Research Article

Journal of Orthoptera Research 2019, 28(1): 3-9

Anaxipha hyalicetra sp. n. (Gryllidae: Trigonidiinae), a new sword-tailed

cricket species from Arizona

JerFrey A. Cote!2, Davip H. Funk?

1 Natural Sciences Division, Pasadena City College, 1570 East Colorado Boulevard, Pasadena, CA, 91106, USA.
2 Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, CA, 90007, USA.

3 Stroud Water Research Center, Avondale, PA, 19311, USA.

Corresponding author: Jeffrey A. Cole (jacole@pasadena.edu)

Academic editor: Juliana Chamorro-Rengifo | Received 28 September 2018 | Accepted 29 December 2018 | Published 11 April 2019

http://zoobank.org/53A484AD-2D25-4DB7-8C36-A9666237829C

Citation: Cole JA, Funk DH (2019) Anaxipha hyalicetra sp. n. (Gryllidae: Trigonidiinae), a new sword-tailed cricket species from Arizona. Journal of

Orthoptera Research 28(1): 3-9. https://doi.org/10.3897/jor.28.30143

Abstract

A new Anaxipha species is described from a locality in southeastern
Arizona adjacent to the border with Mexico. The species is unique among
the North American fauna by virtue of the broad tegmina, distinctive male
genitalia, and calling song phrased in an irregular chirp with a variable
pulse train rate. The possibility that the behavioral repertoire of this spe-
cies includes aggressive song as well as calling song is discussed.

Key words

aggressive song, calling song, mate recognition, Mexico, Neotropical, Sonora

Introduction

Anaxipha is a speciose genus of small crickets with a worldwide
distribution (Walker and Funk 2014, Cigliano et al. 2018). The
systematics of Anaxipha, and of the Trigonidiinae in general, are
in a primitive state (Otte and Perez-Gelabert 2009). Regional sys-
tematic studies have revealed entire faunas of these crickets in re-
cent years (Otte 2006, Otte and Perez-Gelabert 2009, Walker and
Funk 2014), and numerous new species await description (e.g. Tan
2012, DHF unpublished data). A revision of the North American
Anaxipha described two-thirds of the fauna as new to science (8
of 13 species; Walker and Funk 2014). All species known to date
occur in the eastern portion of the continent (Walker and Funk
2014, Cigliano et al. 2018).

While engaged in general Orthoptera collecting in southeast-
ern Arizona in 2013, the first author followed an unfamiliar call-
ing song to discover an Anaxipha population. Apart from the high-
ly disjunct locality, which is adjacent to the border of the United
States with Sonora, Mexico, this Anaxipha immediately stood out
as new to the North American fauna by virtue of the broad tegmina
of males and the variable pulse train rate in the male calling song.
Laboratory study revealed distinct genitalia and stridulatory file
characteristics. No morphologically similar Anaxipha were found
among specimens at the Academy of Natural Sciences of Drexel
University (DHF pers. obs.). The species is described here as new.

Methods

Fieldwork was performed at dusk and at night during 15-16
July 2013, 23 July 2014, and 29-31 August 2014. Triangulation of
calling songs, visual inspection, beating, and sweeping of vegeta-
tion were all employed to secure specimen series. Specimens are
deposited at the Natural History Museum of Los Angeles County
(LACM), at the Academy of Natural Sciences of Drexel University
(ANSP), and at the University of Florida (UFL).

Habitus images (Fig. 1) were made at LACM using a digital
microscope (model VHX-6000, Keyence Corp., Itasca, IL) with
a 10x objective lens. With this microscope, a calibrated anchor
point-based digital measurement function was used to obtain
the following morphological measurements (in mm) for a subset
of dried specimens deposited at LACM: body length (BL, meas-
ured from the front of the head to the end of the tegmina for
males and to the end of the ovipositor for females), body width
(BW, measured at the widest point of the tegmina immediate-
ly behind the hind trochanters), and hind femur length (HF).
Measurements are reported as mean + standard deviation (range)
separately for each sex.

Male genitalia were removed from a pinned specimen that had
been relaxed and cleared in hot 10% KOH. These were mounted
in glycerin on a well slide and photographed using a trinocular
microscope (Labophot, Nikon Inc., Melville, NY) equipped with
a digital SLR (model T1i1, Canon Corp., New York, NY) controlled
with Canon EOS Utility software in Live View mode. Images taken
at several focal points were composited in Adobe Photoshop CS3
software (Fig. 2A). Afterward, the genitalia were transferred to a
genitalia vial with glycerin which now resides with the specimen.

The right tegmen of the above specimen was removed and
slide-mounted in euparal. The tegmen was then photographed
with the trinocular microscope (Fig. 2B). A single digital image
was constructed from multiple fields using Adobe Photoshop.
File counts were made from the photograph and, starting at the
hind margin, a small red dot was added every 10" tooth to fa-
cilitate counting (Fig. 2C). Linear measurement was made from
the first tooth (closest to hind margin) to the junction of the

JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(1)

4 J.A. COLE AND D.H. FUNK

stridulatory vein and the harp vein. This measurement was ex-
pressed as length of cell 5 (sensu Otte 1994) and tooth density
was estimated from this region in order to minimize errors at-
tributable to straight-line measurement of the (slightly curved)
stridulatory vein.

Songs were recorded in the field and in the laboratory using a
digital linear PCM recorder with a built-in condenser microphone
pair (model PCM-D50, Sony Corp., New York, NY). This device
recorded .wav files at a 96 kHz sampling rate and a bit depth of 16
with a flat response range extending to 40 kHz. The temperature
of laboratory recordings was controlled at 24.1 + 0.7°C (Table 1).
Temperature was measured with a digital thermal sensor (model
Trail Pilot 2, Highgear USA, Inc., Fletcher, NC).

Recordings were visualized (Fig. 3) with Raven Lite v. 2.0
(Cornell Laboratory of Ornithology, available from http://www.
birds.cornell.edu/brp/raven/RavenOverview.html) and analyzed
using BatSound (Pettersson Elektronik AB 2001). The Pulse Char-
acteristics feature of BatSound automatically extracted pulse du-
rations, intervals, and peak frequencies from the .wav files. Os-
cillogram thresholds for pulse characteristics extraction were set
between -18 and -10 dB, with higher values necessary to avoid
higher noise floors. Noise below 2 kHz was attenuated with a
Butterworth high pass filter in BatSound before analysis. Spectro-
gram settings for peak frequency measurement were a Hanning
Window employing a 256 Hz fast Fourier transformation. Song
terminology follows that of Walker and Funk (2014). Abbrevia-
tions for song characters are reiterated here for convenience and
are graphically shown in Fig. 3:

pulse duration
PI pulse interval

PP pulse period (=PD+PI)

PR pulse rate (=1/PP)

Pde pulse duty cycle (=PD/PP)

PA. pulse train

PN number of pulses per train
PTD pulse train duration

PTI pulse train interval

PIP pulse train period (=PTD+PTI)
PTR _ pulse train rate (=1/PTP)

PTdc_ pulse train duty cycle (=PTD/PTP)
PTCF pulse train carrier frequency

Statistical analysis was performed in R (R Core Team 2015).
Anaxipha songs are complex, and our analysis considered 12 song
characters. Besides those characters that are mathematically re-
lated (see abbreviations above), additional correlations may exist
among characters. To find statistically independent variables for
analysis, we explored song character data structure with maximum
likelihood factor analysis. The number of components to extract
was determined by examining a scree plot generated with the
nFactors package (Raiche and Magis 2015). A distinct inflection
point (“knee”) in the plot of eigenvalues vs. components indicates
the number of components to extract. The number of components
output by the nFactors scree plot was then input into the factanal
function, with rotation set to varimax. The output of this func-
tion consists of statistically independent factors, together with the
magnitudes and signs of the contributions of the song characters
to each factor. A MANOVA compared song character means be-
tween field and laboratory conditions.

Results

Anaxipha hyalicetra Cole & Funk, sp. n.

http://zoobank.org/1CF3949F-F602-4AEA-ACB6-FF1195B2766D
http://Isid.speciesfile.org/urn:lsid: Orthoptera.speciesfile.
org:TaxonName:505367
Fig. 1: habitus and morphology; Fig. 2: male genitalia, tegmina, and
stridulatory file; Fig. 3: songs, recording 140723_11, available as Suppl.
material 1.

Holotype.-—1 male. USA. ARIZONA: Santa Cruz County; Pena
Blanca Canyon, Coronado National Forest, 31.38230, -111.09251,
elevation 1203 m. 23-VII-2014. J.A. Cole leg. Recording number
140814_00. Prepared with tegmina raised. Right antenna missing
most of the flagellum, otherwise complete (Fig. 1A, B). Deposited
in LACM.

Paratypes.—3 males, 1 female (pinned), same data as holotype
(ANSP); 5 males, 1 female, same data as holotype (FSCA); 4 males,
2 females (pinned), 4 males (DNA vouchers SING0518, SINGO519,
SING0520, SINGO521 in 100% ethanol), same data as holotype
(LACM); 2 males (pinned), same locality as holotype collected
31-VIII-2014 (LACM); 5 males (pinned), 2 males (DNA vouchers
SING0453, SING0454 in 100% ethanol), same locality as holotype
collected 15-16-VII-2013, J.A. Cole and J.E Limon leg. (LACM).

Measurements.—Males (n = 7): BL = 7.63 + 0.24 (7.38 - 7.95),
BW = 3.38 + 0.18 (3.13 - 3.73), HF = 4.20 + 0.14 (4.00 - 4.35);
females (n = 2): BL = 7.31 + 0.27 (7.11 - 7.50), BW = 1.95 + 0.06
(1.91 - 1.99), HF = 4.25 + 0.23 (4.08 - 4.41).

Hind wings.—No specimens among the type series are macropterous.

Seasonal occurrence.—Available records suggest early summer to
midsummer adult activity. Individuals were common from 15-
23 July in two consecutive years. By 31 August 2014, males were
scarce, and no females were found. Males collected 23 July lived in
captivity until 19 August.

Habitat.—The population resides in a north-south trending can-
yon. Within the canyon, individuals are most common in the
creek bed at the canyon bottom but extend a short distance up the
canyon walls into mixed woodland. During both July collecting
events monsoon rains had recently fallen in the canyon and hu-
midity was high. Crickets were found on catclaw acacia (Senegalia
greggii (A. Gray)), on stems of pointleaf manzanita (Arctostaphylos
pungens Kunth), on Yucca, on bunch grass, and on oak leaf litter.
Like other North American Anaxipha, individuals perched within
1 m of the ground (Walker and Funk 2014). Acoustic activity was
observed in the field from 19:41 to 22:05 h. In July and August,
A. hyalicetra was sympatric with Oecanthus californicus Saussure,
an arboreal chirping Oecanthus rileyi Group sp., Gryllus ?persona-
tus Uhler, and two undescribed Gryllus species. Cycloptilum sp. and
Hoplosphyrum sp. scaly crickets (Mogoplistidae) were found in the
same habitat during the August 2014 collecting event.

Diagnosis.—A. hyalicetra has a unique combination of morpho-
logical characters among the North American Anaxipha fauna. The

JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(1)

J.A. COLE AND D.H. FUNK 5

1lmm

-—

imm

Fig. 1. A. Holotype male, dorsolateral view; B. Holotype male, anterior view; C. Paratype male, living habitus; D. Holotype male, left
hind femur; E. Holotype male, face; F. Paratype female, dorsal view; G. Paratype female, lateral view.

JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(1)

6 J.A. COLE AND D.H. FUNK

0.5mm

eu

O00 OF Q2 O03 O4 O5 O86 QO7

08 o9 WO UL

Is 16

SRR en Oe te

2 13° 1A

Fig. 2. A. Paratype male genitalia, oblique posterodorsal view, P = paramere, ML = median lophus; B. Paratype male excised left tegmen
(recording numbers 140811_01 and 140723_15), posterior edge of tegmen at top of image; C. Paratype male, stridulatory file, same

male imaged in B.

basal segment of the hind tarsus is longer than segments 2 + 3
combined, a feature that is also found in A. imitator (Saussure),
A. calusa Walker & Funk, and many Neotropical species. The male
tegmina in A. hyalicetra are broader than in every other Anaxipha
species (Fig. 1B, C), although many Neotropical species have
broad tegmina. With 124 teeth in the stridulatory file (Fig. 2C), A.
hyalicetra overlaps in file characteristics only with A. exigua (Say),
which is a fall species with narrow tegmina found in eastern decid-
uous forests. The male genitalia (Fig. 2A) are unique: the median
lophi are bifurcate and hook inward, each paramere has a hooked
tooth at the anterolateral corner, and the parameres slope antero-
laterally from the midline (rather than a posterolateral slope or
perpendicular orientation, cf. plate 13 Walker and Funk 2014).
The variable PTR in male songs (Table 1, Fig. 3C) is unique among
North American Anaxipha (see Acoustic behavior below).

Etymology.—l. hyalo (glassy) + cetra (a small light shield), referring
to the broad, transparent male tegmina.

Acoustic behavior.—PR in A. hyalicetra is 45.4 s' (Table 1, Fig. 3B), iden-
tical to that of A. fultoni Walker and Funk and close to that of A. imita-
tor. Phrasing differs between these species. The calling song phrasing
of A. hyalicetra is a series of PT of irregular length that may be termed
a chirp (Fig. 3A, C). Phrasing in both A. fultoni and A. imitator is a bro-
ken trill rather than an irregular chirp. The calling song phrasing of the

new species is qualitatively similar to that of A. litarena Fulton from
the beaches of the southeastern United States, except that the latter
produces more regular PTR and PTD. The carrier frequency is 6 kHz.
A series of harmonics extend to 43 kHz at 6 kHz intervals (Fig. 3A).

The scree plot (Fig. 4) has a distinct inflection point at n = 4
components. The four components extracted by maximum likeli-
hood factor analysis (Table 2) may be interpreted as follows. Factor
1 describes PTR (together with mathematically related characters
PTI, PTP, and PTdc). Factor 2 describes PTD, which is determined
by PN. Factor 3 describes PR (with mathematically related charac-
ters PD, PP, and PI) and PTCE Factor 4 describes pulse characteris-
tics Pdc, PD, and PI, which contribute to PTD. The factor analysis
model was rejected as a perfect fit for the data (P = 4.64 x 10°');
the four factors explain cumulatively 91% of variation. PTCF is the
least classifiable character (i.e. highest uniqueness at 0.46).

Songs were significantly different between the field and the lab-
oratory (MANOVA, P = 9.32 x 10°). Notably, males produced more
rapid PTR (P = 3.65 x 10°) due to shorter PTD (P = 8.06 x 10*) in
the laboratory (Table 1). PTCF also differed between field and labo-
ratory (P = 1.21 x 10%; Table 1). Song character differences opposed
relationships predicted by temperature: PR and PTR were faster and
PD and PTD were shorter in the laboratory than in the field, despite
a lower mean laboratory recording temperature (Table 1).

Males sang in aggregations but did not settle into predictable
chorus phase relationships, neither synchronous nor alternating

JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(1)

J.A. COLE AND D.H. FUNK 7

Table 1. Song character descriptive statistics. All means are reported + standard deviation with coefficients of variation (CVs) below.
The P row shows P-value results from a MANOVA test that compared recordings between field and laboratory conditions. Temperatures

between these conditions were compared with a 2-sample t-test.

Temp PR PD PI PP Pdc PN PTR PTD PTI PTP PTdc PTCF
Combined 7.24 223.44 340.6 +
et 24.74+0.9 45.4446 14.3423 7943.2 223423 644490 5.7418 3.5+1.2 43.4 184.9 194.0 37.7+10.8 60+0.4
CV 0.03 0.46 0.38 1.33 0.23 0.01 0.55 0.40 16.10 153.06 110.46 0.03 0.03
: 138.9 + 278.2 + 417.0 +
Field mean 25.140.8 44.143.9 15.2421 7043.0 22.842.1 66.8485 6241.7 2941.1 42.7 927.8 925.4 37.8412.3 5.7+03
CV 0.023 0.351 0.282 1.319 0.196 0.011 0.462 0.436 13.141 186.597 121.789 0.040 0.019
Lab mean 24.140.7 47.545.1 12.42.00 9543.2 21.3424 60.2486 4841.6 43406 86.9420.8 146.74349 233.7431.8 375490 6340.3
CV 0.018 0.544 0.306 1.073 0.261 0.012 0.520 0.078 5.002 8.282 4.318 0.021 0.018
B NS 0.015 NS NS NS NS 3.60 x10% 8.06 x10% NS 0.010 NS 1.21x10°
S Te)
= © Eigenvalues (>mean = 3)
GC .
2 4 Parallel Analysis (n= 3 )
a . .
Be = Optimal Coordinates (n= 3 )
Acceleration Factor (n= 3 )
. i f
Tt w
— oO
2 3”
c
g g
a =
= o
a AN
; # LU
=)
i -
w 10
me]
2 -0.00 &
5 l |
20
i PTD i
$1,386 LS 16 l? 1.8 19 rt 2:1 22 23 24

131 focal male increases he focal male decreases dle
10

Ueda! | | wil mT y w it i |

PT of 2nd male

4 5 6 rd 8 9

time (s)

2
=
oS

amplitude (kU)

oS

«15

i)
w
w

Fig. 3. A. Male calling song, field recording number 140723_11, os-
cillogram (above) and spectrogram (below), 10 s window, 25.4°C.
PTR = 2.2 s?, PICF = 6.3 kHz; B. 1 s window of same recording as
in A, expanded from region surrounded by red box, showing pulse
structure; C. 10 s oscillogram of interaction between two males in
the field, recording 140723_09, 23.9°C. Changes in PTR of focal
male are indicated.

(reviewed in Greenfield 2002). Males walked while singing, and,
if on a stem, circled the stem (JAC pers. obs.). In the field, males
were observed baffling by positioning themselves between twigs
JAC pers. obs.), a behavior that may improve broadcast by reduc-
ing destructive interference due to sound radiation to the rear of
the insect (Forrest 1982, 1991, Greenfield 2002). When in close
proximity in the field, one or both males may have increased their
PTR by shortening their chirps (i.e. reducing PTD; Fig. 3C). To a
human observer, the effect was a staccato chirp that reverted back
to a lower PTR with longer PTD over time (e.g. Fig. 3A). PTdc re-
mained unchanged during alteration of PTR.

Components

Fig. 4. Scree test results from the nFactors R package. The distinct
inflection point of the eigenvalues plot at n = 4 components sug-
gests extraction of that quantity of factors.

Table 2. Loadings of song characters onto four factors as returned
by maximum likelihood factor analysis.

Character Factor 1 Factor 2 Factor 3 Factor 4
PTR -0.75 -0.48
PTI 1.00
PAP 0.98
PTdc -0.70 0.50
PTD 0.91 0.37
PN 0.83
PR -0.99
PP 0299
PD 0.47 0:53 0.70
Jesh -0.47 0.46 0.63
Pdc 0.49 0.87
PTCF -0.44 -0.50

JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(1)

8 J.A. COLE AND D.H. FUNK

Discussion

A trickle of new species discoveries shows that there is still much
to learn about the orthopteran fauna of Arizona, despite much histori-
cal systematic attention (Rehn 1908, Hebard 1935a, b, Ball 1942) and
the popularity of the state as a destination for insect collectors and
hobbyists. Mass collecting methods employed by general entomolo-
gists such as beating, sweeping, and light trapping are not comprehen-
sive sampling strategies for orthopteran diversity and were not effec-
tive in collecting this new species. Macropterous Anaxipha are attracted
to lights, especially in the tropics, but such sampling is not effective
for micropterous species. Seasonality is also a consideration. Collec-
tors generally sample Arizona after the onset of the monsoons in Au-
gust, and thus may overlook species that are active early in the season
such as the Anaxipha described here. Two southern Arizona katydids
described in recent years: Bucrates weissmani Walker (Walker 2014)
and Microcentrum latifrons Spooner (Spooner 1988), are also active as
adults early in the season, the latter being sympatric with A. hyalicetra.

Collecting in the Atascosa and Pajarito Mountains in Santa Cruz
County, Arizona, failed to find this cricket outside of the type local-
ity. Collecting in adjacent Sonora, Mexico, may prove more fruitful.
Anaxipha hyalicetra is evidently part of an incursion of Neotropical
Anaxipha into North America. Once alpha diversity and higher level
taxonomic relationships are better known, the species with long
first hind tarsal segments may be transferred to a new genus. This
set includes A. hyalicetra, A. calusa, and A. imitator among the North
American fauna together with the majority of Neotropical species.

As we observed song character differences between field and lab-
oratory conditions that oppose those predicted by temperature, the
possibility that A. hyalicetra male behavior includes an aggressive chirp
is worth investigating. Specialized aggressive songs in crickets are brief
chirps that may be produced more frequently and at higher amplitude
than calling song chirps (Alexander 1962, Brown et al. 2006). A stac-
cato chirp characterized by fast PTR and short PTD (Table 1, Fig. 3C)
was observed in field situations where two males were in close prox-
imity. We have not performed controlled experiments to determine
whether these song changes function in aggression, but it is possible
that unintentional artificial crowding in the laboratory may have in-
duced aggressive acoustic behavior, where these same changes to PTR
and PTD were also observed. Aggressive songs signal competitive po-
tential (Brown et al. 2006) should an encounter escalate to physical
combat (Jang et al. 2008, Bertram et al. 2011, Bertram and Rook 2012),
a behavior that is often observed in crickets that defend burrows or
shelters (Jang et al. 2008, Bertram and Rook 2012). As A. hyalicetra is
a vegetation-inhabiting cricket that has not been observed fighting,
aggressive signaling behavior may function instead to regulate male
spacing or increase male attractiveness (Mhatre and Balakrishnan
2006, Chamorro-R et al. 2007). An intriguing hypothesis worth test-
ing in A. hyalicetra is that when aggregated, males may increase energy
expenditure to appear more attractive to listening females (Jia et al.
2001, Mhatre and Balakrishnan 2006). PTR adjustments do not alter
PTdc, which has among the lowest CV of any of the 12 measured song
characters (2-4%; Table 1). PI'dc may be a signal component that per-
mits mate recognition (Mendelson and Shaw 2012) despite behavio-
ral adjustment of PTR during intrasexual competition.

Acknowledgments

Thomas J. Walker contributed a valuable critique of an early
version of the manuscript. The Orthopterists’ Society supported
the cost of publishing this work. The manuscript was improved by
the suggestions of two anonymous reviewers.

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Tettigoniidae: Conocephalinae: Copiphorini). Journal of Orthoptera
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Supplementary material 1

Authors: Jeffrey A. Cole, David H. Funk

Data type: WAV file

Explanation note: Male calling song supplementary file.

Copyright notice: This dataset is made available under the Open
Database License __ (http://opendatacommons.org/licenses/
odbl/1.0/). The Open Database License (ODbL) is a license
agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others,
provided that the original source and author(s) are credited.

Link: https://doi.org/10.3897/jor.28.30143.suppl1

JOURNAL OF ORTHOPTERA RESEARCH 2019, 28(1)