The rotundifolia Complex of the Genus Andiltorypha
(Orthoptera: Tettigoniidae): Songs Reveal New Species
T. J. WALKER, T. G. FORREST," AND J. D. SPOONER3
Ann. Entomol. Soc. Am. 96(4): 433 447 (2003)
ABSTRACT Since 1934, the rotundifolia complex of Amblycorypha has consisted of two named
species: Amblycorypha par ; ..... I ei occurring from the Mississippi River westward into the eastern
edges of the Great Plains, and Amblycorypha '. I ,:. occurring in most of the eastern United
States. The latter entity is here shown to consist of at least three species with distinctive .ii, .- songs
and different but overlapping geographical distributions. At25C, A. ....... :" (Scudder) produces
-26 calling song units (phonatomes) per s, whereas Amblycorypha bartrami n. sp. Walker, A.
parnipennis, and Amblycorypha alexander n. sp. Walker produce 10, 5, and 2.8 ph/s respectively.
Amblycorypha rotundifolia occurs from Illinois to New York and southward along the Appalachians
to northern Georgia. Amblycorypha bartrami occurs in the southeastern states, and A. alexander
broadly overlaps the distributions of the other two eastern species. Where A. alexander and A.
S are sympatric, the two occur in similar habitats; where A. alexander and A. bartrami are
sympatric, A. bartrami occurs in more xeric habitats. No morphological characters were found that
reliably ..1. .-i the three eastern species, yet no fewer than three species must be recognized to
provide names for populations that behave as distinct species where they co-occur.
KEY WORDS F'....... -...;I... katydids, calling songs, female phonoresponse, sibling species,
IN THEI 1914 REVISION of North American Amblycory-
pha, Rehn and Hebard divided the genus into two
groups. Group I contained species with the humeral
sinus of the lateral lobes of the pronotum i: im-
pressed, at least rectangulate' and having the "indi-
vidual metasternal lobes not transverse." Group II had
forms with the humeral sinus "less impressed (and not
rectangulate) or subobsolete" and having "individual
metasternal lobes transverse." In Group II, here called
the rotundifolia complex, they placed three forms that
they considered to be geographic races of a single
species, Amblycorypha rotundifolia (Scudder). In
1934, Hebard elevated one of the forms, A. r. parvi-
pennis (Stal) to species status. In 1960, R. D. Alexander
concluded that the species known as A. rotundifolia
consisted of two sibling species that were inseparable
.:.. .1I.. I but had distinctive calling songs. He
., .. II, named the species by their song types and
reported that "rattler" was a northern species extend-
ing southward to the southern border of the Appala-
chian mountains and that "clicker" was a southern
species extending northward into southern Ohio.
When T.J.W. began studying the songs of katydids >40
yr ago, he soon discovered that based on song, Florida
1 Department of Entomology and Nematology, University of Flor-
ida, Gainesville, FL 32611 (e-mail: ..1 :
2 Department of Biology, University of North Carolina at Asheville,
Asheville, NC 28804.
Department ofBiology and Geology, Universityof South Carolina
Aiken, Aiken, SC 29801
was home to two species of the rotundifolia complex.
One species seemed to be Alexander's "clicker"; the
other was a previously unrecognized species that he
dubbed "fast ticker." In the years II. -... he and
J.D.S. accumulated specimens, tape recordings of
songs, and field notes pertaining to populations of the
three presumptive species of A. rotundifolia and, to a
lesser extent, populations of Amblycorypha parvipen-
nis. More recently T.G.F. began studying rattler and
clicker populations near Asheville, NC. This paper
summarizes what we know about four species in the
A. rotundifolia complex and provides formal names for
two that have none.
The rotundifolia complex has at least four species, as
will become evident after their songs, ecology, and
geographical distributions are described. To facilitate
those descriptions, we will first attend to the require-
ments of formal nomenclature.
Amblycorypha rotundifolia (Scudder) 1862: 445
rattler round-winged katydid
(Figs. 1 and 5, 7-10)
Scudder's types of A. .'. .7.' .. are lost, but be-
cause they came from four New England states and
iI .. the name must be assigned to the species that
Alexander termed "rattler."
0013-8746/03!0433-0447$04.00/0 0 2003 Entomological Society of America
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
00 y= 158ffx- 13638
00 0 W = 06478
9 - - - -
18 20 22 24 26
Fig. 1. Phonatome rate as a function oftemperature in A i.:1,.. (24 tapes of 19 males from 10 counties in Georgia,
Illinois, Indiana, New Jersey, Ohio, Pennsylvania, Tennessee, and '. .. .. and A. paroipennis (nine tapes of five males). The
triangles are i. i T.J.W. from Carter Co., MO, and the squares are .i 1.... by J.D.S. from Bell Co., TX. The filled
circle is an A. rotundifolia recording from Dyer Co., TN, with atypical phonatomes. The dashed lines are the trend lines for
A. bartrami and A. alexainleri from Fig. 2.
Material Examined. 47 5, 16 9, Florida State Col-
lection of Arthropods, C .-... .I. FL (FSCA). Geor-
gia: Murray Co.: 2 j, 2 9, ll-VI-67, T.J.W.; Rabun Co.:
1 o, 1-VIII-68, J.D.S. Illinois: Douglas Co.: 8 5, 4 9,
25-VII-65, T.J.W. Indiana: Parke Co.: 4 6, 1 9, 25-VII-
65, T.J.W. New Jersey: Middlesex Co.: 1 o, 17-VIII-67,
T.J.W. New York: Schuyler Co.: 2 6. 12-VIII-65, T.J.W.
and G.K. Morris. North Carolina: Buncombe Co.: 3 e,
4 16-VIII-01, 3 d, 2 9,26-VII-01,3 2 ,4-VII-01,
T.G.F. Ohio: -I.. 11 Co.: 2 6, 12-VIII-67, 1 24-VIII-
67, T.J.W.; Washington Co.: 1 6, 21-VIII-68, 1 S, 21-
VIII-70, T.J.W. ... i .1... Berks Co.: 6 d. 1 9,
21 tu T.J.W.; Butler Co.: 1 S, 11-VIII-65, T.J.W.
Tennessee: Dyer Co.: 1 5, I lit ., T.J.W. Virginia:
Bland Co.: 1 31-VIII-68, T.J.W.; Westmoreland Co.:
2 5, 25-VIII-64, T.J.W. West Virginia: Greenbrier Co.:
4 j, 1-IX-72, TJ.W.
Amblycorypha parvipennis Stal 1876: 58
western round-winged katydid
(Figs. 1, 5, and 10)
A. *.i 1.- 7 parvipennis Rehln and Hebard 1914:
A. brachyptera Ball 1897: 237. [Ames & northwest-
A. iselyi Caudell 1904:50 [Wichita, KS]
Stal's type material was from Texas and is in the
Naturhistoriska Rijksmuseum, Stockholm, Sweden
(Otte 1997). When Hebard (1934) recognized A. par-
vipennis i. 'i 'II. distinctfrom A. rotundifolia, he
indicated that A. parvipennis brachyptera was a north-
ern race of A. parsipennis and that A. iselyi was a
synonym of A. brachyptera.
Otte (1997) placed A. iselyi as a synonym of the
southern race (A. p. parvipennis) rather than the
northern race (A. p. brachyplera). We do not recog-
nize subspecies in A. parvipennis because to do so
would imply that its geographical variation has dis-
continuities or steep lines and we know of neither.
Material Examined. 1 o, MO: Carter Co.: 23-VII-65,
TJ.W., FSCA. T.J.W. also examined specimens at the
University of Michigan Museum of Zoology, Ann Ar-
bor, MI (UMMZ) (29 localities). Academy of Natural
Sciences of Philadelphia, Philadelphia, PA (ANSP; 19
localities), and the National Museum of Natural His-
tory, Washington, D.C. (USNM) (five localities).
These localities plus records from the literature are
documented at http:// ii ., : ,1 .i ,/h00dbase.
htm and mapped in Fig. 10.
Amblycoryplha alexander n. sp., T. J. Walker
clicker round-winged katydid
(Figs. 2, 4, and 6 10)
Holotype. Male, Torreya State Park, 1 :. Co.,
FL, 16-VI-58, ravine forest, WTL-001-1, T.J.W., FSCA.
Green; possessing the characters that distinguish the
complex (Fig. 6A and B); ventral carina of hind femur
with five teeth; hind fernur reaching tip of tegmen.
Pronotal length, 5.9 mm, width, 4.2; tegminal length,
32, width, 9.4; hind femur length, 28; hindwing expo-
.\ii,,l: Female, FL Caverns State Park, Jackson
Co., FL, 14-VI-62, moist broad-leaved forest, T.J.W.
and J.D.S., FSCA. Like the rale but with four and five
teeth on carinae of left and right hind femora; hind
femora exceeding tips of tegmina by 2 mm. Pronotal
length, 5.8 mm, width, 3.9; tegminal length, 28, width,
8.3; hind femur length, 28; hindwing exposure, 3.5;
: ;i., length, 10.4.
_r. .i..'. 31 5, 10 9, FSCA. Florida: Jackson Co.:
6 same data as : : Li Co.: 2 same data
as holotype; 2 e, 17-VI-58, T.J.W., 1 13-VI-62, T.J.W.
Vol. 96, no. 4
WALKER ET AL.: Amblycorypha .' ':. '.
'- 10 -
y= 03447x +08886
y = 0.1693x 0.9026 y0.2508x -20265
R2 =0.3582 R20734
8 20 22
24 26 28 30
Fig. 2. Phonatome rate as a function of temperature in A. bartrami (16 tapes of 10 males, from four counties in Alabama,
Florida, and North Carolina) and A. alexainderi from northern localities (triangles) (17 tapes of 13 males from counties in
Georgia, North Carolina, and Ohio) and southern localities (circles) (13 tapes of seven males from Jackson and 1 .i ..
counties in Florida). The dashed line is the trend line for A. parvipennis from Fig. 1. The black square is a recording of A.
nr. bartrami from near Aiken, SC. The cluster of seven points that are nearly on the line for northern A. alexanderi between
23 and 24C are from Buncombe Co., NC. The two points at 25'
nearby McDowell Co., NC
Georgia: Murray Co.: 1 1 9, 11-VI-67, T.J.W.; Rabun
Co.: 1 20-VII-68, J.D.S. North Carolina: Buncombe
Co.: 4 4, 24-VII-01, 3 1 V 9, 30-VII-01, 1 29-VI-02,
3 S, 1 9, 30-VI-02, 2 S, 4 9, 16-VII-02, 1 S, 2 9,
20-VII-02, 1 6, 27-VII-02, T.G.F.; Macon Co.: 1 o,
20-VII-68, J.D.S. Ohio: Ashland Co.: 3 6, 7-VIII-65,
Other specimens. 6 5, FSCA Tennessee: Sevier
Co.: 6 1, 13-VIII-66, TJ.W.
tr:,,..., -. This species is named for R.D. Alex-
ander, who was first to recognize that it was distinct
from A. rntundifolia.
Amblycorypha bartrani n. sp., T. J. Walker
Bartram's round-winged katydid
(Figs. 2, 3, and 7-10)
Holotype. Male, western ( ..... i. Alachua Co.,
FL, 14-VI-63, interface of xeric hammock and dry old
field, Walker Tape Library (WTL) 005-6a,b, T.J.W.
and R. E. Love, FSCA. Green; possessing the charac-
ters that distinguish the complex; ventral carina of left
hind femur with four teeth, of right, six teeth; hind
femur slightly exceeding tip of tegmen. Pronotal
length, 5.9 mm, width, 4.0; terminal length, 30, width,
8.5: hind femur length, 28; hindwing exposure, 5.5.
Allotype. Female, 0.5 miles N of Alachua, Alachua
Co., FL, 8-VI-24, F.W. Walker, low bushes, mostly
dwarf chinquapin and oak, FSCA. Like the male but
with three and five teeth on carinae of left and right
hind femora: hind femora exceeding tips of tegmina by
2 mm. Pronotal length, 6.5 mm, width, 4.3; tegminal
length, 29, width, 9.0; hind femur length, 28; hindwing
exposure, 2.5; ovipositor length, 9.6.
P .i.. -..- 20 3, 4 9, FSCA. Alabama: Cleburne Co.:
1 i. 29-VIII-64, T.J.W.: Perry Co.: 3 d, 4 9, 9-VI-66.
J.D.S., 6 8, 8-VI-67, J.D.S. Florida: Alachua Co.: 1 S,
'C that nearly overlap those of southern A. alexander are from
29-VI-60, TJ.W., 1 3, 3-VIII-60, T.J.W., 1 3, 5-VI-62,
T.J.W.. 1 S, 3-VII-62, T.J.W., 1 10-VI-63, T.J.W. and
R.E. Love, 1 6, 2-VI-65, TJ.W. and R.E. Love; Leon
Co.: 1 6, l1-VI-62, T.J.W. and J.D.S.; liberty Co.: 1 o,
12-VI-62, T.JW. and J.D.S. North Carolina: Hoke Co.:
1 3,26-VII-64, T.J.W.; -.I ,..: Co.: 1 o, 21-VI-62, T.J.W.
Other specimens. 4 d, 4 9, FSCA. South Carolina:
Aiken Co.: 2 S, 17-' u ,.. 1 ,, 10-VII-87, 1 S, 7-VI-88,
1 j, 21-VI-88,1 9, 14-VI-93,1 9, 21-VI-93, I Edge-
field Co.: 1 9, 7-VI-88, JD.S.
Ft ,!,,I,,._. This species is named for William Bar-
tram, the earliest naturalist to explore the habitats in
Florida where T.J.W. first encountered A. bartrami.
Most fieldwork was at night. Most males were col-
lected by homing on their calling songs; females were
collected in habitats where males were calling by
shining light on foliage or by using a sweep net. Spec-
imens were often held alive with access to water and
fragments of dry dog food in screen cages or small jars
with screen tops. TJ.W. recorded songs full track on
0.25-inch tape at 15 ips with an Ampex 351, Nagra III,
or Nagra IV analog tape recorder and an American
D33 or ElectroVoice 655C dynamic microphone.
T.G.F. recorded songs using a Sennheiser ME66 shot-
gun microphone and a Tascam DAP-1 DAT recorder
at a sampling frequency of 48kHz. J.D.S. recorded
songs half-track on 0.25-inch tape at 7.5 ips with a Uher
Report S or Uher Report L analog tape recorder and
an Electrovoice 655C microphone. Most recordings
were of caged specimens, although a few were made
in the field with the aid ofa 61-cm diameter parabolic
reflector. T.J.W. and J.D.S. measured temperatures
with a mercury laboratory thermometer at the cage or
parabola; T.G.F. used a Tektronix ATP01 temperature
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
15s 24 C
200 ms 260C
*-- H--- ^--.(lm4-t-.-.
200 ms 240 C
200 ms 26 C
200 ms 250C
200 ms 240 C
Fig. 3. Waveforms of the songs of A. bartrami. (A) 15 s of, ii, mean phonatome rate during the long train is 9.1 ph/s
(WTL 005 4a, Leon Co., FL). (B E) Two consecutive phonatomes from prolonged trains of similar phonatomes. (B) Initial
component a single pulse (WTL 005 4d, Leon Co., FL). (C) Initial components two single pulses (WTL 005 4a, Leon Co.,
FL). (D) Initial component a brief pulse train (WTL 0056b, Alachua Co., FL). (E) Initial components two brief pulse trains
(WTL 005-5c, Stanly Co., NC). (F) Pair of phonatomes in a sequence of pairs (WTI, 005-14, Moore Co., NC).
probe. Initially recordings were analyzed with a Kay
Elemetrics model 7029A sound spectrograph. Re-
cently, most of the analog recordings were digitized at
a sample rate of 44,100/s and analyzed with CoolEdit
2000 (- ., 11 ..... Software) running on a Pentium II,
333 MhIz PC.
Male katydids make their. i:',.- songs by moving a
sharp, upturned edge (scraper) of thebase of the right
tegrnen along a series of downward projecting teeth
(file) at the base of the left tegmen. The set of sounds
made by one cycle of tegminal movement is called a
"phonatome." The surest way to learn what sounds are
made during a cycle is to capture the inovernent with
high-speed photography or video and relate the move-
ments to the sounds produced. This has been done
with Amblycortyphaparcipennis (Shaw etal. 1990). For
the other three species of the complex, we inferred
what sounds constituted a phonatome by looking for
repeating patterns in the waveform of the calling song
and comparing the results with what Shaw et al. had
Phonatome rates were calculated by T.J.W. from
times measured with a scale on sound spectrograms or
with a mouse on CoolEdit 2000 displays. For A. ro-
...,. ?. the rate was based on a 2-s sound spectro-
graph of the central portion of a prolonged train (i.e.,
sequence) of phonatomes. For the remaining species,
rates were based on a set of almost uniformly spaced
phonatomes in a prolonged train. For A. alexander and
A. parvipennis songs, measurements were made from
the beginning or end of the first phonatome in the
train to the corresponding point on the last phonat-
ome. For A. bartrami, in which phonatomes at the end
of long trains are suddenly more widely spaced, the
last phonatome measured was the one before the first
phonatome interval that was longer than the previous
one by >50%. In all cases, equal numbers of phonat-
omes and phonatome intervals were included in the
measurement. Linear regression was used to establish
trend lines for the graphs showing phonatome rate as
a function of temperature.
The phonatome rates calculated as described above
were the average rates in prolonged trains. In all spe-
cies,phonatomeratesusually 1 1... -.. 1., 1i1 during
prolonged trains. To quantify the decline, T.J.W. used
CoolEdit measurements to calculate the average pho-
Vol. 96, no. 4
WALKER ET AL.: Amblycorypha .' '..:" '.
natome rates for the first 10 and the last 10 phonatomes
in prolonged trains. The suddenly slowed phonatomes
at the end ofA. bartrami trains were again omitted. We
used the difference in the two average rates, expressed
as a percentage of the initial (faster) rate, to quantify
the degree that the rates declined.
Modal carrier frequencies of songs were deter-
mined with CoolEdit's FFT analysis. The sampling rate
was 44,100/s, and time-sample sizes were adjusted to
minimize variation from reading to reading.
Specimens from all available localities for the three
eastern species were measured in a search for dimen-
sional differences. T.J.W. used a calibrated ocular mi-
crometer of a Zeiss stereo microscope to measure
pronotal length, terminal width, hindwing exposure,
and ovipositor length. Pronotal length was measured
medially. Tegminal width was measured at the widest
point. Hindwing exposure was measured along the
longitudinal axis of the tegmen. Ovipositor length was
the straight-line distance from the apex to where the
ventral edge of the ovipositor disappeared at the sub-
genital plate. Tegminal and femoral lengths were mea-
sured with a dial micrometer.
Stridulatory files were studied after cutting the
stridulatory field from the left tegmen of males. The
inverted field was then magnified to 50 with a Leica
stereomicroscope and digitally photographed with a
Syncroscopy Auto-Montage system (http:/ www.
syncroscopy.com/syncroscopy). The number of file
teeth and length of the files were determined by
T.J.W. from hardcopy of 96X images. To be counted
as file teeth, structures had to be elongate with the
long axis perpendicular to the axis of the file and
evenly spaced or nearly so. File length was the
straight-line distance from the center of the first file
tooth to the center of the last file tooth.
The recordings in WTL will be archived and made
available on the Internet by the Macauly Library of
Natural Sounds at Cornell University. Excerpts of rep-
resentative recordings, including those of the A. ro-
* ." .. complex, are already accessible through
Insects of North America (http://buzz.ifas.
ufl.edu/). Also at this site are spreadsheets with the
data used to develop Figs. 1, 2, 7, and 8 and other
aspects of this paper (http:// i ..i i- ,. : I,
g001a.htm) and a database that substantiates the lo-
calities mapped in Fig. 10 along with detailed infor-
mation about the II .. ,_ sites of most specimens
examined for this study (http:i //I. .. .1,./
Results and Discussion
Male Songs. The calling song feature most useful in
identifying species in the A. i;,r. -.i, .. .1. ,;the
phonatome rate during near-uniform trains of phona-
tomes. Phonatome rates are temperature dependent
in all katydids (Walker 1975), although the differences
among the three eastern species of the A. rotundijolia
complex are so great that different field temperatures
do little to impede identifications by song (Figs. I and
A. rotundifolia has the fastest phonatome rate, av-
eraging -26 ph/s at 250C (Fig. 1). Calling males pro-
duce trains of phonatomes thatlastfrom <1 s to as long
as 7 s. Because the constituent phonatomes come too
rapidly to be heard individually, the human ear ren-
ders A. .. .... 7, trains as "rattles." The song often
begins with several brief trains that lead into a pro-
longed one. The prolonged train is often immediately
*! ii. iby a fewtrains of<10 phonatomes each (Fig.
5A). During trains lasting >1 s, the phonatome rate
decreases slightly. In laboratory recordings of seven
individuals, the phonatome rate near the end of long
trains was 4-14% slower (av. 10%) than the phonat-
ome rate near the beginning of the train. During pho-
natome trains the intensity of the phonatomes often
increases initially (e.g., first four trains in Fig. 5A).
A. alexander has the slowest phonatome rate, av-
eraging <4 ph/s at 25C. Phonatomes can be heard
individually and are rendered as "clicks" by the human
ear. Phonatomes are generally produced at nearly
uniform rates for 5-30 s. During such trains the pho-
natome rate usually declines (16 of 19 recordings),
with the average decline being 7% (range, 1 17%; 16
recordings of 12 individuals). These nearly regular
trains are .... ii preceded by several more widely
spaced clicks (Fig. 4A). During the prolonged trains
the intensity of successive clicks .. :1I '. .. .ini-
tially and then remains fairly constant or, near the end
of the sequence, decreases somewhat. The click rates
for A. alexander from Jackson :..1 i I.. ..: counties in
northern Florida are slower than the click rates for A.
alexanderi from northern Georgia, North Carolina,
and Ohio. The trend lines for phonatome rate as func-
tions of temperature predict a rate of 2.27 for songs
from two counties in Florida and 3.33 for songs from
Murray County, GA, and northward (Fig. 2). Near
Asheville, NC, T.G.F. recently discovered a popula-
tion of the A. rotundifolia complex that produced pho-
natomes at the rate of A. alexanderi but in trains of
8 14 phonatomes with intervals between trains similar
to the duration of the trains. Furthermore, calling
males often inserted ticks between phonatornes
within the trains. This caused TJW to reconsider his
identification, as A. alexander, of a 1966 recording and
six males from Greenbrier Cove, Sevier County, TN.
Until these populations are studied further they
should be excluded from what is here described as A.
A. bartrami produces -10 ph/s at 25C (Fig. 2)
less than half the rate of A. rotundifolia and more than
three times the rate of A. alexanderi. The usual phras-
ing of A. bartrami songs (Fig. 3A) resembles that of A.
i."r (Fig. 5A), but the phonatomes come
slowly enough to be heard as discrete units making the
calling males "fast tickers" rather than "rattlers." The
several more widely spaced sounds that may terminate
a prolonged phonatome train are single phonatomes
rather than trains. During prolonged trains, the pho-
natome rate decreases by 5 32% (average = 13%,
based on 16 recordings of 10 individuals). In his studies
of populations of the rotundifolia complex near Aiken,
SC, JDS originally considered them to be A. bat-trami
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
30 s 25 C
200 ms 25 C
.-.. t ....- 4lt--
200 ms 25 C
---- .. t ------- -t -
200 ms 21 C
2 s 25'C
2 s 230C
Fig. 4. Waveforms of the songs of A. alexander. (A) 30 s of calling; mean phonatome rate after initial seven phonatomes
is 2.1 ph/s (WTL OI1 8c, Jackson Co., FL). (B D) -. i. phonatomes from trains of similar phonatomes. (B and C) Terminal
pulse trains of these two phonatomes differ in length and complexity (WTI, 001 -8c, Jackson Co., FL). (D) Initial components
are a single pulse and two brief pulse trains (WTL 001-16, Murray Co., GA). (E) Sequence of phonatomes with an interpolated
phonatome (WTL 001 5b, Liberty Co., FL). (F) Sequence of phonatomes with two doubled phonatomes (WTL 001 12,
Ashland Co., OH).
because their phonatome rates were intermediate be-
tween A. rotundifolia and A. alexander and they oc-
curred in .. inii i. .i .. .: However, a recent analysis
of a recording of the calling song of a male from near
Aiken revealed a much slower phonatome rate than is
typical for bartrami i,-i 2, filled square). Therefore,
we do not list specimens of Aiken "A. nr. bartrmni" as
paratypes of A. bartrami even though A. bartrami is the
most appropriate available name for the Aiken pop-
ulations. T.J.W. recorded the songs of two A. bartrami
from Moore County, NC, but these are excluded from
Fig. 2 because neither recording contained trains of
more than four phonatomes and 66% of the phona-
tomes were in pairs (Fig. 3F). The phonatome rates
based on these pairs were 11.2 at 24.8C and 8.7 at
19.5C, which c( *.1 I ... *:- 1I to other rates for A.
bartrami and even better when they are adjusted up-
ward for being based on short (fewer than 20 phona-
tomes) rather than long phonatome trains. (When
rates based on short and long trains were compared for
12 of the 16 A. bartra i ... .:- '. 1 'Fig. 2, the
average adjustment factor was 1.18.)
A. parvipennis produces -5 ph!s at 250C, which is
approximately one-half the rate of A. batramni and
approaching twice the rate of A. alexanderi (Fig. 1).
Phrasing usually consists of trains of 20-40 phonat-
omes with inter-train intervals shorter than the trains
(Fig. 5D). As in other species, phonatome rate usually
declines toward the end ofatrain. For eight recordings
of five individuals the rates decreased 0 29% (aver-
age = 12%).
The :... ...... spectra of the calls of the four spe-
cies were broad, varied, and of no use in separating
species. There were usually several ii ...
frequency peaks, and the most intense frequency was
generally between 9 and 14 KHz. The mean most-
intense frequency for songs of each of the four species
. II between 9.8 and 11.1 kHz.
The phonatomes produced by males of the A. ro-
tundifolia complex are noteworthy for their complex-
ity and for their variation within and between species.
A typical phonatome of A. bartrami, A. alexander, and
A. paruipennis consists of one to four brief initial com-
ponents ii .1 by a longer terminal component
Vol. 96, no. 4
WALKER ET AL.: Amblycorypha .' ':. :'.
15s 25 C
200 ms 210 C
200 ms 20C
Fig. 5. Waveforms of the songs ofA. rotundifolia (A-C) and A. paripennis (D-E). (A) 15 s of.r r ..
phonatome rate during long train is 23 ph/s. (WTL 008-la, Berks Co., PA). (B) Four typical phonatomes (WTL 008-13. Parke
Co., IN). (C) Three atypical phonatomes (WTL 008-18, Dyer Co., TN). (D) 15 s ofA. parvipennis i-,. phonatome rate
during trains is 4.9 ph/s (WTL 013 2c, Carter Co., MO). (E and F) -... i phonatomes. (E) WTL 013 2b, Carter Co., MO.
(F) J.D.S. recording PArm2-la, Bell Co., TX.
(Figs. 3B-E, 4B-D, and 5E and F). Each initial com-
ponent has one to several pulses, whereas the terminal
component has 10 or more pulses. Phonatomes of A.
..... :?" ?.. typically lack a longer, terminal compo-
nent (Fig. 5B). In all species, the phonatomes of a
given train usually have a common pattern, as illus-
trated by four and three consecutive phonatomes from
A. .;.. i. !.' trains in Fig. 5B and C and two con-
secutive phonatomes from each of four A. barmrami
trains in Fig. 3B E.
In most of the A. bartrami songs, phonatomes began
with one or two intense exponentially decaying pulses
(initial components) -. :. ..i by the terminal pulse
train, in which the intensity of the pulses first in-
creased and then declined (Fig. 3B and C). As seen in
Fig. 3D, the initial pulse or pulses were sometimes
replaced with a brief pulse train. Males in the northern
half of the range of A. bartrami (Stanly and Moore
counties, NC, and Perry and Cleburne counties, AL)
often made phonatomes with two brief pulse trains
before the longer terminal train (Fig. 3E and F). Such
phonatomes were longer and were often associated
with lower average phonatome rates. In fact, five of
the six points that are- i: :ielow the A. bartrami trend
line in Fig. 2 represent northern localities. It should be
noted that the characteristic slowing of phonatome
rates within prolonged trains, in this and other species,
resulted from longer phonatome intervals rather than
Phonatomes in A. alexander began with 2 4 (usu-
ally 3) brief initial components. They ended with a
terminal pulse train that lasted more than twice as long
as the corresponding train in A. bartrami and included
complex variations in pulse rate and intensity (Figs.
4B-D). alexander phonatomes lasted approxi-
mately three times as long as A. bartrami phonatomes
at the same temperature. This accounted for most, but
not all, of the difference in phonatome rates between
the two species: the rest of the difference was a result
of longer intervals between phonatomes. Two atypical
15s 24 C
200 ms 23 C
200 ms 23
200 ms 230 C
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
s I i., .!. .: .!. F'ig. E and F, demonstrate that the
interval between phonatomes can be as brief as 40
msec, whereas the typical interval at 25C is more than
three times as great.
Phonatomes in A. parvipennis always ended with a
train of 10 or more pulses (Fig. 5E and F). Preceding
this were two or more isolated pulses or n. pulse
trains or a combination of the two.
In A. rotundifolia, the species with the highest pho-
natome rate and the briefest phonatomes, each pho-
natome was I ... .11 brief series of 2 5 (usually 3 or
4) intense pulses. There was no terminal train ex-
cept for the song of a male from western-most Ten-
nessee (Fig. 5C). In this song, each phonatome con-
sisted of a rapid group of 3-4 intense pulses followed
by a train of 10 or more less-intense pulses that had the
same graded changes in pulse intensity seen in the
terminal pulse trains in other species within the com-
plex. The extra train increased phonatome duration,
and lowered the phonatome rate of this recording
relative to the trend line for all recordings of this
species Ii 1, filled circle).
Detection of 1.1.. .. .. within "A. rotundifolia"
long predates this paper. Allard (1912, p. 462) noted
that "The stridulations of Amblycorypha rotundifolia
may consist. i ... :shuffling phrases, sh-sh-sh-sh,
repeated at intervals. At other times the notes become
more lisping and continuous, tsip-i-tsip-i-tsip-i-tsip."
The songs that Allard rendered onomatopoeically are
probably those of A. ,. ,;.. .'. and A. alexander,
respectively. Fulton (1932) described two songs for
North Carolina A. rotundifolia. One is clearly rattler,
"on bushes in mountains"; the other is evidently either
A. bartrami or A. alexander, probably the former,
because it was "on oaks in sand hills." Alexander
(1960) described and illustrated the songs of "clicker"
and "rattler" (his Fig. 16 and selection 5 on an accom-
panying 12-inch LP record.) As mentioned earlier, he
concluded that they were distinct species with over-
lapping geographical ranges. In the overlap area he
sometimes found them in mixed colonies, but he
showed that each was unaffected by the singing of the
other. When caged males of either species called, only
males of the same species were stimulated to join the
The song of A. parvipennis was first described by
Fulton (1928), who was also first to note that neigh-
boring males calling at the same time almost perfectly
synchronize the phonatomes of their overlapping
trains. (- :1i. and :- i, (1991, 1992, 1996) showed
that males that avoid such overlap at the beginnings of
their phrases have an advantage in the competition for
Exactly what movements of the file and scraper
produce the fine structure of the phonatomes are not
known. High-speed recordings of tegminal move-
ments during ..ii.. ..- .. .:., rypha nr. uheri (Walk-
er and Dew 1972) and in >20 species of barbitistine
phaneropterines (Heller 1990) show that the possi-
bilities are enormous. When Shaw et al. (1990) re-
ported that sounds of A. parvipennis equivalent to
those in Fig. 5E were made during a single cycle of
wing movement, they did not parse the sounds be-
tween the opening and closing portions of the cycle.
However, in other studies of phaneropterine tegminal
movements during ,ii, most or all of the sounds
have been made on closure. In any case, each pulse in
the phonatorne probably represents the engagement
and release of a single file tooth, with the amount of
energy stored before release determining the intensity
of the pulse.
Female Answers. Unlike most other katydids, fe-
males of most species in the .,1 !.,,i Phaneropteri-
nae make brief, nondescript answers (ticks) to songs,
or to certain songs, of ... :.. ,i; males (Spooner
1968). The details of phaneropterine pair forming sys-
tems vary (Spooner :* *'.,, and even within a genus
some species may employ female answers and others
not (Heller and von Helversen 1993).
Shaw et al. (1990) were first to report on female
acoustic responses for a species in the A. rotundifolia
complex. Using specimens of A. parvipennis from
Ames, IA, they found that females made ticks that i. II
between the phonatomes of the male's phonatome
train. I i .1...1. females produced an average of 3.2
ticks per phonatome train with the last tick coming
after the last phonatome in the train 36% of the time.
At 24-25C, the delay between the beginning of a
phonatome and the tick produced in the following
interval averaged 120 ms. Galliart and Shaw (1991)
found that when afermale phonoresponded, I. -i.,
male .:i. I to the female if both were unrestrained.
C- .11I. I and : i, (1992) reported that if the male was
restrained and the female was free, the responding
female moved to the calling male. Galliart and Shaw
(1996) used computer-generated calls to further study
female phonoresponse and phonotaxis in A. parvipen-
In 1966, J.D.S. recorded acoustic exchanges be-
tween caged males and females of A. bartrami from
Perry County, AL. In these recordings the males usu-
ally made successive trains of five or six phonatomes
and the females ticked one or more times after each
train. At 23C, the time between the end of the pho-
natome train and the first female tick varied from 138
to 688 ms (n = 10, mean = 416). When a male pro-
duced a longer train, the answering female would
sometimes start ticking before the train ended. In
these cases, the females showed no tendency to time
their ticks relative to the phase of the phonatome
Years later, J.D.S. studied acoustic pair formation in
A. nr bartrami II. -. i .: near Aiken, SC. In a 1988
recording of a caged male and female (J.D.S.-Paml
8,-9; 25.3oC), the female at first ticked only in phase
with the phonatornme intervals of prolonged trains. To
trains of 21, 24, and 21 phonatomes, she did not begin
answering until the seventh phonatome in the train or
later. The timing of the answers (always a single tick)
varied little relative to the beginning or end of the
:-.... ..1,. phonatorne. In a sample of 10, the delay
from the beginning was 103-140 ms (mean = 134) and
from the end. 21-43 (mean = 35). No more than 6
phonatomes in a train were answered with the earliest
Vol. 96, no. 4
WALKER ET AL.: Amblycorypha .' '. :" '.
answer coming after the seventh phonatome and the
last phonatome being answered in two of the three
trains analyzed. When the male began to produce
trains of 4 7 phonatomes, the female would answer
one or two phonatomes keeping the timing as before
When the male started to produce long trains again,
the female would sometimes produce multiple ticks
with little or no regard to the phase of the phonatome
period and at other times would deliver single ticks in
the intervals as earlier. Overall, in this recording, A. nr.
bartrami resembled A. parvipennis more than A. bar-
trami in both phonatome rate (5.20/s) and in the
timing and placement of female ticks.
In a report summarizing his studies of A. nr. bam-
trami, J.D.S. (Spooner :. *.! noted that responsive
females, after hearing the first few short trains in a
song, made one to three ticks at the end of each
additional short train. When a long train was pro-
duced, they would begin to vigorously tick before it
ended a, I I II .,, the end, make several more ticks.
(These phonoresponses resemble those of the Perry
Co., AL bartrami.) If either sex was restrained, the
unrestrained sex moved to the restrained. If both male
and female were freed as they called and ticked 4 mr
apart, they moved toward each other and met about
midway between their release points.
The phonatome rates and intervals of the remaining
two species fall on either side of those treated above.
The phonatome rate of A. :.......^ .. is more than
double that ofA. bartrami, suggesting that female ticks
would not be restricted to a : .. i;- part of the pho-
natome period and would more likely come at the end
of pulse trains. In keeping with these expectations,
when a caged female from Berks County, PA, an-
swered the rattle of a male from the same locality, in
12 of 17 cases the ticks began after the rattle ended. In
the other five cases, the ticks began during the rattle,
.. *i **near the end. When the ticks began after the
rattle, the delays averaged -130 ms (range, = 91-366;
24.70C WTL-8 lb). T.G.F. recorded >220 tick an-
swers from four A. rotundifolia females duetting with
three different A. rotundiolia males (Buncombe
County, NC; at 22.3-23.7C). Twenty-six percent of
the answers occurred during the rattle; the rest oc-
curred during the intervals between phrases, usually
between the short phrases following the prolonged
portion of the song. On average the delays were
-200ms but were highly variable and ranged from 140
The phonatome rate of A. alexander is little more
than half that ofA. parvipennis, suggesting that females
should tick in the intervals between phonatomes. In
two recordings of a pair of A. alexander from Bun-
combe County, NC, T.G.F. found that the female's
answers (N = 69) -,.. :i occurred between the pho-
natomes in the last half of the male's series of clicks and
never during the first four clicks in the series. At 22.7
and 23.3C, the delays in the female's answers aver-
aged -270rs (range, 29 335ms). The variation in the
delay was in part because of the variability in the
period of the males' clicks. When the female's delays
are expressed as a percent of the cycle period of the
clicks, 42 of the 69 answers were produced during the
last 15% of the phonatome cycle. For six of the nine
trains answered, the female responded to the final
phonatome in a train.
Ecology. Throughout their ranges, A. rotundifolia
and A. alexander occur in the understory of broad-
leaved woods and in weedy roadsides and fencerows
(Alexander 1956, 1960; current study). No differences
in habitat preferences have been identified, and the
two species sometimes occur in mixed colonies (Al-
exander 1960). In 4 yr, near Asheville, NC, where
T.G.F. studied five sites with A. ..,. -' '... popula-
tions and five with A. alexander populations, he nei-
ther found a site with both species nor developed a
means of predicting which species would occur at an
occupied site. On the access road to Black Rock Moun-
tain in Rabon County, GA, J.D.S. found A. alexander
and A. rotundiflia in similar habitats but 0.4 km
In west Florida, where A. alexander and A. bartrami
co-occur, the former species was found exclusively in
mesic, broadleaved woods (beech-maple in Jackson
County and ravine forest in Liberty County), whereas
the latter was found exclusively in more xeric, sandhill
habitats. Sandhills habitats are characterized by fre-
quent fires and, now that most of the longleaf pines
have been lumbered, .. -. 11I are dominated by tur-
key oaks. Most other collections of A. bartrami were
also in sandhills, but in Perry and Cleburne counties,
AL, and Stanly County, NC, the species was found in
the hardwood understory of : I.! .:1 pine woods.
The original habitat of A. parvipennis was appar-
ently prairie and its interface with deciduous wood-
land. It is now numerous in weedy pastures and old
fields as well as prairie remnants (Isely 1941; Shaw et
al. 1981, 1990). It feeds avidly on I. 1 and on buds
and young leaves of a variety of plants (Isely 1941).
The seasonal life cycles of the four species are so
similar that in localities where two of the species
occur, no differences have been rioted. Overwintering
is in the egg stage. Adults first occur in June in the
southern states and in July to early August farther
north. Adults soon reach a maximum abundance and
then decline. No second peak of adult abundance
occurs, which suggests a univoltine life cycle. How-
ever,because many ,* .1.1 ..1. i. ,.],. K', rypha
oblcngifolia (De Geer), have eggs that require >1 yr
to hatch, a longer life cycle cannot be ruled out (Han-
cock 1916, Ingrisch 1986).
All females of the A. rotundifolia complex probably
lay their t ... i I i .. (1941) described oviposition
of A. parcipennis in north-central Texas, in which the
female uses her mrouthparts to help guide the ovipos-
itor into the soil. With her ovipositor fully inserted, she
continues to hold it with her jaws while as many as 14
eggs are deposited in the soil in a loose cluster. J.D.S.
found that captive females of A. rotundifolia and A. r.
bartrami would lay in soil, although in this case no
alternative oviposition substrates were offered. The
role of the arrays of acuminate teeth on the distal
margins of ovipositors in the rolundifolia complex is
unknown (Fig. 9). No similar teeth occur on the ovi-
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Fig. 6. Id-,riti iir characters of the Amblycorypha ro-
tundifolia complex, as illustrated by the holotype of A. alex-
anderi. (A) Left lateral lobe of the pronotum. Arrow points
to shallow humeral sinus. () Right metasternal lobe (spec-
imen is inverted with the head to the right; pin pierces right
mesosternal lobe). Arrow points to the lobe's caudal margin,
which is truncated or nearly so. The width of the lobe,
measured from the ventral midline and perpendicular to it,
exceeds the length of the lobe as measured perpendicular to
the caudal margin.
positors of members of the A. oblongifolia complex,
which also lay in soil.
Morphological Characters. Each of the species pos-
sesses the morphological characters that distinguish
members of the A. rotundifolia complex from other
species of Amblycorypha-namely, a shallow humeral
sinus and transverse metasternal lobes. Because the
state of neither of these characters is easy to judge
without specimens to compare, we have illustrated
them in Fig. 6A and B.
Only one of the four species in the complex can be
reliably identified on the basis of morphological char-
acters. In A. parvipennis the tegmina conceal the hind-
wings at rest, whereas in the other three species the
tips of the hindwings, when normally developed, pro-
trude 2 6 mm beyond the tips of the tegmina. A series
of seven specimens of A. bartrami from Perry County,
AL, 9 June 1966, prompted the caveat about normal
Table 1. Measurements in millimeters of the three eastern
species ofthe Amblycorypha rotundifolia complex. Specimens of A.
alexander from Florida and from more northern localities are
listed separately because the means of their measurements differ
A. A. alexander A
Mrotundfolia Northern Florida bartrami
mean (n) 5.4(31) 5.4(11) 5.6(12) 5.7(21)
range 4.8-5.9 4.5-5.5 5.0-6.0 5.2-6.2
mean (n) 26 (30) 26(11) 30(11) 29 (19)
range 23-29 24-28 27-32 24-32
mean (n) 3.0(28) 3.0(10) 3.3(11) 3.3(19)
range 2.7-3.4 2.8-3.2 3.1-3.7 2.9-3.5
mean (n) 4.3(27) 4.1(11) 5.2(9) 4.9 (14)
range 3.1-5.2 3.3-4.7 4.6-5.8 3.1-6.0
mean (n) 6.1 (12) 6.1(7) 5.9 (1) 5.8 (5)
range 5.8-6.4 5.8-6.4 5.4-6.5
mean (n) 26 (12) 25(7) 28 (1) 26(4)
range 23-28 23-27 23-29
mean (n) 3.0 (12) 3.1(7) 3.3(1) 3.2(4)
range 2.8-3.3 2.9-3.2 3.0-3.4
mean (n) 2.4(12) 2.4(7) >3.4 (1) 2.5(1)
range 1.6-3.2 1.9-3.0
mean (n) 9.9 (11) 10.1 (7) 10.2 (1) 9.7 (5)
range 9.3-11.0 9.7-10.9 9.3-10.0
development: five had shriveled hindwings about half
the length of the tegmina. In A. parvipennis the hind-
wings are normally developed and their tips nearly
reach, but do not exceed, the tips of the tegmina.
Routine measurements of the remaining three spe-
cies provided no identifying (nonoverlapping) fea-
tures (Table 1). The measurements of A. alexander
are noteworthy in that those from northern popula-
tions resemble those of A. rotundifolia, the northern
species of the trio (first two columns of data in Table
1), and those from Florida populations resemble those
of A. bartrami, the southern species of the trio (last
two columns in Table 1). Indeed, the only populations
that differed enough to be identifiable by their mea-
surements were the northern and Florida populations
of A. alexander. However, for univoltine species of
tettigoniids with extensive distributions in eastern
United States, clinal increase in size from north to
south is commonplace. Therefore, if we had had spec-
imens of A. alexander from intermediate localities,
they would presumably have been intermediate in
Because the songs of the eastern trio are so differ-
ent, we looked for differences in characters of the
stridulatory files. We found none that were of use in
separating the species. Both in file length and number
of file teeth the species broadly overlap (Fig. 7).
Vol. 96, no. 4
WALKER ET AL.: Amblycorypha .' '. '" '.
A rotundifolia 0
*bartrami A A .
S90 A *
1.50 1.70 1.90 2,10 2.30 2.50
Fig. 7. Length of stridulatory file and number of file teeth
for males of the three eastern species of the A. ..... ..
complex. A. alexanderi (n = 6; four counties in Florida,
Georgia, and Ohio), A. = .... (n = 7; six counties in
Georgia Illinois, Ohio, Pennsylvania, and Tennessee), and A.
barr-ami (n = 6; five counties in Alabama, Florida, and North
Characters of the ovipositor sometimes are useful in
differentiating phaneropterine species, but our study
of the ovipositors of A. ., \ alexander, and
A. bartrami revealed no species-i..1, 1*i._ --. features.
Neither the absolute length of the ovipositor nor the
length relative to the pronotal length was species spe-
cific (Fig. 8). However, the ovipositors of Aiken A. nr.
bartrami were longer than those of any other speci-
mens examined. In the southern portion of its range,
ovipositors ofA. rotundiflia (n = 10, North Carolina,
Georgia) had the marginal teeth produced into fine
points, similar to those ofA. alexander and A. bartrami
(Fig. 9C). Farther north the teeth on A. rotundifolia
ovipositors (n = 5, Illinois, Pennsylvania) lacked the
fine, sharp points (Fig. 91). Thus the apparent geo-
graphic variation in ovipositor teeth of A ..
as in the geographic variation in the overall size of A
alexander, is in the direction that makes separating
sympatric A. rotundifolia and A. lexanderi more dif-
.i ,- th ,, th,,-I. l1 C.
Geographical Distribution. Figure 10 shows the dis-
tribution of the four species as currently understood.
A. .. .' r. -. occurs farther to the north and west
than the two new species with which it was confused.
Records for Indiana and Illinois are abundant because
of extensive published records that were deemed valid
without confirming song data, as required in areas of
sympatry. Most Ohio records are from the field notes
of E. S. Thomas and R. D. Alexander, as made available
to T.J.W. by Alexander (personal communicationn.
The paucity of records in Kentucky and Tennessee
may reflect scanty fieldwork in those states.
All records of A. parvipennis are from west of the
Mississippi River. A study of calling songs of "rotun-
difolia" in I ::.. and of "parvipennis" in eastern Iowa
might reveal a geographical overlap not suspected
from studies of museum specimens. For example, A.
parvipennis could vary geographically in wing length
but keep its reproductive integrity by virtue of its
A. alexander is ... i sympatric with A. rotundi-
folia in the north but extends much farther south. Its
southern-most records are from a region of Florida
known for its endernic plants and animals and for
being the southern limit of many northern species
(Hubbell et al. 1956). The Florida populations may be
disjunct from those farther north, or. more i,
populations in habitats suited to A. alexander along
the Chattahoochee River, may connect the Florida
populations with those in the southern Appalachians.
A. bartrami probably occurs wherever there are
sand-hill habitats in southeastern United States. How-
ever, the three records that mark its northwestern
lirrits (Stanly County, NC, ard Cleburne and Perry
Counties, AL) are from more mesic habitats. This
suggests that A. bartrami may occur farther to the
north and west than the limits indicated in Fig. 10.
Studies of calling songs of North American crickets
and katydids have revealed many previously unrec-
ognized species and resolved several taxonomically
* orotundifolia difficult groups (e.g., Thomas and Alexander 1962,
Salexanderi (fl) Walker et al. 1974). The usefulness of i:.. songs
a alexander (n) stems from the direct evidence they give of repro-
5o bartrami (fl, al) ductive isolation among populations. Populations with
nr. bartrami (sc) different calling songs are using .1... .. signals to
a mediate sexual pair formation and hence are :.: i to
I be reproductively isolated. When laboratory crosses
AA A produce hybrids between populations with different
S B songs, the hybrids have intermediate songs (Alex-
5 o ander 1968). Thus, a failure to maintain reproductive
o < isolation between sympatric populations of song-de-
0 --fined species would be apparent from the occurrence
5.0 5.5 6.0 6.5 of intermediate calling songs.
Pronotal length (mm) Once putative species have been defined on the
8. Ovipositor length as a function ofpronotallength, basis of calling songs, morphological differences that
ack triangle is for the allotype and sole southern female correlate with the song differences are usually found.
ten ofA. alexander. The black squares are for females This makes possible identification of specimens that
populations near Aiken, SC. lack song data, thereby aiding further study and fa-
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Fig. 9. Ovipositors. (A) A. alexander, allotype. (B) A. bartrami, allotype. (C) A. rotundifolia Murray Co., GA. (D) A.
rotundifolia, Douglas Co., IL.
cilitating the description of the species that were first alexander and A. bartrami in the southern portion of
recognized by their songs. the range of A. alexander. When populations that
On the basis of their calling songs, nearly all males differ in calling songs are in local contact and do not
of the A. rotundifolia complex can be unambiguously hybridize (no intermediate calling songs), they vali-
assigned to one of the four species we recognize here, date their status as species. To deny them formal
but our studies of these males and associated females names would make it unlikely that field biologists
have uncovered no morphological characters that per- would know that two species of rotundifoliaa" might
mit definite identification of the three eastern species, exist in their study areas, and those who did so realize
Our naming two new species before being able to would be hindered in reporting their findings.
separate them other than by song is driven by the Female answers are apparently essential to pair for-
sympatry of A. rotundifolia and A. alexander in most nation for species in the A. rotundifolia complex. This
of the range of A. alexander and the sympatry of A. means that males must make songs that have features
Vol. 96, no. 4
WALKER ET AL.: Amblycorypha rotundifolia
Fig. 10. Gr 2, .1phi( .lA di t i hiiit fi nt the species in tl. \ i ... .1, ...... ... i,.1i!.i.., complex. In the map for each species,
dots represent county records and shading indicates probable geographical limits. The open circle in the A. bartrami map
is for a population near Aiken, SC, that has lower phonatome rates and longer ovipositors than populations elsewhere.
that stimulate their females to tick. Among the species
here recognized, phonatome rate and phonatome
structure are probably important. However, the ticks
of females have no distinguishing physical features but
must be delivered at times during the song that render
them meaningful to their males. This suggests that
studies to determine what features of calling songs
make females tick and what timings of ticks stimulate
males to approach will shed light on which of the
variations in these two features noted here are likely
to have roles in maintaining reproductive isolation
among sympatric populations.
When considering species concepts, it is important
to realize that all species are local (Lloyd 2002). This
means, for example, that we cannot be certain that A.
alexander in the southern Appalachians and A. alex-
anderi in northwestern Florida are the same species.
We know that we can distinguish most specimens from
the two areas by their sizes (and judge that to be of no
consequence), yet we do not know whether individ-
uals from the two areas would mate and produce
fertile offspring were they given the chance. However,
we have strong evidence that individuals we identified
as A. alexander from Torreya State Park in Liberty
County, FL, belong to a different species from those
individuals we identified as A. bartrami from the same
place. Similarly, we know thatA. alexander is a species
distinct from A. rotundifolia at numerous sites in the
southern Appalachians. In contrast, we can notbe sure
whether the sole species of the A. rotundifolia complex
known from near Aiken, SC, is the same as or different
from the species named A. bartrami with its holotype
coming from Alachua County, FL. In this paper we
have recognized a minimum number of species and
described variations that we have found among the
populations assigned to each of the four names.
It is important to understand that when sympatric
populations have different songs they are almost cer-
tainly different species, but that when allopatric pop-
ulations have the same song they are not necessarily
conspecific. The case of the cricket genus Pictonemo-
bius (Gross et al. 1990) exemplifies the difficulty of
extending locally defined species to distant areas. Pro-
duction of four types of calling songs among Pic-
tonemobius populations in the vicinity of Gainesville,
FL, indicated four species, and the reproductive iso-
lation of the four was confirmed by laboratory crosses
and electrophoretic analysis. Two of the species were
morphologically distinct. However, when Pictonemo-
bius populations to the north and west of Gainesville
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
were studied, their identities relative to the Gaines-
ville species were different depending on whether
appearance, songs, or allozymes were used to make
the assignments. L local species in Gainesville did not
necessarily correspond to any of the local species at
places a few hundred kilometers away. Because all
Pictonenobius are flightless and some species live in
habitats that are :. .,i distributed, it is easy to un-
derstand how microgeographic variants might de-
velop and remain distinct. So far as we know, all
members of the A .... 7.'" .. complex are likewise
flightless and the sandhill habitats occupied by A.
bartrami are often discontinuous.
An intriguing aspect of differences among the songs
of the four species in the A. rotundifolia complex is that
their phonatome rates approximate a doubling series.
This is surprising because closely related species of
ensiferan Orthoptera usually have .ii songs more
modestly divergent in phonatome rates, making the
differences i. attributable to gradual increases or
declines (Otte 1992). Expected phonatome rates at
25C, as calculated from the regressions depicted in
Figs. 1 and 2, are 2.8 ph/s (mean for A. alexanderi, 4.8
(parvipennis), 9.5 (bartrami), and 26 (0. .. ?.. .
The rates thus increase by factors of 1.7, 2.0, and 2.7.
[In theory, the temperature selected for the compar-
ison should not affect these factors because with re-
peated measurements under controlled conditions the
phonatome versus temperature regressions for closely
related ensiferan species tend to converge at some
temperature for an expected rate of 0 e.g., 11C for
Orchelimum species (Conocephalinae) (' tII ,
S ).] Doubling of phonatome rates among closely
related species is rare but not unprecedented among
ensiferans: The rates for two closely related species of
Cyrtoxipha (Trigonidiinae) are 19 and 39 ph!s (Walk-
er 1969), and Otte (1992) reported two examples
among Hawaiian Trigonidiinae. Alexander and
Thomas (1959) noted that the courtship phonatome
rate for Nemobius tinnulus Fulton is about half that of
the calling song rate. [In cricket songs, pulses and
phonatomes are equivalent ]
If most of the ,llt .- among the phonatome
rates in the A. rotundifolia complex resulted from
doubling or halving the rates during speciation, vari-
ations in those directions should occur in the songs
that are currently produced. A potential for doubling
can be seen in the excerpts from A. alexander songs
in Figs. 4E and F. A. alexander not only has the longest
phonatome (-140 ms at 250C), it also has the longest
phonatome interval (-300 ms s. ; and the great-
est proportion of the phonatome period (phonatome
+ phonatome interval) occupied by the interval
(-68%). In other terms, the slow phonatome rate is
attributable both to long phonatomes and long delays
between phonatomes. The long delays could be a
result of dropping alternate phonatomes. The mini-
mum interval between successive phonatomes in A.
alexander (Fig. 4) is brief enough to support this
In some species of phaneropterines, males produce
different songs to induce females to approach (pho-
notaxis) and to make answering ticks (phonore-
sponse). Spooner (1968) designated the two as calling
songs and female tick elicitors. Males of the A. rotun-
difolia complex make no such separate songs. What we
have termed the .il.-. song in our study of the A.
:.." ', !..:1. probably has both functions in all
four species, although the calling function has only
been demonstrated in A. parvipennis and Aiken A. nr.
bartrami (Galliart and Shaw 1991, Spooner I *' ).
We believe that the four species treated here are
monophyletic. In keeping with this, we used "round-
winged katydid" (from Blatchley 1920) in each of the
four vernacular names we proposed. However, those
wishing to assign Armblycorypha specimens to the
round-winged katydid group of species should be
warned that most have tegmina that are little if any
more rounded than specimens in other species groups
of Amblycorypha. Rehn and Hebard (1914) recog-
nized three morphological modes among North Amer-
ican Amblycorypha. Their Group I, included two
modes, a group of four robust species t :..i.. by A.
.. _. ..... 1 a single species of small size, A. uhieri.
As mentioned earlier, their Group II was the A. ro-
tundifolia complex, which they distinguished from
Group I by the states of the humeral sinus and the
metasternal lobes. T,. also used differences in the
caudal margin of the pronotal disk and the proportions
of the tegmina as key characters. However, we have
found these to be of little use. In regards to tegminal
proportions, Rehn and Hebard (1914, p. 319) stated
that the tegmina in Group II are "rarely over two and
four-fifths times as long as greatest width," whereas
our Table 1 shows that few of the specimens we ex-
amined had wings that round. All our mean values of
tegminal ratio equaled or exceeded the 3.0 that Rehn
and Hebard set as the minimum value for their Group
We thank these persons for their assistance: R. D. Alex-
ander and Ken Shaw for advice on the manuscript; R. E. Love
for lI .... ., :.., G.K.M orrisfor i: :.. I I.
O'Brien for help with specimens of A. parvipennis and F. W.
Walker's field notes; Chris Cleasby for help with i ,-
T.J.W. and J.D.S. were supported by National Science Foun-
dation during some of their field work. T.G.F. was supported
in part by a grant from the Grass Foundation. Florida Agri-
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Received for publication 28 October 2002; accepted 7 Feb-