTH
a
post-territorial
as
of the behavioural differences between
pre- and
xanthostoma is that
s.
them
are
male’s ability
clines with age, older males
pretation
to
species (MILLER, 1983;
have been recorded
never
(see
a
be
is that smaller males
holding potential (RHP),
individual
suggested
of C.
true
also tend
Thus, the difference
season.
simply
other odonate
in male behaviour
(FORSYTH &
calopterygids
some
population
a
true, smaller individuals may have been observed
were
resource
the
in
case
pre-territorial males, but would
ther
1989); this is also
males may
possibility
odonates, size of
(PENN, 1951; BANKS &
season
earlier in the
post-territorial
emergence. A second
territories,
FINCKE,
the
1). Therefore, the oldest males in
(Fig.
the differ-
post-territorial behaviour,
seasonal effect. In many
a
1985; MICHIELS & DHONDT,
in size between pre- and
C.
to
decreases
commonly
THOMPSON,
the
be correlated with pre- and
size may
a
outcome.
‘conHow-
the aggressors in
explanation.
males
Simi-
normally
win
Non-territorial
and
fights
may be
resident males.
displace
these
important,
behaviour of pre- and
a
to
capture
brief defensive
may be
flights
flight
from
cases,
territory
holders. A
site.
site
to
poorly defended,
individual territories, sometimes
Pre-territorial males may therefore be
well
as
as
the
territorial males,
on
aggregated,
with
from. One
reason
male that
may
displaced
be that
from their
1987). Finally,
it may
it could be that males
In
are
conclusion, the results of this
between
adopt
RHP
any decline in
male
be of
to
to
re-gain
successful
to
vacate
the
high reproductive
‘recognised’
as
it (TSUBAKI & ONO,
sneaking
on
the holder and
are
at
study
demonstrate
their own
therefore
and behavioural
physical
post-territorial males; notably
very different
and male
were
neighbours.
differences between pre- and
ritorial males
that
displaced
pygmaea males that had been dis-
are more
still
at a
reproductive
flights
of
assessment
refusal of the
territory
dragonfly Nannophya
avoid conflict with
to
a
represent
random
males have information about the
territory repeatedly attempted
territories, particularly if they
able
at a
had been
territory they
rapid
a
if the male considers the
value. In the libellulid
placed
post-territorial
at
those territories. Post-
defending
for the
them that would enable
performance. Alternatively,
territory, especially
patrol flights
territories
assessing
patrol
undefended,
even
the other hand, showed non-territorial patrol
significant preference
a
or
at-
only
all territories present
visiting
of the males
fighting ability
not
were
elicited
flights
function of these
possible
territories. Pre-territorial males executed non-territorial
with respect
learning
the differences in
patrol flights
non-territorial patrol
information about
gather
to
and
experience
explain
to
of non-territorial
majority
most
unlikely
are
males.
post-territorial
females. In
in
Although changes
179
xanthosloma
Calopleryx splendent
alone
changes
The results show that the
tempts
in
behaviour
that pre- and post-ter-
territory acquisition strategies.
territory acquisition
strategies
is
The relationship
discussed elsewhere
(PLAISTOW & SIVA-JOTHY, 1996).
ACKNOWLEDGEMENTS
I
am
very
grateful
to RICHARD BAILEY, JANE RUSDEN, JOCK RUSSELL, ELLY SAUNDERS,
and NICOLA SEAL for their excellent
Dr MIKE SIVA-JOTHY, Dr
criticism
BBSRC
which
assistance in the field. DAVE BLAKE for technical assistance.
MATT SPENCER
greatly improved
earlier drafts
and Dr
of the
NICK COLEGRAVE,
manuscript.
This
provided
work
was
constructive
supported by
a
studentship.
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