Chrysoperla Mohave (Banks) Neuroptera: Chrysopidae: Two Familiar Species in an Unexpected Disguise

Abstract

The vibrational courtship song of Chrysoperla mohave (Banks)
from several widely separated sites in California is described and
compared to C. downesi (Smith) and to a C. mohave-like population
of the recently described species. C. johnsoni Henry, Wells,
and Pupedis. Unexpectedly, C. mohave is shown to have a song
very much like that of C. downesi, in contrast to the very different
song of mohave-like C. johnsoni. Yet the C. downesi and C. johnsoni versions of "C. mohave" are physically indistinguishable.
Therefore, C. mohave is not a monophyletic taxon, but instead it is
a color morph that has arisen independently at least twice in separate lineages of Chrysoperla. Both C. downesi and C. johnsoni are thus highly variable species, encompassing populations that range
from solid dark green or bright green with no markings to light
yellowish-green with tergal spottig and dark crossveins. It is
argued that C. downesi should be taken from synonymy with C.
carnea (Stephens) and recognized as a valid biological species or
species complex, defined by a unique type of courtship song.

Full text

CHRYSOPERLA
MOHAVE
(BANKS)
(NEUROPTERA:
CHRYSOPIDAE):
TWO
FAMILIAR
SPECIES
1N
AN
UNEXPECTED
DISGUISE
BY
CHARLES
S.
HENRY
Department
of
Ecology
&
Evolutionary
Biology
University
of
Connecticut
Storrs,
Connecticut
06269
ABSTRACT
The
vibrational
courtship
song
of
Chrysoperla
mohave
(Banks)
from
several
widely
separated
sites
in
California
is
described
and
compared
to
C.
downesi
(Smith)
and
to
a
C.
mohave-like
popula-
tion
of
the
recently
described
species,
C.
johnsoni
Henry,
Wells,
and
Pupedis.
Unexpectedly,
C.
mohave
is
shown
to
have
a
song
very
much
like
that
of
C.
downesi,
in
contrast
to
the
very
different
song
of
mohave-like
C.
johnsoni.
Yet
the
C.
downesi
and
C.
john-
soni
versions
of
"C.
mohave"
are
physically
indistinguishable.
Therefore,
C.
mohave
is
not
a
monophyletic
taxon,
but
instead
it
is
a
color
morph
that
has
arisen
independently
at
least
twice
in
sepa-
rate
lineages
of
Chrysoperla.
Both
C.
downesi
and
C.
johnsoni
are
thus
highly
variable
species,
enconpassing
populations
that
range
from
solid
dark
green
or
bright
green
with
no
markings
to
light
yellowish-green
with
tergal
spotting
and
dark
crossveins.
It
is
argued
that
C.
downesi
should
be
taken
from
synonymy
with
C.
carnea
(Stephens)
and
recognized
as
a
valid
biological
species
or
species
complex,
defined
by
a
unique
type
of
courtship
song.
Key
Words:
systematics,
song,
sibling
species,
courtship,
reproductive
isolation
My
interest
in
green
lacewings
began
in
the
fall
of
1968.
The
preceding
year,
as
a
Junior
in
college,
I
had
enrolled
in
an
exciting
undergraduate
course
on
the
biology
of
insects,
taught
by
Professor
Frank
M.
Carpenter.
Carpenter
had
been
approachable
and
encour-
aging,
so
I
asked
him
to
support
an
undergraduate
honors
thesis
Manuscript
received
21
August
1992
291

292
Psyche
[Vol.
99
project
in
biology.
He
agreed
to
serve
as
my
advisor,
and
sug-
gested
that
I
talk
to
his
graduate
student,
Lee
Miller,
for
some
additional
ideas.
Miller
was
studying
the
structure
and
physiology
of
the
unique
ultrasound
detector
in
the
forewing
of
the
common
green
lacewing,
known
then
as
Chrysopa
carnea
Stephens
(Miller
1970).
He
proposed
a
trade:
if
I
took
over
the
care
of
his
lacewing
colonies,
he
would
let
me
use
his
lacewing
stocks
and
some
of
his
laboratory
space
and
equipment
for
my
project.
Before
long,
I
was
deeply
involved
in
a
study
of
courtship
and
mating
behavior
in
this
insect.
By
the
end
of
the
academic
year,
my
undergraduate
project
was
completed.
It
included
the
first
description
of
the
vibrational
courtship
songs
that
are
now
known
to
be
typical
of
the
lacewing
genus
Chrysoperla.
Although
years
would
pass
before
I
returned
to
acoustical
communication
in
insects,
this
study
set
the
stage
for
all
of
my
later
research
on
songs,
systematics
and
speciation.
In
those
days,
the
taxonomy
of
the
common
North
American
chrysopids
seemed
relatively
straightforward.
Chrysoperla
(for-
merly
Chrysopa)
carnea
was
considered
to
be
a
widespread,
hol-
arctic
species:
Tjeder
had
recently
synonymized
nearctic
C.
plorabunda
(Fitch)
with
palearctic
C.
carnea,
unable
to
find
con-
sistent
anatomical
differences
between
the
two
(Tjeder
1960).
Thereafter,
it
was
assumed
that
the
results
of
studies
on
American
populations
would
apply
equally
to
European
or
Asian
populations
mand
vice
versa.
Thus,
findings
within
local
populations
of
C.
carnea
sensu
lato
were
broadly
generalized
to
the
entire
species
in
investigations
of
photoperiod
and
diapause
(Honek
and
Hodek
1973,
Sheldon
and
MacLeod
1974;
Tauber
and
Tauber
1969,
1970),
dispersal
flight
(Duelli
1980,
1981),
defensive
responses
to
bat
sonar
(Miller
1975),
and
resistance
to
common
insecticides
(Bartlett
1964,
Plapp
and
Bull
1978).
In
addition,
lacewings
identi-
fied
as
C.
carnea
were
shipped
freely
all
over
the
world
in
com-
mercial
and
government-sponsored
programs
of
biological
control
(Jones
and
Ridgway
1976,
Ridgway
and
Jones
1969,
Tulisalo
and
Korpela
1973,
Tulisalo
and
Tuovinen
1975,
Zeleny
1965).
How-
ever,
those
most
familiar
with
the
species
were
acutely
aware
that
this
unified
view
of
C.
carnea
was
a
reflection
more
of
ignorance
than
of
cogent
empirical
data
(Adams
1983,
Bickley
and
MacLeod
1956,
Bram
and
Bickley
1963;
Adams,
pers.
comm.).
Recognition
of
taxonomic
diversity
within
C.
carnea
of
North
America
(=C.
plorabunda
s.
lat.)
actually
began
years
earlier,
with

1992]
Henry
293
the
description
of
several
species
from
local
geographical
areas
that
were
later
synonymized
with
or
included
as
varieties
of
C.
plorabunda.
These
have
included
C.
robertsonii
(Fitch),
C.
califor-
nica
(Coquillet),
C.
vegata
(Naves),
C.
downesi
(Smith),
and
C.
mohave
(Banks),
among
others
(Bickley
and
MacLeod
1956).
Each
was
thought
to
be
distinguishable
from
"true
C.
plorabunda"
(or
C.
carnea)
by
different
shades
of
green
or
by
unique
markings
on
the
body
and
wings.
Of
these
taxa,
only
C.
downesi
and
C.
mohave
are
commonly
recognized
and
discussed
today,
although
neither
has
official
species
status
anymore
(see
below).
Unfortunately,
colors
fade
badly
in
lacewings
no
matter
how
they
are
preserved,
so
museum
holotypes
and
paratypes
have
been
of
little
use
in
resolv-
ing
taxonomic
questions
that
concern
the
closely
related
taxa
of
the
Chrysoperla
carnea/plorabunda/downesi
complex.
Even
in
liv-
ing
specimens,
ground
color
and
markings,
setation,
body
size,
wing
shape,
and
venation
have
been
shown
to
vary
markedly
within
single
presumed
species
across
relatively
small
geographi-
cal
areas
(Tauber
and
Tauber
1986a;
Tauber
and
Tauber
1975,
1981).
Increasing
awareness
of
songs
produced
by
Chrysoperla
green
lacewings
has
completely
changed
how
we
perceive
species
boundaries
within
the
genus.
These
songs
are
vibrational
tremula-
tion
signals,
produced
by
rapid
vertical
oscillations
of
the
insect’s
abdomen,
and
are
transmitted
between
individuals
through
compli-
ant
substrates
such
as
leaves,
grass
blades,
or
conifer
needles
(Henry
1979,
1980b,
1983a).
No
drumming
is
involved.
Both
sexes
sing,
and
copulation
depends
upon
a
male
and
a
female
success-
fully
establishing
a
duet
in
which
each
partner
alternately
produces
the
same
song
phrase.
Because
the
songs
of
each
partner
must
exactly
match
for
this
to
happen,
populations
which
differ
acousti-
cally
from
one
another
will
be
unable
to
interbreed
and
should
be
considered
separate
biological
species.
By
this
criterion,
C.
carnea
is
not
holarctic
and
monolithic,
but
instead
consists
of
distinctive
complexes
of
several
species
each,
in
both
Europe
and
North
America
(Henry
1983a).
The
C.
carnea
complex
of
Europe
remains
to
be
analyzed
(Henry
1985b),
but
extensive
song
analysis
and
behavioral
tests
of
North
America’s
equivalent
C.
plorabunda
complex
have
supported
the
recognition
of
three
sibling,
cryptic
species
within
it:
C.
plorabunda,
C.
adamsi
Henry,
Wells,
and
Pupedis,
and
C.
johnsoni
Henry,
Wells,
and
Pupedis
(Henry
1993,

294
Psyche
IVol.
99
Henry,
Wells,
and
Pupedis
1993,
Wells
and
Henry
1992).
These
newly
defined
species
are
not
clearly
comparable
to
any
of
the
species,
subspecies,
races
or
varieties
of
nearctic
Chrysoperla
pre-
viously
described
in
the
literature.
A
very
different
type
of
song
characterizes
C.
downesi,
strongly
supporting
its
taxonomic
valid-
ity
and
arguing
against
Garland’s
decision
to
synonymize
it
with
C.
carnea/plorabunda
(Garland
1985).
In
fact,
variation
on
the
C.
downesi
song
theme
in
different
parts
of
North
America
suggests
that
this
taxon
may
itself
turn
out
to
be
a
complex
of
cryptic
bio-
logical
species
(Henry
1985a).
C.
mohave
is
a
special
case.
It
appears
to
be
closely
allied
to
the
three
described
species
of
the
C.
plorabunda
complex,
but
it
is
uniquely
characterized
by
numerous
darkly
pigmented
crossveins
and
gradate
veins
in
the
wings
(Banks
1938,
Tauber
and
Tauber
1973).
Also,
the
body
is
usually
light
yellowish-green,
with
more
or
less
numerous
brownish
or
reddish-brown
spots
on
the
thoracic
and
abdominal
terga
(Henry
1993).
This
suite
of
features
is
unknown
in
the
C.
plorabunda
or
C.
downesi
complex
outside
of
the
American
southwest.
Other
traits
that
have
been
associated
with
some
populations
of
C.
mohave
include
semi-predaceous
adult
food
habits
and
prey-mediated
control
of
reproductive
dia-
pause
(Tauber
and
Tauber
1981).
Although
it
was
finally
reduced
to
the
rank
of
a
variety
or
ecotype
of
C.
plorabunda
(Tauber
and
Tauber
1973),
it
has
enough
distinctive
traits
to
suggest
that
it
might
constitute
a
cohesive
genetic
unit.
Certainly
C.
mohave
is
a
candidate
for
careful
analysis
of
courtship
songs
across
its
known
geographical
range.
The
present
study
examines
the
songs
of
Co
mohave
collected
from
widely
separated
parts
of
California
and
identified
as
that
species
(or
ecotype)
by
Phillip
Adams,
California
State
University,
Fullerton.
Results
are
compared
to
song
data
collected
earlier
on
a
special
population
of
mohave-like
insects
from
the
San
Francisco
Bay
area
(Henry
1993),
and
to
the
songs
of
C.
downesi,
C.
plorabunda,
C.
adamsi
and
C.
johnsoni.
The
systematic
status
of
C.
mohave
is
then
discussed
and
assessed,
based
on
these
comparisons.

1992]
Henry
295
MATERIALS
AND
METHODS
Collection
and
Rearing-
Living
adults
of
C.
mohave
were
col-
lected
20-28
September,
1987,
from
three
relatively
hot,
dry
sites
in
California,
covering
a
north-south
range
of
nearly
1000
km
(Fig.
1).
Those
referred
to
as
"Garberville"
were
found
on
the
evening
of
20
September
in
mixed
stands
of
young
Douglasfir
and
scrub
live
oak
along
a
side
street
of
Garberville,
California,
elevation
-280
m,
about
22
km
southeast
of
the
Humboldt
Redwoods.
"Bartlett"
specimens
were
obtained
21
September
near
the
transi-
tion
from
desert
to
forest,
elevation
1000
m,
in
an
open
area
of
Springs
Canyon
(Clare o
Forest
Figure
1.
Collecting
sites
in
California
of
the
"C.
mohave"
yellow-green
color
variant
of
Chrysoperla
johnsoni
and
C.
downesi.
Claremont,
marked
with
a
triangle,
is
the
type
locale
of
C.
mohave
(Banks).

296
Psyche
[Vol.
99
mixed
Douglasfir
and
pine,
along
a
dirt
road
from
Route
20
to
Bartlett
Springs,
California,
ten
km
northeast
of
Clear
Lake.
"For-
est
Home"
insects
were
taken
26-28
September
in
an
open,
diverse
forest
at
the
Forest
Home
Campground,
California,
elevation
1650
m,
in
the
San
Bernardino
Mountains.
Collecting
data
for
other
mohave-like
lacewings,
obtained
in
Strawberry
Canyon,
Berkeley,
California
and
assigned
to
C.
johnsoni,
can
be
found
in
another
paper
(Henry
1993).
Adults
of
C.
downesi
were
procured
in
September
and
October,
1977-1983,
from
a
second-growth
forest
of
red
spruce
and
from
a
young,
open
stand
of
red
pine
at
the
Huyck
Nature
Preserve,
Rens-
selaerville,
New
York,
elevation
650
m,
about
32
km
southwest
of
Albany.
Other
specimens
identified
as
C.
downesi
were
col-
lected
in
1987
at
various
locations
in
western
North
America,
including
the
Bartlett
and
Forest
Home
sites
described
above.
All
insects
were
returned
to
Storrs,
Connecticut
for
mainte-
nance,
rearing
and
song
analysis.
Sexes
were
kept
apart
and
sup-
plied
with
water
and
a
WheastTM-based
diet.
In
the
laboratory,
long
day
photoperiods
(17"7
hr
L:D)
terminated
reproductive
diapause
in
field-collected
individuals
and
brought
on
sexual
receptivity
and
spontaneous
singing
after
1-6
weeks.
Progeny
of
selected
mated
pairs
were
raised
under
short
day
conditions
(10:14
hr
L:D)
using
established
methods
(Henry
1979,
1983b,
1991).
Upon
eclosion,
all
adults
were
maintained
under
a
long
day
photoperiod
regime.
This
rearing
protocol
was
designed
to
satisfy
any
obligatory
dia-
pause
requirements
of
the
insects
(Nechols,
Tauber,
and
Tauber
1987,
Tauber
and
Tauber
1973b)
and
to
facilitate
rapid
acquisition
of
sexual
receptivity.
Analysis
ot"
Songs-
At
least
five
complete
courtship
songs
(short-
est
repeated
units
or
SRUs)
of
each
presumed
member
of
C.
mohave
from
each
locality
were
recorded
on
cassette
tape
and
then
digitized
and
analyzed
with
MS-DOS-based
computer
hardware
and
software,
using
methods
described
in
previous
papers
(Henry
1979,
1980b,
1990;
Henry
and
Johnson
1989,
Henry
and
Wells
1990).
Males
and
females
were
induced
to
sing
by
playing
back
previously
recorded
or
artificially
synthesized
songs
through
a
loudspeaker
(Henry
1989).
A
similar
protocol
was
applied
to
at
least
five
SRUs
of
each
individual
of
C.
downesi.
Comparisons
to
other
species
and
populations
used
the
results
of
analyses
per-
formed
previously
and
tabulated
in
earlier
papers.

1992]
Henry
297
The
songs
of
Chrysoperla
green
lacewings
consist
of
frequency-
modulated
volleys
of
abdominal
vibration
repeated
at
regular
inter-
vals.
Some
taxa,
such
as
C.
plorabunda
and
C.
adamsi,
are
characterized
by
relatively
simple
songs,
composed
of
single-vol-
ley
SRUs
repeated
many
times
(Henry,
Wells,
and
Pupedis
1993).
Other
species,
such
as
C.
downesi
and
C.
johnsoni,
produce
more
complex
songs
that
consist
of
much
longer,
multisyllabic
(multi-
volley)
SRUs
and
perhaps
even
two
or
more
distinctly
different
types
of
volleys
(Henry
1980a,
Henry,
Wells,
and
Pupedis
1993).
The
populations
of
C.
mohave
analyzed
here
have
complex
songs,
very
similar
to
those
of
C.
downesi;
I
defined
and
measured
ten
song
features,
chosen
for
maximum
compatibility
with
the
song
measurements
published
for
other
species
(Table
1,
Fig.
2).
The
following
measurements
were
taken
for
each
of
the
two
types
of
volleys,
long
and
short,
found
in
these
songs:
(1)
duration
of
each
volley,
(2)
interval
between
the
start
of
one
volley
and
the
start
of
the
next,
(3)
frequency
(pitch)
of
each
volley,
and
(4)
number
of
volleys
per
SRU.
In
addition,
it
was
necessary
to
measure
fre-
quency
(3,
above)
separately
for
beginning
versus
terminal
short
volleys
in
each
SRU.
Total
length
of
the
SRU
(5)
was
also
tabulated.
In
total,
comprehensive
analyses
were
performed
on
the
songs
of
18
C.
mohave
individuals
from
California
(13
from
Garberville,
3
from
Bartlett,
and
2
from
Forest
Home).
The
sex
ratio
(male:
female)
of
the
total
sample
was
0.44.
For
C.
downesi
from
New
York
State,
the
songs
of
19
individuals
were
analyzed
(sex
ratio
0.47).
Comparisons
were
made
to
31
additional
mohave-like
indi-
viduals
of
C.
johnsoni
from
Strawberry
Canyon,
described
in
another
paper
(Henry
1993).
Temperature
strongly
influences
song
characteristics,
and
data
must
be
treated
in
some
way
to
compensate
for
this
if
they
are
col-
lected
at
different
temperatures.
Fortunately,
all
measurements
of
C.
mohave
were
taken
within
half
a
degree
of
25C.
However,
data
for
C.
downesi
were
obtained
over
several
years,
at
temperatures
ranging
from
19-31
C.
Therefore,
for
that
species,
all
individual
means
at
different
temperatures
were
transformed
(normalized)
to
a
standard
temperature
(25C).
This
was
done
by
regressing
tem-
perature
against
each
song
feature.
When
the
temperature
effect
was
significant
(P<0.05),
as
it
was
for
most
features,
the
slopes
of
the
resulting
regression
equations
were
used
to
modify
the
original

298
Psyche
[Vol.
99
Table
1.
Means
and
standard
errors
of
song
features
of
Chrysoperla
downesi
(eastern
U.S.A.)
and
mohave-like
C.
downesi
(California).
Unless
otherwise
noted,
time
is
in
milliseconds
and
frequency
is
in
Hertz.
Also
shown
are
the
results
of
a
T-
test
to
determine
significant
differences
between
the
populations.
C.
downesi
C.
"mohave"
(N=19) (N=18)
T-Value
P-Value
Duration,
long
volleys
311.66
485.47
9.86
0.000
+14.91
+8.96
Interval,
long
volleys
633.50
675.63
2.19
0.035
+10.23
+16.52
Duration,
short
volleys
126.35 155.45
4.13
0.000
+5.34
+4.32
Interval,
short
volleys
290.88
241.88
-5.20
0.000
+_5.81
+_7.50
Frequency,
long
volleys
82.50
80.02
1.44
0.163
+_2.02
+_0.66
Frequency,
1st
short
volleys
Frequency,
last
short
volleys
74.41
73.56
-0.56
0.583
+_1.59
+_0.73
64.81
64.20
-0.44
0.664
+_1.27
+_0.78
Number
of
long
volleys
4.52
5.75
3.54
0.001
+_0.21
+_0.28
Number
of
short
volleys
Song
Duration
(total,
in
seconds)
13.36
23.90
6.52
0.000
_+0.99 +_1.26
6.23
9.66
5.91
0.000
_+0.28
+-0.52
values;
otherwise,
data
were
left
unchanged.
Calculations
of
basic
statistics
and
Student’s
T-tests
were
then
performed
on
the
new
data
set
of
temperature-compensated
and
raw
values.
When
appro-
priate,
tabulation
of
the
residuals
from
the
linear
regression
analy-
sis
was
used
to
predict
the
mean
and
standard
error
at
25C
for
each
song
feature
(Table
1).
I
used
the
PC
software
package
CSS:
Statistica
3.1
(StatSoft
Inc.,
Tulsa,
OK)
for
statistical
analyses.
Acquisition,
display,
tern-

1992]
Henry
299
0
2
3
4
5
6
7
8
9
10
11
12
Time
(seconds)
Figure
2.
Digitized
oscillographs
of
the
shortest
repeated
unit
(SRU)
of
the
vibra-
tional
song
of
mohave-like
C.
johnsoni
from
Strawberry
Canyon,
California,
dark
green
C.
downesi
from
New
York
State,
and
mohave-like
C.
downesi
from
Gar-
berville,
California.
All
were
recorded
at
25C.
poral
and
spectral
analysis,
and
artificial
synthesis
of
songs
were
executed
using
Waterfall
3.18
and
Spike2
4.0
(Cambridge
Elec-
tronic
Design,
Cambridge,
U.K.)
and
DADiSP
2.00B
(DSP
Devel-
opment
Corp.,
Cambridge,
MA).
The
graphs
and
other
figures
were
produced
in
SigmaPlot
5.0
(Jandel
Scientific,
Corte
Madera,
CA)
and
CorelDRAW
3.0
(Corel
Systems
Corp.,
Ottawa,
Canada).
All
programs
ran
under
MS-DOS
on
an
80286
computer.
Analysis
of
Morphology:
All
living,
field-collected
individuals
of
C.
rnohave
were
examined
under
a
dissecting
microscope,
to
iden-
tify
external
morphological
features
that
might
vary
among
geo-
graphical
populations
or
between
morphs.
The
genitalia
of
two
preserved
C.
mohave
males
from
Garberville
and
two
more
from

300
Psyche
[Vol.
99
Forest
Home
were
prepared
for
closer
study
by
clearing
the
last
few
segments
of
the
abdomen
in
hot
10%
KOH
for
15-30
min,
as
described
by
Bram
and
Bickley
(1963).
Sclerotized
regions
were
enhanced
by
staining
for
two
minutes
in
5%
Chlorazol
Black
E
aqueous
solution.
The
genitalic
structures
were
then
everted
using
a
small
pipette
and
placed
in
a
drop
of
glycerin
for
examination
under
a
compound
microscope.
These
preparations
were
compared
to
those
of
eastern
C.
downesi
and
mohave-like
C.
johnsoni
geni-
talia
made
earlier
(Henry,
Wells,
and
Pupedis
1993).
Voucher
specimens,
pinned
or
in
70%
ethanol,
have
been
deposited
in
the
personal
collection
of
Charles
S.
Henry
or
in
the
collection
of
the
Connecticut
State
Museum
of
Natural
History,
the
University
of
Connecticut,
Storrs.
RESULTS
Oscillographs
of
the
vibrational
songs
(SRUs)
of
a
mohave-like
C.
johnsoni
from
Strawberry
Canyon,
a
C.
downesi
from
New
York
State,
and
a
C.
mohave
from
Garberville,
California
are
shown
in
Fig.
2.
Each
exhibits
the
volley
duration,
interval,
and
amplitude
structure
typical
of
its
population
at
25C.
The
songs
of
C.
downesi
and
Garberville
C.
mohave
are
compared
further
in
Fig.
3,
which
shows
the
frequency
(pitch)
structure
of
the
songs
as
computer-generated
sonographs.
The
figures
establish
that
the
song
of
C.
mohave
from
sites
outside
of
Strawberry
Canyon
is
very
similar
to
that
of
classic
C.
downesi,
and
bears
no
resemblance
to
the
song
of
its
closest
look-alike,
C.
johnsoni
from
Strawberry
Canyon.
When
the
temperature-corrected
means
of
each
song
fea-
ture
were
compared
between
C.
mohave
and
C.
downesi,
the
strik-
ing
similarity
of
the
two
populations
was
confirmed.
Even
though
the
comparison
was
between
populations
on
opposite
sides
of
the
continent,
T-tests
revealed
only
minor
differences
(Table
1).
Basi-
cally,
C.
mohave
has
a
lengthier
song
than
C.
downesi,
distin-
guished
by
more
and
longer
volleys
of
both
kinds,
and
the
interval
between
its
short
volleys
is
less.
But
without
seeing
the
striking
color
differences
that
separate
them,
it
would
be
easy
to
consider
C.
mohave
and
C.
downesi
a
single,
homogeneous
species.
In
a
parallel
manner,
mohave-like
C.
johnsoni
from
the
Berkeley
area
is
acoustically
indistinguishable
from
other,
more
typically
pig-
mented
C.
johnsoni
from
the
Pacific
Northwest,
central
and
south-
ern
California,
and
southwestern
Arizona
(Henry
1993).
Its
song
is

1992]
Henry
301
140
.
oo
0
.-
60-
I.I.
20-
C.
downesi,
eastern
U.S.A.
.:
i:"i
"5-:.".-
::’::’:
i:::!:::
:::::
".-
.:::’:’:
!:.q:
:!r
i::i
iii"
i.:.::
:!"
:ii
140
-r-
100-
0
60-
20
"C.
mohave,"
Garberville,
California
Time
(seconds)
Figure
3.
Computer-generated
sonographs
of
the
SRU
of
typical
dark
green
C.
downesi
from
New
York
State
and
mohave-like
C.
downesi
from
Garberville,
Cali-
fornia,
at
25C.
The
sonograph
shows
frequency
(tonal
or
pitch)
changes
with
time,
for
the
duration
of
each
song.

302
Psyche
[Vol.
99
so
different
from
that
of
C.
mohave/downesi
that
it
is
not
clear
how
to
begin
any
comparison.
Two
other
sibling
species,
C.
plorabunda
(sensu
stricto)
and
C.
adamsi,
were
not
considered
further
in
this
study,
because
neither
exhibits
mohave-type
morphology.
Also,
they
are
even
less
similar
to
C.
mohave/downesi
in
the
structure
of
their
songs
than
is
C.
johnsoni
(Henry
1993).
Externally,
all
C.
mohave
featured
some
degree
of
yellowish
ground
color,
tergal
spotting,
and
darkening
of
crossveins.
So
also
did
all
individuals
of
mohave-like
C.
johnsoni
from
Strawberry
Canyon,
and
in
fact
the
latter
could
not
be
distinguished
from
the
former
using
external
morphology.
More
detailed
studies
of
the
male
genitalia
of
several
specimens
from
each
taxon
failed
to
reveal
any
uniquely
different
character
states
that
could
be
of
sys-
tematic
use.
DISCUSSION
The
results
clearly
show
that
C.
mohave,
collected
over
a
wide
geographical
range
in
California,
shares
its
song
with
C.
downesi
and
is
almost
certainly
a
member
of
that
species.
However,
earlier
results
just
as
emphatically
placed
similarly
defined
"C.
mohave"
in
the
new
species
C.
johnsoni,
again
based
on
song
structure
(Henry
1993).
Yet
there
is
little
resemblance
between
C.
downesi
and
C.
johnsoni
in
any
other
way:
not
only
are
their
songs
com-
pletely
dissimilar,
but
the
former
is
typically
a
much
deeper
green
color
than
the
latter.
Two
conclusions
emerge
from
this.
Neither
conclusion
is
new
or
original
at
first
glance,
but
each
demands
an
important
revision
in
the
way
we
should
approach
the
systematics,
ecology,
population
genetics,
and
general
biology
of
Chrysoperla
in
the
future.
First,
C.
mohave
is
not
a
valid
clade,
and
therefore
should
not
be
recognized,
even
as
a
variety
or
ecotype.
It
is
polyphyletic,
hav-
ing
arisen
in
two
independent
(though
very
closely
related)
lin-
eages.
Of
course,
C.
mohave
even
now
has
no
official
validity,
having
been
synonymized
with
C.
carnea
(=C.
plorabunda
s.
lat.)
some
years
ago
(Tauber
and
Tauber
1973).
However,
it
continues
to
figure
in
detailed
models
of
nutritional
and
photoperiodic
con-
trol
of
seasonal
diapause,
evolution
of
seasonal
adaptations
and
life
history
traits,
geographic
polymorphism
in
ecophysiological
responses,
and
sympatric
speciation
(Tauber
and
Tauber
1982,

1992]
Henry
303
1987,
1989;
Tauber
and
Tauber
1973a).
Without
sure
knowledge
of
the
species
affiliation
of
the
particular
"mohave"
being
investi-
gated,
even
the
most
careful,
elegant
experiments
will
generate
misleading
or
meaningless
results.
Thus,
much
of
the
Taubers’
work
on
Strawberry
Canyon
C.
mohave
probably
applies
to
C.
johnsoni,
in
the
C.
plorabunda
species
complex,
but
is
not
at
all
applicable
to
other
California
populations
of
C.
mohave
belonging
to
the
C.
downesi
complex.
Nor
is
it
certain
that
C.
mohave
col-
lected
at
any
given
site
is
the
same
entity
from
year
to
year
or
even
from
individual
to
individual:
Chrysoperla
spp.
are
renowned
for
their
dispersal
capabilities
(Duelli
1980),
and
several
different
species
are
often
sympatric
or
syntopic
at
western
North
American
sites
(Henry
1991).
Every
field-collected
individual
must
be
placed
in
its
correct
species,
based
on
courtship
song,
before
any
other
work
can
be
done.
The
second
conclusion
is
that
acoustically
defined
species
such
as
C.
johnsoni
and
C.
downesi
are
not
necessarily
homogeneous
in
their
external
appearance
or
biology.
Again,
the
polymorphic
nature
of
several
species
within
Chrysoperla
has
already
been
rec-
ognized
and
described,
but
that
was
when
many
different
forms
were
still
included
within
a
monolithic,
holarctic
C.
carnea
(Tauber
and Tauber
1986b)
and
C.
downesi
was
defined
by
color
alone
(Tauber
and
Tauber
1981).
It
is
now
known
that
C.
johnsoni,
formerly
considered
part
of
C.
plorabunda
s.
lat.,
encompasses
populations
whose
colors
vary
from
light
yellowish-green
to
bright
green,
some
of
which
remain
green
in
winter
diapause
and
others
of
which
turn
brown
while
hibernating
or
aestivating
(Henry
1993).
More
surprisingly,
the
present
study
shows
that
C.
downesi
is
also
extremely
variable
in
color.
It
is
no
longer
acceptable
to
assign
all
dark
green,
nearctic
Chrysoperla
with
carnea-type
male
genitalia
to
C.
downesi.
In
fact,
some
are
among
the
yellowest
pop-
ulations
yet
encountered
in
the
genus.
Throughout
this
paper,
I
have
assumed
that
C.
downesi
is
a
valid
species,
even
though
it
is
presently
in
synonymy
with
C.
carnea--and
therefore,
presumably,
with
some
unspecifiable
mem-
ber
of
the
C.
plorabunda
species
complex.
Support
for
its
validity
comes
from
the
unique
structure
of
its
courtship
song
(Henry
1980a),
which
is
so
different
from
the
songs
of
the
three
species
in
the
C.
plorabunda
complex.
The
C.
downesi
song
varies
region-
ally,
but
is
always
recognizable
in
populations
across
the
entire

304
Psyche
[Vol.
99
width
and
breadth
of
the
North
American
continent.
Based
on
what
we
know
of
the
songs
of
hybrids
produced
between
eastern
C.
downesi
and
C.
plorabunda
in
the
laboratory
(Henry
1985b),
C.
downesi
seems
not
to
hybridize
with
other
species
in
nature:
indi-
viduals
with
hybrid
songs
have
not
been
found
in
the
field,
in
spite
of
extensive
collecting.
Thus,
all
evidence
indicates
that
C.
downesi
is
a
monophyletic
clade,
and
that
it
should
have
its
species
status
reinstated.
But
as
in
C.
johnsoni,
distinctive
color
forms
like
"C.
mohave"
probably
do
not
respect
species
boundaries
within
the
C.
downesi
species
complex.
Instead,
single
biological
species
may
show
a
wide
range
of
color
patterns
and
life-history
traits
in
differ-
ent
regions,
perhaps
through
adaptation
to
local
conditions.
The
reasons
for
the
parallel
evolution
of
mohave-like
morphol-
ogy
in
at
least
two
lineages
of
lacewings
are
unknown.
One
can
speculate
that
these
morphs
are
restricted
to
dry
areas
with
highly
seasonal
precipitation
patterns,
and
that
the
yellow-green
ground
color
and
dark
crossveins
camouflage
the
insects
against
desic-
cated
vegetation.
Yet
one
of
the
driest
regions
collected,
the
Kofa
Mountains
of
southwestern
Arizona,
supports
a
bright
green
popu-
lation
of
C.
johnsoni
that
remains
green
even
during
the
winter
(Henry
1993).
Also,
mohave-type
individuals
of
C.
downesi
at
the
dry
Bartlett
Spring
site
coexist
with
typical
dark
green
C.
downesi,
which
seems
contrary
to
the
camouflage
hypotheses
(C.S.H.,
unpublished
data).
Clearly,
much
hard
work
remains
to
be
done
on
the
systematics
and
ecology
of
the
C.
downesi
complex.
Recogniz-
ing
the
true,
biologically-defined
species
boundaries
within
this
and
the
other
sibling
species
complexes
of
Chrysoperla
will
greatly
facilitate
our
understanding
of
all
aspects
of
their
biology
and
prevent
potentially
costly
scientific
mistakes.
ACKNOWLEDGEMENTS
This
study
was
supported
in
part
by
National
Science
Founda-
tion
Award
BSR-8508080
to
Charles
S.
Henry
and
by
the
Research
Foundation
of
the
University
of
Connecticut.
I
thank
the
following
for
help
in
collecting
and
maintaining
living
lacewings
from
North
America:
Marta
Martinez
Wells
and
Julie
J.
Henry
(University
of
Connecticut,
Storrs);
J.
B.
Johnson
(University
of
Idaho,
Moscow);
Phillip
A.
Adams
(California
State
University,
Fullerton);
Ray-
mond
Pupedis
(Yale
University);
and
Norman
D.
Penny

1992]
Henry
305
(California
Academy
of
Sciences,
San
Francisco).
I
dedicate
this
paper
to
my
mentor
and
friend,
Professor
Frank
M.
Carpenter,
who
has
managed
so
often
to
recognize
and
encourage
hidden
strengths
in
his
students
and
associates,
thereby
coaxing
from
them
their
very
best
work.
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