Research Article |
Corresponding author: Jeffrey A. Cole ( jefreecoal@gmail.com ) Academic editor: Juliana Chamorro-Rengifo
© 2019 Jeffrey A. Cole, David H. Funk.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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
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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 species includes aggressive song as well as calling song is discussed.
aggressive song, calling song, mate recognition, Mexico, Neotropical, Sonora
Anaxipha is a speciose genus of small crickets with a worldwide distribution (
While engaged in general Orthoptera collecting in southeastern Arizona in 2013, the first author followed an unfamiliar calling song to discover an Anaxipha population. Apart from the highly 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.
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 vegetation 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.
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 T1i, 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.
The right tegmen of the above specimen was removed and slide-mounted in euparal. The tegmen was then photographed with the trinocular microscope (Fig.
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
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 mean | 24.7 ± 0.9 | 45.4 ± 4.6 | 14.3 ± 2.3 | 7.9 ± 3.2 | 22.3 ± 2.3 | 64.4 ± 9.0 | 5.7 ± 1.8 | 3.5 ± 1.2 | 117.2 ± 43.4 | 223.4 ± 184.9 | 340.6 ± 194.0 | 37.7 ± 10.8 | 6.0 ± 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 |
Field mean | 25.1 ± 0.8 | 44.1 ± 3.9 | 15.2 ± 2.1 | 7.0 ± 3.0 | 22.8 ± 2.1 | 66.8 ± 8.5 | 6.2 ± 1.7 | 2.9 ± 1.1 | 138.9 ± 42.7 | 278.2 ± 227.8 | 417.0 ± 225.4 | 37.8 ± 12.3 | 5.7 ± 0.3 |
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 | 24.1 ± 0.7 | 47.5 ± 5.1 | 12.8 ± 2.0 | 9.5 ± 3.2 | 21.3 ± 2.4 | 60.2 ± 8.6 | 4.8 ± 1.6 | 4.3 ± 0.6 | 86.9 ± 20.8 | 146.7 ± 34.9 | 233.7 ± 31.8 | 37.5 ± 9.0 | 6.3 ± 0.3 |
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 |
P | NS | 0.015 | NS | NS | NS | NS | 3.60 ×10-3 | 8.06 ×10-4 | NS | 0.010 | NS | 1.21×10-3 |
Recordings were visualized (Fig.
A. Male calling song, field recording number 140723_11, oscillogram (above) and spectrogram (below), 10 s window, 25.4°C. PTR = 2.2 s-1, PTCF = 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.
PD pulse duration
PI pulse interval
PP pulse period (=PD+PI)
PR pulse rate (=1/PP)
Pc pulse duty cycle (=PD/PP)
PT pulse train
PN number of pulses per train
PD pulse train duration
PI pulse train interval
PP pulse train period (=PTD+PTI)
PR pulse train rate (=1/PTP)
Pc pulse train duty cycle (=PTD/PTP)
PF pulse train carrier frequency
Statistical analysis was performed in R (
1 male. USA. ARIZONA: Santa Cruz County; Peña 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.
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, SING0519, SING0520, SING0521 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.F. Limón leg. (LACM).
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).
No specimens among the type series are macropterous.
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.
The population resides in a north-south trending canyon. 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 humidity 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 (
A. hyalicetra has a unique combination of morphological characters among the North American Anaxipha fauna. The 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.
l. hyalo (glassy) + cetra (a small light shield), referring to the broad, transparent male tegmina.
PR in A. hyalicetra is 45.4 s-1 (Table
The scree plot (Fig.
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 | |||
PTP | 0.98 | |||
PTdc | -0.70 | 0.50 | ||
PTD | 0.91 | 0.37 | ||
PN | 0.83 | |||
PR | -0.99 | |||
PP | 0.99 | |||
PD | 0.47 | 0.53 | 0.70 | |
PI | -0.47 | 0.46 | 0.63 | |
Pdc | 0.49 | 0.87 | ||
PTCF | -0.44 | -0.50 |
Songs were significantly different between the field and the laboratory (MANOVA, P = 9.32 × 10-3). Notably, males produced more rapid PTR (P = 3.65 × 10-3) due to shorter PTD (P = 8.06 × 10-4) in the laboratory (Table
Males sang in aggregations but did not settle into predictable chorus phase relationships, neither synchronous nor alternating (reviewed in
A trickle of new species discoveries shows that there is still much to learn about the orthopteran fauna of Arizona, despite much historical systematic attention (
Collecting in the Atascosa and Pajarito Mountains in Santa Cruz County, Arizona, failed to find this cricket outside of the type locality. 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 laboratory 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 (
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.
Data type: WAV file
Explanation note: Male calling song supplementary file.