Download how is interbreeding between species

Adv. Odomtol.
1:
Review
251-265
of
December
reproductive
31,
isolating barriers
1982
in
Odonata
K.J.
1949 Hickory
Premating
and
habitat,
ences
tive
in
a
as
researched,
and tactile
tandem
type
ence
of
Groups
mental
on
idae,
and
mating
has not been
dragonflies
which
not exist
Polythoridae,
the
isolation
are
is
need to be
effec-
sense
of
in
interpreted
some
barriers have not been
have been
reported.
actual data. Studies
and
genitalic
expected
Gomphidae,
classical
on
for
functions of the
which
sense.
experi-
Macromiidae,
various
are
the exist-
nongenitalic
but
Aeshnidae,
and
Megapodagrionidae, Pseudostigmatidae,
Protoneuridae. The exact
still
been
stimuli in Anisoptera
isolation in both
and
Separation of potential
isolating barriers far outweighs
ethological
important
in the
demonstrated.
has
Differ-
exchange.
gene
most
isolation,
(nongenitalic union)
importance of tactile
studies do
duliidae,
in
on
mechanical
in
the
comprise
several hybrid
although
the
stimuli
Odonata involve temporal,
interspecific
of mechanical isolation. Postmating
Speculation
needed
to
United States
35630,
barriers in
isolating
order. Mechanical
genitalia,
during
groups
Florence, A1
barriers
within the
incompatible
mates
reproductive
ethological
visual
barriers
Avenue,
Tennessen
Cor-
Platystict-
structures used
clarified.
INTRODUCTION
The
essence
interbreeding
such
groups
of
the
biological
populations
(MAYR
are
1970), prompts
a
by
was
devised
term, reproductive
cause
it
of
anism.
Knowledge
closely
related species
pered
gene
isolating
does not imply
derstanding
animal
isolation
The term
by DOBZHANSKY (1937)
which interspecific
an
the
of
exchange
a
is
major
to
is
function
barriers
exact
animals
groups
from
how is
encompass the
the
does the
isolating
does not exist.
especially
stumbling
more
used in this
as
in
block
of
other
isolating mechan-
prevented;
barriers (RIBs)
evolved
communication,
studies;
isolated
fundamental question:
interbreeding between species prevented?
ism
that
concept,
species
reproductively
the
ways
current
paper be-
term mech-
majority of
Difficulties
in
un-
insects, have hamhas
been how
iso-
252
K.J.
lating barriers arise
of
(modes
Tennessen
and
speciation)
the
roles of
relative
natural selection and chance in these processes.
Given
taxonomic status of the Odonata,
the advanced
size and
atively large
ease
of recognition
in the
field,
a
their rel-
fair knowledge
of RIBs in
dragonflies would be expected. Despite considerable infor-
mation
reproductive
on
needed,
was
there
of this review
poses
odonate RIBs,
to
gaps in
briefly
to
are
identify
in
guidelines
some
JOHNSON’S review (1962) of
which he pointed out
large
are
and
behavior,
isolation in
reproductive
of the group. The pur-
summarize
present knowledge
is
where study
areas
the
approaching
where research
areas
knowledge
of
study
needed,
RIBs
in
and
to
on
give
dragonflies.
STATE OF KNOWLEDGE
All
RIBs.
in
published
Almost
studies
nothing
I reviewed
Odonata, probably because dragonflies
and
will
(1965)
The
not
show promise
study
of
where
groups
crosses
are
of
ity
mate when
readily
that
RIBs
postmating
could
postmating
from
yet
barriers
eggs
known
be
to
interspecific copulations
two
species
of
be
especially
in
produced
rewarding
occur,
100 percent
Odonata. WAAGE (1975)
operative
as
mortal-
interspecific
by
in
copul-
demorsa, the only example of
based
on
his
assumed
observations
of
and the absence of obvious hybrids between
Calopteryx.
Discoveries
of
possible
hybrid
I) further point to the need for studies in this
(Table
techniques
frequently
JOHNSON (1975) reported
hatching
mortality
hybrid
difficult to hold captive
JOHNSON’S
attempts
ations between Ischnura damula and I.
premating
damselflies may be cultured.
some
tandem
with
postmating mechanisms
are
confined.
at least
interspecific
possible.
larvae
concerned
were
is known of possible
specimens
area.
PREMATING ISOLATING BARRIERS
The
cates
dominance of
its potential
mates,
and
barriers
in
vision
among
importance
indeed this
addition
to
a
as
appears to
optical
JOHNSON (1962).
In the
of RIBs,
the examples
are
senses
operative.
are
also
following
are
of
dragonflies
of recognizing
not intended
ation of the literature, but rather
erate.
means
be
stimuli
nised by
given
the
On
conspecific
the other hand,
operating,
as
discussion of the
as
a
indi-
recog-
types
complete compil-
used to show how the RIBs op-
Reproductive
isolation in
Table
A
list
of
Kiauta
dragonfly species
1967; Asahina
reported
to
have
253
Odonata
I
individuals
hybrid
produced
(main
sources:
1974)
GOMPHIDAE
GOMPHIDAE
Gomphus fraternus
fraternus
X
G.
G. externus
extemus
9
Calvert
Calvert
1901
1901
Gomphus
X
G. graslinellus
G.
graslinellus
(J
�
Williamson
Williamson
1903
Aeshna confusa
confusa
X
A.
A.
<J
�
Calvert
Calvert
1956
19S6
Anax imperator
X
A.
A. parthenope
parthenope
6
�
Bilek
BUek
1955
Anax
nigrofasciatus
Anax nigrofasciatus
X
A.
parthenope
A. parthenope
julius
julius
66
<J<3
Hiura
Hiura
1968
1968
Anax
nigrofasciatus
nigrofasciatus
X
A. parthenope
julius
julius
99
Hiura
1971
Anax nigrofasciatus
nigrofasciatus
X
X
A. parthenope
julius ��
julius
lividus
lividus
AESHNIDAE
AESHNIDAE
diffinis
diffinis
66
Asahina
Asahina
1974
1974
LIBELLULIDAE
L1BELLULIDAE
X
L. intacta
Intacta
�
6
Tennessen
Tennessen
1981
1981
X
X
L. luctuosa
luctuosa
larva
Wilson
Wilson
1920
elatum
elatum
X
S.e. eroticum
66 9
Asahina
Asahina
1974
pedemontanum elatum
Sympetrum pedemontanum
elatum
X
S.e.
5. e. eroticum
9
Miyazaki
1972
Sympetrum
Sympetrum
baccha matutinum
X
X
S.e.
S.e.
eroticum
eroticum
6
�
Yamamoto
1965
Sympetrum
Sympetrum
risi risi
X
S.e.
eroticum
eroticum
5
Asahina
Asahina
1974
1974
Bilek
BUek
1963
1963
Leucorrhinia
glacialis
Libellula pule
helia
pulchella
pedemontanum
Sympetrum
<sd9
<J<5V
COENAGRIONIDAE
6
�
Coenagrion pulchellum
X
C.
puella
puella
civile
Enallagma
Enallagma civile
X
X
E.
E.
carunculatum
carunculatum
12 different
SUMMARY:
SUMMARY:
species
species
pairs
involved,
24+
66
��
Williamson 1906
1906
specimens.
Temporal isolation.
The two modes of temporal isolation, i.e. differences in
and
son
diel
adult mating
reproductive
An
can
lier
example
overlap
appear
be
to
disappear
seasonal
related species
few
cases
would
have
of G.
where
,
wherein adults of G.
antilope
they
(unpublished
differences
through
time and
been
carried out,
the North Ameri-
furcillata
appear
flight
antilope
adults
G.
of
The literature
do
ear-
sea-
not
furcillata
is replete
with
relevance to
Data
relative
abundance
maturation, including
frequencies
ignored.
on
conspecific and heterospecific encounters,
are
inter-
Odonata.
although
species,
reproductive
prevent
adults
before
observations).
between
has
on
mature
G.
sea-
during the
species
1940). Although
sympatric,
are
reproductively
isolation
studies
a
effectively
flight
the time of
documented in
been
(GLOYD
in
nonoverlap
of seasonal isolation may exist in
ductive
of
a
Gomphaeschna
than those
sons
of
However,
genus
involve
readiness. Obviously the absence of
mating activity
breeding.
periods,
the importance
of
are
season
repro-
needed. Until such
in
Odonata repro-
254
K.J.
Tennessen
ductive isolation remains unknown.
On
diel
a
basis,
an
species of Corduliidae
agrionid
by
congeners
though
of E.
females
two
ity periods of
were
in
the
be
day
The
isolated
partly
(TENNESSEN
coen-
its
from
al-
1975),
observed being taken into tandem by males
MAY (1980)
pollutum.
to
appears
earlier
mating
isolation between two
offered by PAULSON (1973).
dubium
Enallagma
of temporal
example
was
differences in the daily activ-
reported
four species of Micrathyria, although
much overlap
oc-
curred
and
(1974)
showed distinct temporal differences in three damselflies, but
the
exact
species
isolation
Temporal
mating
is probably
lead
changes
to
in
widespread
not
Variation
overlap
function secondarily
appear to
have
evolved
not
to
be
a
Odonata,
in
breeding
directly
RIBs in
as
such
as
per-
amongst individuals
another barrier would be necessary. Temporal
case
probably
the isolation appeared
study
the smaller species.
relatively inefficient.
environmental
which
in
it is
and
on
pressures
MIZUTA’s
given.
not
were
congeneric
predation
haps because
and
not
were
response to
times
mating
in
times,
differences
some
groups,
(PAULSON
but
1973).
Habitat isolation
Often
to
referred
ulation
of
sympatric
differences may be
be
swarm-feeding
of
tinctly
species
separate
is
initiated;
exceedingly
(CORBET
habitat.
or
microgeographic
isolation,
differences in the microhabitats where cop-
simultaneously
present
types
ecological
as
habitat isolation involves
small.
during
in
terms of
certain
even
though
adult
microhabitats,
may
occur
larvae
behavior
species
such
activities,
1963), but mating
Conversely,
these
distance,
Adults of related
in
may
may
as
may
mixed
different
live
in
dis-
obliterate
the
difference.
Isolation
an
due
of
recognition
ethological
specific males
effected
in
via
a
habitat differences results
barrier.
differences
involve
placed
to
which prompted
and
in
because
However,
behavior
responses to
habitat
a
special
classify
it
hetero-
(i.e. isolation
and
is
as
does not
between
case
stimuli),
extrinsic
to
isolation
encounters
during
females, it represents
specific
from adult behavior,
GARRISON (1979)
is
therefore
distinct category.
Groups in which evidence of habitat isolation has been found include
phus
Enallagma
(BYERS
(JOHNSON
1962),
(TENNESSEN
1939),
1966,
1975, GARRISON
Tetragoneuria (TENNESSEN
VERDONK
Somatochlora (TAKETO
isolation is
an
inefficient
barrier,
1979),
1960).
as
the
1979), Progom1977), Ischnura
Leucorrhinia
In
these
potential
(PAJUNEN
examples,
habitat
for individuals to
Reproductive
transgress
habitat
encounters.
The
“boundaries”
is great,
making
likely interspecific
results of JOHNSON (1966)
experimental
that habitat
suggest
255
isolation in Odonata
changes could
cause
a
strongly
breakdown of segregation.
This breakdown necessitates the presence of another RIB.
Ethological isolation
Several
elucidated.
mates)
acts
In
meet
that
(based
examples of ethological
on
(1)
these
but
elicit
mating is
A wide
Visual stimuli.
found to evoke
others
described
in
1963;
CONSIGLIO
HOLTZ
they
ing
are
in
cused
displayed
on
their role
been
and
are
groups, but
(CORBET
in which
and ways
in
one
pair-form-
study has fo-
isolation
reproductive
BUCH-
(
Hemiphlebiidae
critical
only
HEY-
1961;
Platycnemididae
Color patterns
in
has been
Calopterygidae
1973), Chlorocyphidae
1966),
and
dragonflies,
families:
to the female
many of these
in the
means
be achieved.
Courtship
JOHNSON
1955;
1974),
1913).
can
in
response
discovered.
HEYMER
1956;
(TILLYARD
“recognition”
Zygopteran
1951,
1973; WAAGE
four basic
are
variety of visual stimuli have
to be
four
(BUCHHOLTZ
MER
There
the mating
bound
are
(potential
individuals
intrinsic differences
by
response.
by which species
isolation have been
behavioral
prevented
the copulatory
the senses)
or
heterospecific
barriers,
(WAAGE
1975).
Studies
and
nition”
ITO
1960;
of only
large,
dimorphism
and
the
one
of
acquisition
shown
a
mate
that
in
important
are
PARR & PARR
courtship
“recog-
sex
(JACOBS
1974). However,
I
(PAJUNEN
family
1964).
He
1955;
am
demonstrating visual isolation
study
diverse
have
Libellulidae
on
sexual
aware
in this
concluded
that males of two similarly colored Leucorrhinia species
females by
recognize
females
appear
sight stimuli
libellulid
to
prior
genera
and color-pattern
differences in abdominal
recognize
to
but
they
by
come
to
mind
differences
in
exist
which striking
amongst
color
the species
Libellula, Micrathyria, Rhyo-
have not been studied.
JOHNSON (1975)
proposed
that
male-like
andro-
morphs within Ischnura offer increased reproductive
lation,
as
he
and
tandem attempts. Numerous other
( Celithemis, Erythrodiplax,
themis),
males
size,
undetermined
found
male
mating
preferences
in
isotwo
256
K.J.
species.
Many
barrier.
in
likely
to be
and
UV
and
eyes
experimental
ocelli
of
differ-
as
discrimination,
are
patterns of ultravioseveral species
receptors
work
such
cues,
form
have been found in
1979),
compound
for
and
species.
for study
species
visual
detect
not
Enallagma
two
discovered. Distinctive
reflections
need
behavior
flight
BERGLIED
in
Other
did
(1979)
in
males
by
contain polymorphic
groups
possible
ences
let
GARRISON
However,
preferences
morph
this
Tennessen
have
(MENZEL
(SIL-
been
found
The
1979).
these various possible
on
modes of visual isolation is obvious.
(2)
Tactile
uli
stimuli.
function
Experimental
RIBs
as
males of Lestes
having
and
Ischnura
experimentally
NESSEN (1975)
by
was
havior
in
ERSON
(1982),
in the
altered
of males by
These
using
from
respect
of
experimental
on
from
a
under
1970).
female.
in
Tactile
tial
mates.
the
differences
contacting
a
stimuli
series of signals
step
In
the
The
in
study
to
(BARTH
differences
some
in
tac-
or
arise
rejection
groups
are
just
exchanged between poten-
Odonata studied
appear
one
of the organ
the behavior, which evokes acceptance
the
shape.
organ in
particular
species
one
with
In studies of other insects,
the
in
ap-
mechanism described
DOBZHANSKY
similar
bethat
differ in
isolation
the shape
be
1968;
approach
superior appendage
tactile
the tactile
even
and
ROBERTSON & PAT-
though
manner,
TEN-
provided
of Enallagma
stimuli consist of males “using”
specific
copulate
appendages.
evidence of tactile recognition
females based
examples
Fewith
shape of these appendages
an
other groups of insects.
may
LOIBL
by
release copulatory
the species.
found strong
Enallagma,
by
refused to
females of two species
specific differences
tile
tactile stim-
concluded that tactile stimuli
pear to be isolating
a
presented
the male superior appendages
basic
that
and KRIEGER & KRIEGER-LOIBL (1958).
(1958)
males
evidence
first
arise
thus far,
from
the
however,
shape
of the
organ(s), which is detected and discriminated
by females. This hypothesis involves behavior and merits
classification
appears to
serve
as
be
the
as
an
very
ethological
effective,
ultimate
barrier. Tactile isolation
and
premating
in
some
RIB
odonates may
should
other
de-
Reproductive isolation
in
257
Odonata
vices break down.
Olfactory
(3)
(4)
and
substances
sound
communicative
signals
in
effort
these
stimuli.
Auditory
nor
waves
in
is
areas
have
Odonata,
almost
visual acuteness of dragonflies
to the
approaches
long-range
ity
of
fact
study
females
detecting
material may
bond.
Is
material
contain
this
be
a
on
the
these
determined.
problems
proaches,
To
my
such
summary,
interspecific
between
superior
has
(see
ata
BICK &
may
exchange,
related
easily
BICK
the
be isolated by
there
and
could
or
the
could
it
are
of Enallagma
function has
devised,
and
may
be
other
ap-
of sound detection in
may
few
not
solving
be rewarding.
investigated.
thoroughly
to
for
procedures
possibility
been
not
species thought
1981).
their
chemical analysis,
as
that
the tandem
fluid,
appendages
1975) but
not
a
that appear to be secretory
ethological isolating barriers
gene
closely
are
knowledge,
dragonflies
In
Cells
seminal
Experimental
as
reported
mesostigmal plates
phenomenon
displaced
males (TENNESSEN
been
female
function in strengthening
dried,
in
rule out chemo-
the chemical and
producing
the
common
a
not
species of Argia. He postulated
pheromone?
found
were
be
weeks (any phero-
even
WILLIAMSON (1906)
it.
left
males of several
the
or
dissipated) does
males may
white substance
by
attention. The
deserves
attempt tandem when presented females
would have
as
of the
dominated
unlikely, the possibil-
seems
that have been dead for days
reception
as
research
Recognition
has obviously
chemoreception
that males
mone
nil.
implicated
although
of their biology. And although
chemoreception
contact
Neither chemical
been
effectively prevent
reports of copulation
ethologically
isolated
The majority of cognate species of Odon-
ethological
means, with visual and tactile dif-
ferences playing the major roles.
Mechanical isolation
Also
called
the
“lock-and-key”
under much criticism,
withstood
strated
scrutiny.
convincingly
ated that in
as
hypothesis,
few supposed
examples
this
in
Incompatibility of genitalia has
in any odonate
idea
has
come
the Insecta have
not been demon-
species. WATSON (1966) specul-
Tramea male hamules interlock with the female genitalia;
JOHNSON’S study (1972)
showed
that
these structures
in
Hagenius
258
K.J. Tennessen
support the penis
probably
on
lock
a
and key
how the genitalia
during
and held
engaged
are
in
tandem
because
appendages
of
CORBET (1963)
dence of this type of
mental evidence
of
supportive
the
Based
I
studies,
isolation
species,
on
genera
into tandem
upon
specific males, performed “refusal motions” and
had
no
1975). When presented
“difficulties
female
from
a
from
a
them.
PAULSON interpreted
isolation.
on
to the
“difficulty”
inappropriate
stim-
Separation of such pairs gives the
contact stimuli
mechanical barrier when
different
type
of
mechanical
ap-
behavior
and
isolation may be
using WAAGE’s discovery (1979a) that males
the
female’s
would work
but
bursa
as
copulatrix
follows;
a
A,
sp.
B; barring
sperm of the sp. A
remove
transferred. The hypothesis
the previous
remove
sperm
fertilizing
The
least
of
eggs
hypothesis
tandem
previous
is
taken
of isolation,
hypothesized
sperm stored in
remove
The
matings.
A mates with
into tandem by
copulation
is
the
(see
also
following
some
species
last
male;
no
sperm
would not
male’s
sperm
apply
the
facilitate
sperm
productive
a
barrier
male of
male of
or
length
from the sp. B male
if
a
are
male does not have
in order to transfer his
own
be-
male to mate appear to have precedence
WAAGE
1979a for
would not hold for
copulation.
If
this
discussion
and
those species
hypothesis
is
in
references).
ovipositing
in
for
at
correct
pairs, it would fit the classical definition of mech-
anical isolation in that genitalic incompatibility
much of
a
attempted;
the intromittent organ is not of the correct shape
however,
cause
she
other type
any
from
female of species
before oviposition,
sp.
to
the
However,
involved.
A
to
hetero-
released (TEN-
were
fitting of the male appendages
female discrimination based
heterospecific male.
pearance of
are
mechanical
as
Females
by
heterospecific females, males
dead
clasping
associated with the
thorax”
could arise
uli
in securely
difficulty
and
has been inter-
tactile stimuli.
being taken
experi-
observations
what
that
involve
may
presented
that he considered
own
my
believe
provide evi-
to
considered
coenagrionid
is
preceding
union
JOHNSON (1962)
isolation. PAULSON (1974)
hypothesis.
mechanical
NESSEN
on
how do hamules
females with anal
clasp
differences).
cited several studies
other
of two Enallagma
securely
to
anatomical
from several
of
understanding
as
needed in Anisoptera
linkage, the nongenitalic
copulation (i.e. inability of males
preted
is
together (i.e.
Another mechanical barrier which has been proposed
function?).
incompatibility
and
translocation, casting doubt
sperm
More work
operation.
is involved.
However,
morphology of the odonate penis probably evolved
displacement
isolation may be
(WAAGE
only
an
1982),
and
any
resulting
occasional consequence.
to
re-
isolation in
Reproductive
259
Odonata
BREAKDOWN AND REINFORCEMENT
The above premating
how
interspecific
riers
fail
ing
in
gene
is unknown.
sequence,
RIBs
is
habitat differences may isolate two sympatric
poral
or
most
part;
if
visually and avoid mating.
inefficient,
a
tactile
mechanical
or
these barriers would result in
mating
RIBs,
zygote
or
such
hybrid
bridization.
increase
in
specific
mate.
foregoing
selection
example,
males
distinguishable.
and
sites,
to
attempt
in these
tandem
chance of obtaining
wered
questions
speciate?).
isms
riers arise
into
come
little
or
are
supportive
possible
tropic
may
benefit
The
biggest
anisopteran
are
also
be
female,
such
been
evidence has
a
favor
females
are
those
versus
this
as
furtive
(i.e.
unans-
how do organ-
that isolating bar-
incipient species
1978),
been found (PATERSON
1978).
other
Further
gene
exchange
evolved
incidental
or
very
It
of
studies
of
is
pleio-
functions (ROBERTSON
advancement
population-genetic
are
for
the
increasing
stem from
or
near
1975, WHITE
to
an
con-
advantageous
more
thereby
postulated
1970, BUSH
&
speciation
theory
isolating
barriers
1981).
gap in
our
knowledge
and effectiveness
families Aeshnidae,
needed
females
it
(MAYR
to
heterospecific
how RIBs originate
has
securing
seem
via natural selection after
of
prevent hy-
phenomenon in
to
perfected
from
importance
would
may
CONTINUING THE QUEST
the
may
rare
Although
1982.
(TEMPLETON
a
devoted
Nondescript
mate. Problems
a
consequences
post-
case
different groups, females may
with any
that many barriers
PATERSON,
or
female. However, owing to dif-
groups it
concerning
contact
absent
Failure of
sequence, there appears to be
and energy
ferent sexual selection pressures in
breeding
inferiority
or
discriminate against
visually
are
1981).
time
of
For
stimuli
interspecific mating, in which
those that attempt to mate with any
be
tem-
species for the
intervene.
may
mating is apparently
in the
wastage
If visual
RIB
mortality, sterility
Interspecific
With each step
not
operat-
chromosome incompatibility (no fertilization),
as
Odonata (BICK & BICK
males that
as
bar-
heterospecific individuals meet, they may “recognize”
differences
may
should supposed
reinforcing another. Accordingly,
one
and
100% effective,
not
prevented
Nevertheless, RIBs have been viewed
series,
or
probably
are
exchange
on
of
-
STUDY NEEDS
of RIBs
ethological
Gomphidae
Cordulegasteridae,
in Odonata
barriers in
concerns
the
large
and Corduliidae; studies
Macromiidae and
Synthemist-
260
K.J.
Observations
idae.
these groups
ly has
to
the entire
mating
due to
tandem is achieved.
these
general,
hyaline
uli
and
is
a
belong
made.
once
tandem pairs
involving
(BICK & BICK
groups in
to
which
the
1981).
wings
the species,
RIBs
might
and
in
these
much
be
are
stimonce
aeshnids,
indicating that tactile
the effective
macromiids
barriers.
Male
uni-
remarkably
are
in libellulid genera, and discover-
as
is
groups
number of
large
a
shaped,
uniquely
are
cordulegasterids
form amongst
in
appendages
corduliids
of
appendages
the
of
prior
unreliable, and that isolation is effected
or
Male
mechanical differences
and/or
number
sexes
from the site
courtship behavior is unknown, evidence that visual
gomphids and
ing
However,
in
species
nonexistent. Rare-
within these groups
flight, and disappearance
species
nonexistent
are
contact
are
for any species
sequence
individuals have been collected
heterospecific
In
closely-related
data
the unknown whereabouts of the
their rapid
meeting,
of
behavor
mating
on
scanty, and experimental
are
been observed,
Tennessen
especially
Much
challenging.
more
study of Libellulidae is needed.
Very
ferences
exist
coveries
await
in
the
well
as
world.
Acanthagrion,
is
knowledge
undertake studies of
large
and
Argia
available.
Epallagidae of
in
genera
Ischnura
neo-
color
dif-
and
dis-
Polythoridae,
Platystictidae and Protoneu-
and
Chlorocyphidae
are
and
Intriguing problems
Pseudostigmatidae,
the
There
species.
who
investigators
as
behavior and isolation of
on
which many unusual structural
between related
Megapodagrionidae,
ridae,
published
little has been
tropical Zygoptera,
for
other parts of
Coenagrionidae,
the
which
little
very
such
as
behavioral
Much remains to be learned in the Caloptery"
gidae.
'
The terms used by
phological
be
structures
directed
minimize
clarify
and
in
not uniform
terminology
difficulties.
importance of each
and
mor-
and
confusion
needed
document
to
T.
Takeuchi
on
are
being
groups.
Fe-
possible barriers
contact
published work
studies
although
two
are
possibility of
deserves attention. No
appeared,
Obana and Mr.
prevent
is
barrier in certain
sperm removal
further study. The
has
will
that
Research
and
1972) and effort should
(BICK
difference between tactile and mechanical isolation,
polymorphism
Zygoptera
riers
the
defining the
quiring
a
interpretive
and
male
toward
are
'
for behavioral acts
various workers
re-
in
chemoreception
on
bar-
postmating
conducted by
Dr.
Sympetrum in Japan (K. Inoue,
S.
pers.
comm.).
The
exact
male accessory
of
genitalia
and
need of clarification. There
ture
of these
the
functions
parts
within
different
how
is
a
the
they
are
structures
used
large degree
order,
and
in
comprising
mating
are
the
still in
of variation in the struc-
different functional roles
isolation
Reproductive
for
structures
homologous
JOHNSON
of the
understanding
mechanical
cover
reconstruct
tions
and
(1972),
in
exist.
may
WAAGE
Studies
(1979a,
sperm-transfer
261
Odonata
is
rarely possible,
and
as
rarely
un-
from having
arise
relationships of male and female parts,
during copulation
(1971),
furthered
have
Such research may
process.
barriers. Difficulties
isolating
by PFAU
1982)
to
direct observa-
do
in
pairs stay
the wheel position after being collected.
PROBLEM-SOLVING GUIDELINES
Temporal
and
under
study.
and
useful.
dicates
through time,
between
Any overlap
either
initially
in the group
maturation
periods,
considered. Marking tech-
described by
as
for heterospecific
potential
a
changes
movement must be
such
analyses,
particularly
of isolation afforded by
Population
mating frequencies and
niques
isolation should be investigated
ecological
to determine the extent
GARRISON (1978),
in time
species
or
are
space in-
males and females to meet, and
other barriers must be sought.
In
such
ethologically-isolated
questions
ponse
in
males?
of the different
fishing-line
PAULSON
do males
visually
species? Models
technique
there is
model,
on a
nothing
in
favor
tandem by
males
and/or
dead
sented
achieve
dence
and
sult
their
a
from
&
with
copulation
males
models
in
tandem
be
the
be
observed. If
be
be
be
females
into
and
taken
though
even
perform
males
are
pre-
able
experimental
of the male
with
difficult
to
to
evi-
appendages
fe-
conspecific
control.
Evi-
(1) separation of heterospeachieved
(this
could
also
re-
however); (2) separation
of refusal motions by the female; (3)
without sperm transfer.
can
broken; (2) when
attempts
lat-
problematical.
Further
shape
may
1958;
the
Evi-
species,
type of tactile requirement,
lack
can
cooperate
them.
can
With
in tandem, the females
tandem
orven-
male’s attempting
a
(1)
tandem is
hold
techniques
secure
can
of different
altering
success
not
using the
1952),
1975).
res-
females
mechanical barrier
includes:
fly
answer
KRIEGER-LOIBL
or
two
do
and
securely
such
a
MOORE
WAAGE
mechanical barrier would be:
some
pair
isolation
can
tandem
a
to males
presented
1934;
tactile
a
motions and
obtained by
pairs before
of the
in
be
and
although
dence of
cific
female
observing
males,
be
interfere with
to
latter
the
them
clasp
refusal
tandem
can
tactile
heterospecific
able to
are
escape
of
models
details of tandem behavior
so
differentiating
presumed;
of
use
discriminate between
(KRIEGER
visual isolation is found lacking,
dence
can
1974; TENNESSEN 1975;
technique,
to land
(2)
the
species,
aspects of the female elicit
what
technique (ST. QUENTIN
trally-supported
ter
(1)
as
a
break
262
K.J.
If
interspecific
the possibility
that
vious conspecific
artificially
or
using
species
may be
every
Hybrids
fertilized
were
with
used
attempt should be
result, although
may
with sperm
from
in
seen
a
pre-
Therefore, attempts
from other species
sperm
previously
tandem. The techniques
.
not be ruled out.
can
eggs
observed,
larvae
rear
the eggs
mating
fertilize
especially
is
copulation
made to obtain eggs and
Tennessen
to
needed,
are
copulation
interspecific
by FREMLING (1967) for mayflies
applicable.
CONCLUDING REMARKS
Visual
(1)
in
important
needed
stimuli
color
distinctive
in
in
the
or
isolating
uniformly-colored
of
understanding
mechanical
morphological
of
range
character displacement
of RIBs (WAAGE
species;
and
groups;
more
in
efforts
serve
to
as
a
work
of which
some
vary
research should
the importance
focus
on
are
within
possible
of selection in the
origin
1979b).
be used
difficult taxonomic problems; such application
solve
is
Odonata.
amongst populations
Knowledge from reproductive isolation studies should
(3)
with
species
isolation
characteristics,
isolating species, often
geographic
barriers
patterns, whereas tactile differences may be equally
Behavioral and
(2)
be important
clear-winged
the
important
may
test of conventional
in
could
taxonomy.
ACKNOWLEDGEMENT
I thank
JONATHAN K. WAAGE for
many
valuable comments
on
the manuscript.
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