Research Article Journal of Orthoptera Research 2020, 29(2): 137-201 Calling songs of Neotropical katydids (Orthopiera: Tettigoniidae) from Panama HANNAH MI. TER HorsTeDe!®, LAUREL B. Symes!?, SHARON J. MARTINSON!, TONY ROBILLARD?, PAUL FAURE’, SHYAM MIADHUSUDHANA2, RACHEL A. PAGE? 1 Dartmouth College, Department of Biological Sciences, 78 College Street, Hanover, NH 03755, USA. 2 Cornell University, Lab of Ornithology, Center for Conservation Bioacoustics, 159 Sapsucker Woods Road, Ithaca, NY 14850, USA. 3 Institut de Systématique, Evolution et Biodiversité (ISYEB), Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, 57 rue Cuvier, CP 50, 75231 Paris Cedex 05, France. 4 McMaster University, Department of Psychology, Neuroscience & Behaviour, 1280 Main Street West, Hamilton ON, L8S 4K1, Canada. 5 Smithsonian Tropical Research Institute, Apartado 0843-03092, Balboa, Ancon, Republic of Panama. Corresponding author: Hannah M. ter Hofstede (Hannah.ter.hofstede@dartmouth.edu) Academic editor: Klaus-Gerhard Heller | Received 7 September 2019 | Accepted 7 March 2020 | Published 4 December 2020 http://zoobank.org/392BD4E1-5F50-47B9-8C19-C83E77AD3845 Citation: ter Hofstede HM, Symes LB, Martinson SJ, Robillard T, Faure P, Madhusudhana S, Page RA (2020) Calling songs of Neotropical katydids (Orthoptera: Tettigoniidae) from Panama. Journal of Orthoptera Research 29(2): 137-201. https://doi.org/10.3897/jor.29.46371 Abstract Understanding the ecology and evolution of animal communication systems requires detailed data on signal structure and variation across spe- cies. Here, we describe the male acoustic signals of 50 species of Neotropi- cal katydids (Orthoptera: Tettigoniidae) from Panama, with the goal of pro- viding data and recordings for future research on katydid communication, evolution, ecology, and conservation. Male katydids were recorded indi- vidually using an ultrasound-sensitive microphone and high-sampling rate data acquisition board to capture both audible and ultrasonic components of calls. Calls varied enormously in duration, temporal patterning, peak frequency, and bandwidth both across and within subfamilies. We confirm previous studies showing that katydid species within the subfamily Pseudo- phyllinae produced short calls (<250 ms) at long intervals and we confirm that this is true for species in the subfamily Phaneropterinae as well. Species in the Conocephalinae, on the other hand, typically produced highly re- petitive calls over longer periods of time. However, there were exceptions to this pattern, with a few species in the Conocephalinae producing very short calls at long intervals, and some species in the Phaneropterinae producing relatively long calls (1-6 s) or calling frequently. Our results also confirm previous studies showing a relationship between katydid size and the peak frequency of the call, with smaller katydids producing higher frequency calls, but the slope of this relationship differed with subfamily. We discuss the value of documenting the diversity in katydid calls for both basic stud- ies on the ecology, evolution, and behavior of these species as well as the potential conservation benefits for bioacoustics monitoring programs. Keywords acoustic signals, bioacoustics monitoring, bushcrickets, insect communi- cation, ultrasound Introduction Understanding the ecology and evolution of animal commu- nication systems requires detailed data on signals and how they vary across species (Cocroft and Ryan 1995, Endler et al. 2005, Arnegard et al. 2010, Liénard et al. 2014, Tobias et al. 2014). In many animal taxa, males produce conspicuous acoustic signals to attract females for mating (Myrberg et al. 1986, Catchpole 1987, Gerhardt and Huber 2002, Smotherman et al. 2016), providing opportunities for both basic studies on communication and ap- plied studies through bioacoustic monitoring (Sueur 2002, Chek et al. 2003, de Solla et al. 2005, Gasc et al. 2013, Krause and Farina 2016, Grant and Samways 2016). Acoustic signal production by males is particularly conspicuous and ubiquitous in the Orthop- tera (ROmer 1998, Gerhardt and Huber 2002), making species in this taxon ideal for the types of studies mentioned above (e.g., Diwakar and Balakrishnan 2007a, Schmidt et al. 2012, Jain et al. 2014, Frederick and Schul 2016, Roca and Proulx 2016, Bailey et al. 2019). Here we describe male acoustic signals of 50 species of Neotropical katydids (Orthoptera: Tettigoniidae) from Panama, with the goal of providing data and recordings for future research on katydid communication, evolution, ecology, and conservation. Katydids, also known as bushcrickets, are a highly diverse group of insects (Mugleston et al. 2018) in which males produce acoustic signals, or calls, to attract females. In most subfamilies, males call and females walk to males by tracking the source of the sound, a be- havior called phonotaxis (Bailey et al. 1990, Schul and Schulze 2001, Guerra and Morris 2002, Kowalski and Lakes-Harlan 2011, Dutta et al. 2017). In the subfamily Phaneropterinae, however, males and females usually produce an acoustic duet, with the female produc- ing a call in a short, and species-specific, latency after the male call (reviewed in Bailey 2003, Heller et al. 2015). Phaneropterine males walk to the replying female or, in some phaneropterine species, both sexes move toward each other (Heller et al. 2015). Male katy- dids call by rubbing a plectrum on one forewing across a file on the underside of the other forewing (Bailey 1970, Montealegre-Z and Mason 2005), a form of sound generation termed stridulation. De- pending on the species, sound can be produced during wing clos- JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 138 Temporal Parameters H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Spectral Parameters 80 > Call Duration O Cc sc) a eo co) — LL Fig. 4. Photograph and calling song spectrograms of Agraecia festae. A. Female (red bump on pronotum C is tag for identifying individuals); B. and C. Spectro- JO gram (top panel) and oscillogram (bottom panel) of | .N one call (B) and two pulses from the same call (C). tL Photo credit: L. Symes. esis > oO Cc Ld) a oO co) , Li 143 with a mean of »2 s (Table 1). The peak frequency of the entire call is 40 kHz with a -20 dB frequency range spanning 32-52 kHz, giv- ing a bandwidth of 20 kHz (Table 1). The amplitude of the pulses is similar across the call, although the first few pairs of pulses are usually of a lower amplitude than the rest of the pulses in the call (Fig. 4B). Individuals will call frequently at night and are com- monly recorded in the forest on BCI. Pulses are arranged in pairs, and individual tooth strikes are visible on the oscillogram (Fig. 4B, C). The duration of the first pulse in a pair is shorter than the second pulse (Table 2). The spec- tral properties of each pulse type are the same (Table 2). This appears to be the first description of the call of this species. 80 60 40 20 it 0.04 0.06 0.08 Time (s) 0.02 Table 2. Call pulse parameters of Agraecia festae (3 individuals, 15 calls; mean + SD); n = number of pulses measured. Pulse Type (n) Pulse Duration _—_ Pulse Period (ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1(199) 14.741.1 ID he ae ev A 40.0 + 0.7 33.0 + 0.8 50.4 + 1.9 | pe = db 2 (199) 22/3 3 26.0 + 2.8 39.8 + 0.8 32.04 0.5 pl; Ree 19 LOLS JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 144 Copiphora brevirostris Stal, 1873 Fig. 5 [MNHN-SO-2019-329, -330, -331, -332, -333, -334, -335, -336]| Copiphora brevirostris is a large (1.63 + 0.31 g, n = 51), green katydid with a broad, flat, and yellow face and a powerful bite (Fig. 5A, B). Unlike many other species of Copiphora, the fastigi- um is not elongated (i.e., no cone-like structure on the top of the head). In females, the ovipositor is longer than the body (Fig. 5B). This species is known from Panama (Nickle 1992) and Colombia (Cigliano et al. 2020). The call consists of 1-4 pulses (Fig. 5C, D) with a mean call duration of 30 ms (Table 1). Pulses usually increase in ampli- tude across the call, and relatively high-amplitude wing-opening sounds can be seen before some pulses (Fig. 5D). The call has strong harmonics with the fundamental (~16 kHz) and first har- monic (~33 kHz) produced at similar amplitudes (Fig. 2). The first harmonic usually has more energy than the fundamental, but in some calls the fundamental can be the same or slightly higher in amplitude than the first harmonic. The peak frequency of the har- monic is 33 kHz with a -20 dB range spanning ~28-35 kHz, giving A C H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE a bandwidth of 7 kHz (Table 1). Males call very rarely and tend to call more frequently after midnight. Pulse durations are typically 6-9 ms (Table 3), although pulse durations are highly variable and can range from 2-12 ms. The second pulse is often slightly longer than the first pulse. Pulse peri- ods are ~15 ms (Table 3). The spectral properties of the individual pulses are very similar to each other and the entire call (Table 3). The peak frequency of the fundamental is ~16 kHz, and the first harmonic is ~32 kHz, with a -20 dB range spanning 29-34.5 kHz, giving a bandwidth of 5.5 kHz (Table 3). The bandwidth reported here is just for the first harmonic. The fundamental was usually of a lower amplitude than the first harmonic, but the difference in amplitude was highly variable across calls and pulses. Each pulse is frequency modulated, either sweeping from higher to lower fre- quencies (~34 to 30 kHz) or shaped like an upside-down U (rang- ing from ~33 up to 35 and down to 30 kHz; Fig. 5D). Calls of this species were previously described by Belwood and Morris (1987), Belwood (1988a), Morris et al. (1994), Falk et al. (2015), and Symes et al. (2016). In addition to acoustic signals, both males and females produce vibrational signals (described in Belwood 1988a). — 80 60 40 20 Frequency (kHz Oo & DD oO CO G&G © G& Frequency (kHz) >) 0.00 0.02 0.04 0.06 0.08 Time (s) Fig. 5. Photographs and calling song spectrograms of Copiphora brevirostris. A. Male (photo credit: H. ter Hofstede); B. Female (photo credit: C. Wilson); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Table 3. Call pulse parameters of Copiphora brevirostris (8 individuals, 115 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) LG 5:) Gt 1 SLA tel 2 29.8 + 1.7 34.741.2 5.0 + 2.3 2 (113) 8.4+1.5 Ie Cae al Io) 32.7 + 1.0 29) Le 41e5 34.241.2 5.14+1.8 3 (55) 7.0 + 1.4 16.0+ 1.4 BL 24 1.2 26.6 + 1.8 34.04 2.4 7.4 + 3.6 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Eppia truncatipennis Stal, 1875 Fig. 6 [MNHN-SO-2019-611, -642, -646] Eppia truncatipennis is a large (1.18 + 0.15 g, n = 2), mottled, brown katydid with abruptly truncated wings, a black face, and red mouthparts (Fig. 6A). This species was redescribed by Naskrecki (2000). It is known from southern Mexico, Costa Rica, Panama, and Colombia (Cigliano et al. 2020). The call consists of a sequence of “chirps” (term used in Naskrecki 2000) composed of 10-12 pulses produced with al- most no silence between them (Fig. 6B, C). Chirps are produced at very regular intervals, with a chirp period of ~280 ms (Table 1). Sequences of chirps are produced for long periods of time (11.3- 44.7 s with a mean of 21.3 s; Table 1). The peak frequency of an entire sequence of chirps is ~50 kHz with a -20 dB frequency range spanning ~37-63 kHz, giving a bandwidth of ~26 kHz (Table 1). The chirps are all very similar in their temporal and spectral properties. Chirp durations are 114.4 + 9.1 ms (3 individuals, 7 Fig. 6. Photograph and calling song spectrograms of Eppia trun- catipennis. A. Male (photo credit: L. Symes); B. and C. Spectro- gram (top panel) and oscillogram (bottom panel) of the start of one call (B) and one chirp from the same call (C). 145 11 calls, 110 chirps). There is always an even number of pulses within a chirp, usually 10 or 12 (mean 11.4 + 1.1). Pulse dura- tions within a chirp range from ~7-14 ms. It is possible that sound is produced both during the wing opening and wing closing movements, resulting in pulses that vary in amplitude but have almost no silence between them (Fig. 6C). High-speed video of males singing would be helpful in confirming that this is the mechanism responsible for these chirps that lack silence between pulses. The peak frequency of the chirps is 49.7 + 2.5 kHz with a -20 dB frequency range spanning 37.5 + 1.2-63.0 + 8.3 kHz, giving a bandwidth of 25.5 + 8.1 kHz (3 individuals, 11 calls, 110 chirps). There is also significant energy at 10-12 kHz, and, in some calls, this frequency range is the same or greater in amplitude than the typical peak frequency of ~50 kHz. Calls of this species were previously described by Naskrecki (2000), but they were recorded at a lower sampling rate that did not capture the higher frequencies described here. 80 60 40 20 Frequency (kHz) Frequency (KHz) © 0.20 0.15 0.10 Time (s) 0.05 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 146 Erioloides longinoi Naskrecki & Cohn, 2000 Fig. 7 [MNHN-SO-2019-649, -650, -651] Erioloides longinoi is a small (0.36 + 0.07 g, n = 8), cylindrical, green katydid with blue mouthparts, red and yellow markings on the ventral surface of the abdomen, and an agile bite (Fig. 7A, B). This species is known from Mexico, Costa Rica, and Panama (Cigliano et al. 2020). The call consists of a rapid series of pulses (Fig. 7C, D) with a total call duration ranging from 1.0-1.9 s and a mean of 1.4 s (Table 1). The peak frequency of the entire call is 30 kHz with a -20 dB frequency A C H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE range spanning 25-37 kHz, giving a bandwidth of 12 kHz (Table 1). The amplitude of the pulses gradually increases for the first 10-15 pulses and then remains constant for the rest of the call (Fig. 7C). The pulses in the call are all very similar in their temporal and spectral properties. Pulse durations are 4.4 + 0.7 ms (3 individuals, 41 calls, 410 pulses) and pulse periods are 8.9 + 0.5 ms. The peak frequency of the pulse is 30.2 + 1.9 kHz with a -20 dB frequency range spanning 26.6 + 2.9-38.2 + 4.3 kHz, giving a bandwidth of 11.6 + 7.2 kHz. Each pulse is frequency modulated, sweeping from ~32 to 28 kHz (Fig. 7D). This appears to be the first description of the call of this species. 80 iy =< 60 oO ) on) Time (s) Fig. 13. Photographs and calling song spectrograms of Anapolisia colossea. A. Male (photo credit: C. Kernan); B. Female (photo credit: H. ter Hofstede); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call (C) and one pulse from the same call (D). Table 10. Call pulse parameters of Anapolisia colossea (9 individuals, 116 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (116) 13.441.4 TOF 9.6 + 0.7 26.3 54.0 16.7+0.8 2 (116) 11.64 1.4 316.02 ThiZ 19:3 20:6 5.80.9 26.4 + 1.0 LG. eer, 2 3 (116) 11.341.5 394.0 + 18.7 19.8 + 0.7 9.7+0.8 26.441.1 16.741.1 4 (112) 11.641.5 423.54+19.5 19.8 + 0.6 9.6 + 0.7 26.5 + 1.0 16,9:2°1.1 5 (84) 11.341.9 435.7 425.1 199° £025 9.8+0.9 26,3:-4.10 16.6 + 1.3 6 (59) THO. 221 460.9 + 23.3 19.8 40.5 9.6 + 1.0 263 4c 16.6213 7 (35) TZ pel: 516.1 + 69.6 19-3 °+)4 2 9.841.5 2G 6-e le 16.7 42.5 8 (10) Ede ol 465.9 + 96.6 19.4 + 0.7 9.8 401.7 26.8 + 1.4 17.0 + 3.0 9 (3) 13.541.1 519.2 + 15.4 20.14+0.5 952,00 27.4+0.5 18.2+0.5 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 154 Anaulacomera furcata Brunner von Wattenwyl, 1878 Fig. 14 [MNHN-SO-2019-232, -233, -234] Anaulacomera furcata is a very small (0.14 + 0.04 g, n = 43), green katydid with narrow wings, a solid green face, three black spots on the posterior edge of the pronotum, light yellow stripes along the dorsal margins of the pronotum, and male cerci that are forked, having two branches at the end (Fig. 14A, B). This A C H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE species is known from Costa Rica, Panama, and Colombia (Cigliano et al. 2020). The call consists of two short pulses of equal amplitude pro- duced ~20 ms apart (Table 11; Fig. 14C, D). The peak frequency of the entire call is »29 kHz, with a -20 dB range spanning ~24-36 kHz, giving a bandwidth of ~12 kHz (Table 1). The two pulses have similar temporal and spectral properties (Table 11). This appears to be the first description of the call of this species. — 80 N eo?) © 40 Frequency (KH Frequency (kHz) Time (s) Fig. 14. Photographs and calling song spectrograms of Anaulacomera furcata. A. Male; B. Female; C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Photo credit: C. Kernan. Table 11. Call pulse parameters of Anaulacomera furcata (3 individuals, 53 calls; mean + SD); n = number of pulses measured. Pulse number (n) Pulse Duration _—_ Pulse Period (ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (53) 0.8+0.1 pls Ee eal eo 24.84 1.1 35.0 + 0.7 10.2 + 1.3 2 (53) Ot Oy1 20 Joel 9 29.84 O-F 25.0 + 0.7 35.8 + 0.3 10.8 + 0.7 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE 155 Anaulacomera “goat” temporary species name “goat” due to the unique eye patterning. Fig. 15 The calls recorded from these individuals are all the same and can be readily distinguished from the other species of Anaulacomera we Anaulacomera “goat” is a very small (0.16 + 0.02 g n= 12), collected in Panama. green katydid with narrow wings, a dark line through the eye, and The call consists of a single pulse with a duration ~2 ms (Ta- a dark brown stridulatory area in males (Fig. 15A, B). We were ble 1; Fig. 15C, D). The peak frequency of the call is ~27 kHz, with not able to identify these individuals to species and provide the a-20 dB range spanning 23-33 kHz, giving a bandwidth of 10 kHz. A C 80 N =< 60 > 2 40 g ae = 0 0.0 05 1.0 1.5 2.0 B D_ N a x > O a ®o Fa | oO © LL 0.00 0.02 0.04 0.06 0.08 0.10 Time (s) Fig. 15. Photographs and calling song spectrograms of Anaulacomera “goat”. A. Male (photo credit: C. Wilson); B. Female (photo credit: C. Kernan). Inset shows dark line through eye; C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at dif- ferent time scales. JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 156 H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Anaulacomera “ricotta” identify these individuals to species and provide the temporary spe- Fig. 16 cies name “ricotta” due to the unique white mottling on the body. The call consists of two short pulses of equal amplitude pro- Anaulacomera “ricotta” is a very small (0.12 + 0.02 gn = 7), duced ~60 ms apart (Table 12; Fig. 16C, D). The peak frequency green katydid with narrow wings, a white and green mottled body, of the entire call is ~34 kHz, with a -20 dB range spanning ~29-39 and male cerci that are forked, having two branches at the end, one_ kHz, giving a bandwidth of ~10 kHz (Table 1). The two pulses of which ends in a spiral coil (Fig. 16A, B). We were not able to have similar temporal and spectral properties (Table 12). A C — 80 60 si | 20 Frequency (kHz) oO 0 Frequency (kHz) Time (s) Fig. 16. Photographs and calling song spectrograms of Anaulacomera “ricotta”. A. Male (photo credit: H. ter Hofstede); B. Female (photo credit: C. Kernan); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Table 12. Call pulse parameters of Anaulacomera “ricotta” (3 individuals, 16 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (16) TO 092: 36 a A pee 29532201 38.0 + 0.2 8.4+1.9 2 (16) LOSE 0.2: 580.4. 1.9 33.9 + 1.5 30.1+ 1.6 38.0 + 0.1 Cot LS JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE 157 Anaulacomera spatulata Hebard, 1927 The call consists of two short pulses of equal amplitude pro- Fig. 17 [MNHN-SO-2019-238, -239, -240] duced ~40 ms apart (Table 13; Fig. 17C, D). The peak frequency of the entire call is ~25 kHz, with a -20 dB range spanning ~22- Anaulacomera spatulata is a small (0.30 + 0.08 g, n=129), green 29 kHz, giving a bandwidth of ~7 kHz (Table 1). The two pulses katydid with very narrow wings. Males have a dark brown stridula- have similar temporal and spectral properties (Table 13). tory area and spatulate cerci (Fig. 17A, B). This species is known This appears to be the first description of the call of this species. from Panama, Colombia, and Suriname (Cigliano et al. 2020). A C (oe) i) NO o>) © i) Frequency (kHz) pS © oO © a ee NO ee) Frequency (kHz) iS i) 0.00 0.02 0.04 0.06 0.08 0.10 Time (s) Fig. 17. Photographs and calling song spectrograms of Anaulacomera spatulata. A. Male; B. Female; C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Photo credit: C. Wilson. Table 13. Call pulse parameters of Anaulacomera spatulata (3 individuals, 59 calls; mean + SD); n = number of pulses measured. Pulse number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (59) 134.2 24.6 + 3.2 22.2 + 3.3 29.3 + 2.5 ipa a: sal a 2 (59) 1.4+0.3 41.44 2.3 24.3 + 3.7 22.0 + 3.3 29.3 + 2.0 7.341.4 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 158 H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Anaulacomera “wallace” The calls recorded from these individuals are all the same and can Fig. 18 be readily distinguished from the other species of Anaulacomera that we collected in Panama. Anaulacomera “wallace” is a very small (0.22 + 0.05 g, n = 28), The call consists of three short pulses of equal amplitude pro- green katydid with narrow wings, a green and white mottled face, duced ~16 ms apart (Table 14; Fig. 18C, D). The peak frequency eyes that are half green and half white, and highly reduced cerciin of the entire call is 25 kHz, with a -20 dB range spanning ~20- males (Fig. 18A, B). We were not able to identify these individuals 31 kHz, giving a bandwidth of ~11 kHz (Table 1). The three pulses to species, and we provide the temporary species name “wallace.” have similar temporal and spectral properties (Table 13). A C 80 60 40 20 H Frequency (KHz) 0 D> So © NO © Frequency (kHz) KK © Time (s) Fig. 18. Photographs and calling song spectrograms of Anaulacomera “wallace”. A. Male; B. Female; C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Photo credit: C. Kernan. Table 14. Call pulse parameters of Anaulacomera “wallace” (4 individuals, 19 calls; mean + SD); n = number of pulses measured. Pulse number (n) Pulse Duration Pulse Period (ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (19) 12 O12. O29 0 aad Nk 21.4 4+ 0.7 29. T2.6 Mesto Stl: 2 (19) 10s). 16.5 + 0.7 25.0 + 0.6 21.5 + 1.3 29.7 43.4 8.2 + 4.3 3 (18) 0.9+0.2 1622 611 25.1 + 0.9 21.7+1.2 29.9+3.4 8.2 44.2 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE 159 Arota festae (Griffini, 1896) Fig. 19 [MNHN-SO-2019-241, -246, -247, -248, -249, -250, -251, -252, -253, -254| Arota festae is a mid-sized (0.98 + 0.15 g, n = 34), light green katydid with broad, rounded tegmina that cover nearly all of the hindwings (<3 mm visible beyond the apex of the tegmina) (Fig. 19A, B). This species is known from Panama, Colombia, and Suriname (Cigliano et al. 2020). The call consists of a series of 7-10 (mean: 8) short pulses (Fig. 19C, D) with a total call duration ranging from ~15-28 ms and having a mean of 21 ms (Table 1). The peak frequency of the entire call is ~13 kHz with a -20 dB frequency range spanning A C Frequency (kHz) Frequency (kHz) ~8-19 kHz, giving a bandwidth of ~11 kHz (Table 1). The ampli- tude of the pulses varies across the call. In most cases, the pulses increase in amplitude (Fig. 19D), but sometimes they increase and then decrease in amplitude. Pulse durations are short and increase slightly in duration over the call, whereas pulse period stays constant across the call (Table 15). The peak frequency of each pulse increases across the call (Table 15). The low and high frequencies of each pulse also increase across the call, with bandwidths ranging from 5-9 kHz, depending on the pulse (Table 15). The calls of this species were previously described by Symes et al. (2016). 80 60 40 20 | Time (s) Fig. 19. Photographs and calling song spectrograms of Arota festae. A. Male; B. Female; C. and D. Spectrogram (top panel) and oscil- logram (bottom panel) of one call at different time scales. Photo credit: C. Wilson. Table 15. Call pulse parameters of Arota festae (10 individuals, 83 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (83) 8a. 2 8.7 + 0.3 6.7 + 0.6 15.8 + 2.2 Paige ng 2 (83) 1.0%. O32 3.0+0.5 8.8 + 0.2 7.34 0.4 14.3 + 2.2 7.0 + 2.4 3 (83) 10+ 0:2 2.7+0.4 9.2+0.3 7.7 +0.2 13.2: 42.2 5.542.3 4 (83) 1.0 + 0.2 2.7 + 0.4 9.7+0.4 8.0 + 0.3 a7 S21 5.7+2.0 5 (83) 1.00+0.1 2.7+0.5 10.6 + 0.5 8.4 + 0.3 15.0+2.2 6.642.1 6 (83) 0.9-4-0.1 2.8 + 0.4 11.9+ 1.0 9.1 + 0.6 17.142.4 8.0 + 2.4 7 (83) 2 0.3 2 eB O35 13.5+0.8 10.4 + 1.3 17.9 41.5 7.5 42.0 8 (62) ARS 20.2 2.9 +0.3 14.4+0.9 10.9 + 1.0 19:3 £:2.2 8.442.5 9 (26) 1440.2 2.8+0.5 14.3 + 0.8 11.44 0.4 LS. 4 12 6.7 + 1.0 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 160 H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, $8. MADHUSUDHANA AND R.A. PAGE Arota panamae (Hebard, 1927) Fig. 20 [MNHN-SO-2019-255, -256, -288, -289, -290, -291, -292, -293, -294, -295] Arota panamae is a mid-sized (0.57 + 0.11 g, n = 68), light green katydid with broad wings and hindwings that extend >3 mm be- yond the apex of the tegmina (Fig. 20A, B). This species is known from Panama and Colombia (Cigliano et al. 2020). The call consists of a series of 3-6 (mean: 5) short pulses (Fig. 20C, D) with a total call duration ranging from ~8-25 ms and having a mean of ~15 ms (Table 1). The peak frequency of the entire call is ~24 kHz with a -20 dB frequency range span- A C w O Frequency (kHz) Frequency (kHz) ning ~15-33 kHz, giving a bandwidth of ~18 kHz (Table 1). The amplitude of the pulses varies across the call. The pulses either increase in amplitude or they increase and then decrease in ampli- tude across the call (Fig. 20D). Pulse durations are short and increase over the call, where- as pulse period stays constant (Table 16). The peak frequency of each pulse increases across the call (Table 16). The low and high frequencies of each pulse also increase across the call, with bandwidths ranging from 10-20 kHz depending on the pulse (Table 16). The calls of this species were previously described by Falk et al. (2015) and Symes et al. (2016). 80 60 40 20 0.00 0.02 0.04 0.06 Time (s) Fig. 20. Photographs and calling song spectrograms of Arota panamae. A. Male (photo credit: C. Wilson); B. Female (photo credit: M. Ayres); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Table 16. Call pulse parameters of Arota panamae (10 individuals, 156 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (156) 0.4+0.1 12.1414 7540.9 25.2 44.1 17.8 + 4.0 2 (156) 0.5+0.1 3.3 +038 14.3 42.5 De ae ae 4 26,9:+'3.3 17.1 43.1 3 (156) 0.5+0.1 3.6+0.5 18:2 +:3.0 Pe ard 28.9 + 3.5 15.74+3.4 4 (151) 0-7-4052 3.6 + 0.4 DS act Se: 16.3 + 3.0 31.6 + 2.5 15.3 + 3.2 5 (118) 1.0 + 0.3 3.6+0.5 263054) ex 20). 222324; 34.743.5 14.4+5.1 6 (48) Les 0.2 3.3 + 0.3 27.9 + 1.0 23.0 + 1.5 34.7 + 3.6 11.7 + 4.7 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE 161 Ceraia mytra Grant, 1964 Fig. 21 [MNHN-SO-2019-302, -303, -304, -305, -306, -307, -308, -309, -310] Ceraia mytra is a large (1.30 + 0.28 g, n = 31), green katydid with narrow wings, reddish cerci, and reddish-purple hindlegs (Fig. 21A, B). This species is known from Costa Rica, Panama, and Colombia (Cigliano et al. 2020). The call consists of a series of 6-13 (mean: 10) short pulses (Fig. 21C, D) with a total call duration ranging from ~40-96 ms and having a mean of ~76 ms (Table 1). The peak frequency of the entire call is ~11 kHz with a -20 dB frequency range spanning ~7-20 kHz, A C. .. 86 N Frequency (kH & © Frequency (kHz) aa © giving a bandwidth of ~13 kHz (Table 1). The amplitude of the pulses varies across the call. The pulses either increase in amplitude (Fig. 21D) or they increase and then decrease in amplitude across the call. The pulses in the call are all very similar in their temporal and spectral properties (Table 17). Pulses sometimes have silent gaps within them, making it look like there are two shorter pulses sepa- rated by a very short silent period (e.g., pulse six in Fig. 21D). The peak frequency of each pulse decreases slightly across the call (Ta- ble 17). The low and high frequencies of each pulse also decrease slightly across the call (Table 17). The calls of this species were previously described by Falk et al. (2015). o>) © NO i) =) CO i) (ep) © NO a 0.00 0.02 0.04 0.06 0.08 0.10 Time (s) Fig. 21. Photographs and calling song spectrograms of Ceraia mytra. A. Male (photo credit: H. ter Hofstede); B. Female (photo credit: L. Symes); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Table 17. Call pulse parameters of Ceraia mytra (9 individuals, 71 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (71) 1440.5 AP Oh LED EOD 17.441.7 9.542.2 Dita) 1.6 + 0.6 7.94+1.2 11.9 + 1.0 8.0 + 1.1 17.7 41.9 9.742.5 S371) 1740.3 7.6 + 0.6 T1609) 7.8 + 1.0 17.441.4 OM ote mel 4 (71) 15+ 0.3 8.0 + 0.7 1S ORF C6 EO 16.9 4:1.3 Deel 5 (71) 134073 8.0+0.5 11.1+0.9 8.0 + 0.7 16.141.4 > oo Ae ol Wg 6 (71) 1440.4 8.0+0.5 11.1+0.8 7.6 + 0.6 16.5 41.7 8.9 + 1.6 7 (68) 1.6 + 0.6 8.1+40.5 10.9 + 0.7 7.8 + 0.8 Na DBs eae 3: + 228 8 (63) 1.64 0.4 8.4 + 0.6 10. #150, 7.5 + 0.7 16.841.5 a ae oe 9 (48) 18.0.6 8.5+0.8 9.8+0.8 tel, £06 15.3 42.3 8.24 2:8 10 (16) 1.140.4 Soa e035 9.2+0.8 6.7 + 0.4 14.64+1.2 Pi Me as JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 162 H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S. MADHUSUDHANA AND R.A. PAGE Chloroscirtus discocercus Rehn, 1918 Fig. 22 [MNHN-SO-2019-311, -312, -313, -314, -315, -316, -317, -318, -319, -320, -321, -322] Chloroscirtus discocercus is a mid-sized (0.59 + 0.22 g,n=79), green katydid with narrow wings, sometimes with light yellow stripes along the dorsal margins of the pronotum (Fig. 22A, B). This species is known from Costa Rica, Panama, and Colombia (Cigliano et al. 2020). The call consists of a series of 4-8 (mean: 6) short pulses (Fig. 22C, D) with a total call duration ranging from ~85-173 ms and having a mean of ~140 ms (Table 1). The peak frequency of the entire call is ~20 kHz with a -20 dB frequency range spanning ~11-26 kHz, giving a bandwidth of ~15 kHz (Table 1). Pulses are A C Frequency (kHz) fairly constant in amplitude, but the first or last pulse is often of a lower amplitude than the rest of the pulses. The first pulse in the call is longer in duration than the oth- er pulses, which are similar in duration (Table 18). The first pulse period is also longer in duration than the other periods, which are similar in duration (Table 18). The pulses in the call are all similar in their spectral properties (Table 18). Pulses are usually frequency-modulated, with the first half consisting of a constant frequency component at ~13 kHz, with visible tooth strikes in the oscillogram, followed by a frequency-modulated sweep up to ~20 kHz, often followed by a steep vertical tail at the end (Fig. 22D). The calls of this species were previously described by Symes et al. (2016). 80 60 40 20 oO © Oo Oo NO -) Frequency (kHz) & =) 0.10 Time (s) 0.00 0.05 Fig. 22. Photographs and calling song spectrograms of Chloroscirtus discocercus. A. Male; B. Female; C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Photo credit: H. ter Hofstede. Table 18. Call pulse parameters of Chloroscirtus discocercus (12 individuals, 157 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration _—‘ Pulse Period (ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (157) 13.7+2.5 1 Qeeeee7 10.0-4-1 .2: 26.9 42.5 16.9 +2.8 2 (157) Se foke 2) 30.0 + 3.1 18.8 + 2.5 1d Dee) 26.1 43.5 13.9 + 4.4 35157) 7442.1 24.2 + 1.3 18.5 + 2.8 13.0 + 2.4 25.5 43.1 12.4445 4 (157) Fale 1.8 22.7 £7120 As al Roches 13. D222 24.8 + 3.1 11.7 + 4.3 5 (156) 7.341.7 22.1 41.1 19.0 + 2.2 13.142.1 24.5+2.5 11.4 + 3.8 6 (148) Ais eal hs 2273. SE ANG 19.24+2.4 12.74 1.5 25.5 + 2.6 128356 7 (60) 7.64 1.4 Masset 1S 18.5 + 2.2 12.241.5 25.442.1 jh Ml Ibe eo JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE 163 Dolichocercus latipennis (Brunner von Wattenwyl, 1891) Fig. 23 [MNHN-SO-2019-344, -345, -346] Dolichocercus latipennis is a very small (0.21 + 0.03 g, n = 40) and mostly brown katydid with hind wings that extend signifi- cantly beyond the tips of the sharply-angled and narrow tegmina, reminiscent of a wind-dispersed seed (Fig. 23A, B). The dorsal sur- face of the abdomen is bright green. This species is known from Costa Rica, Panama, and Colombia (Cigliano et al. 2020). The call consists of a series of 14-17 (mean: 16) short pulses (Fig. 23C, D) with a total call duration ranging from ~282-370 ms and having a mean of ~330 ms (Table 1). The peak frequency of the entire call is ~26 kHz with a -20 dB frequency range span- ning ~21-32 kHz, giving a bandwidth of ~11 kHz (Table 1). Pulses usually increase in amplitude over the call with the last two pulses then decreasing in amplitude (Fig. 23C). The pulses increase in duration across the call (Table 19). The pulse periods are similar in duration (Table 19). The pulses in the call are all similar in their spectral properties (Table 19). Each pulse is ashort, downward frequency modulated sweep from ~28-21 kHz (Fig. 23D). In some calls, some pulses have silent gaps within them. This appears to be the first description of the call of this species. A C ~— 80 = < 60 > 2 40 g o 20 LL D Frequency (kHz) Table 19. Call pulse parameters of Dolichocercus latipennis (3 individuals, 19 calls; mean + SD); n = number of pulses measured. Fig. 23. Photographs and calling song spectrograms of Dolichocercus latipennis. A. Male; B. Female; C. and D. Spectrogram (top pan- el) and oscillogram (bot- tom panel) of one call (C) and nine pulses from the same call (D). Photo cred- it: H. ter Hofstede. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (19) 1.0+0.5 DEM DD 22 + OO 32,0 $21 8.84 1.4 2( 19) 1.0 + 0.4 22.5+0.9 26.5 + 1.8 DIA se 13 32.04 1.4 ve Rp pee el S769) 1.1+0.3 22.9 4 led 26.1 +131 22.9 + 0.7 31:3; 4 1:2 8.4 + 1.0 4 (19) 1.1+0.3 Zoe BALD 26.0 + 0.9 2S Ot OD 31 let OD 8.1+0.8 5b D)) 1.14+0.3 232, £:09 26.1 + 0.8 22.8 + 0.8 <9 ie lie sad $.3:£0.6 6 (19) 1.3 + 0.3 23.04 1.1 26.0 + 0.6 22.6 + 0.6 31.1 +0.9 8.5 + 0.6 Fi eGS>) 1.4+0.2 23.14 1.0 25.9+0.4 22.9 + 0.7 30.9 + 0.7 8.0+0.5 8 (19) 1.5+0.3 22.9. £018 26.1 + 0.6 22.9 +0.5 30.8 + 1.1 Co £2 9 (19) 1.64+0.2 22.8 + 1.0 26.0 + 0.6 22.6 + 0.4 30.44 1.1 ZOE 1S 10 (19) 1.8+0.5 D229 + 0.¢ 26.0 + 0.6 22.4+0.4 30.6 + 1.3 8.241.5 11 (19) 1.9+0.5 22.8°4 Oe7 25.9 + 0.6 22.3 + 0.4 30.74+1.5 8.441.5 12° (19) 2.2+0.8 22 SIO 26.1 + 1.0 AMT + OD 30:38: 1.9 Of £19 13: (19) 23. +:0:8 21.8 + 1.5 25.9 + 0.9 21.6 + 0.1 30.8 + 1.8 ele eal Us 14 (19) 2.4+0.8 OA leis De iad nd 25.5+0.8 21.4 + 0.0 30.9 + 2.3 9.542.3 ES E(ahG) 2.6 + 1.0 19.9.4 270 25.0 + 0.4 21.0 + 0.2 31.54+2.2 10.5 + 2.4 16 (13) 31 0.3 Li. 0.6 23.8 41:0 AEs SA Le oe Jo. 2 14.0 + 2.9 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 164 H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Ectemna dumicola Saussure & Pictet, 1897 Fig. 24 [MNHN-SO-2019-347, -348, -608, -609, -610] Ectemna dumicola is a mid-sized (0.66 + 0.11 g, n = 10), green ka- tydid with narrow wings and a thin white and purple stripe running from the eyes, across the lateral surface of the pronotum, and con- tinuing on the leading edge of the tegmen (Fig. 24A, B). This species is known from Panama and Colombia (Cigliano et al. 2020). The call consists of a series of 3-14 (mean: 10) short pulses (Fig. 24C-F) with a total call duration ranging from ~123-678 ms and having a mean of ~466 ms (Table 1). The peak frequency of the entire call is ~15 kHz with a -20 dB frequency range spanning ~10-26 kHz, giving a bandwidth of ~16 kHz (Table 1). Pulses usu- ally increase in amplitude over the call with the last two pulses often decreasing in amplitude (Fig. 24C-F). The pulses increase slightly in duration from across the call (Table 20), whereas pulse periods decrease over the call (Table 20). The pulses in the call are all similar in their spectral proper- ties (Table 20). This appears to be the first description of the call of this species. E ) S A — 80 + =< 60 oS < 40 S ceed LL B 60 Frequency (kHz BES oO Frequency (kHz) 0.0 0.2 Time (s) Frequency (kHz) 0.4 0.6 Time (s) Fig. 24. Photographs and calling song spectrograms of Ectemna dumicola. A. Male (photo credit: L. Symes); B. Female (photo credit: H. ter Hofstede); C. and D. Spectrogram (top) and oscillogram (bottom) of one call with 11 pulses at different time scales; E. and FE. Spec- trogram (top) and oscillogram (bottom) of one call with three pulses at different time scales. Table 20. Call pulse parameters of Ectemna dumicola (5 individuals, 83 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (83) 5.5 + 0.6 14.04+1.4 10.0 + 0.8 24.9 +2.2 14.9 + 2.3 2 (83) 6.207 60:0°+3.7 14.74 1.8 10.3 + 1.1 24.8 +3.5 14.4 + 3.7 3 (83) 6.2+0.8 56.1 + 2.3 14.84+1.9 10.4+1.0 24.9+3.4 14.5 + 3.6 4 (81) 5.9 + 0.8 57.4+1.6 14.9 + 2.0 10.4+1.0 24.8 + 3.3 14.443.5 5 (81) 6.0 + 0.9 56.5 + 3.1 15.0 + 2.4 10.4 + 0.9 24.7 + 3.2 14.3 + 3.4 6 (81) 6.44+1.2 54.6 + 3.7 15.3.4 2.5 10.5+0.8 25.6 + 1.7 15.2 41.6 7 (80) 6.841.5 52.9 + 4.6 15.2 42.3 10.4 +0.8 26.0 + 1.6 15.64 1.4 8 (70) 6.9+1.3 48.54+4.4 15.3 +2.5 10.3 + 0.7 26.04 1.4 15.74+1.4 9 (58) 7.4+0.9 41.8+4.3 15.9 + 2.6 10.3 + 0.8 26.0+ 1.2 15.7 41.3 10 (53) 7.0 + 1.0 3912: B36 15.94+2.9 1G 2+ 09 25.7 41.1 15.441.5 11 (27) 2 Ore Os9 38.4 + 2.2 15.7+2.9 10.3 + 1.0 24.641.5 14.3 42.1 L212) 6.9+1.3 38.44 1.7 14.14+0.2 10.7 + 0.6 23.5 44.3 12.9+4.9 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE 165 Euceraia atryx Grant, 1964 Fig. 25 [MNHN-SO-2019-662, -663, -666] Euceraia atryx is a mid-sized (0.67 + 0.15 g, n = 56), green katy- did with narrow wings, orange tarsi, and a pronotum that is yellow on the sides and brown on the dorsal surface (Fig. 25A, B). This species is known from Costa Rica, Panama, Colombia, and Suri- name (Cigliano et al. 2020). The call consists of a series of 11-17 (mean: 14) short pulses (Fig. 25C, D) with a total call duration ranging from ~0.7-1.7 s and having a mean of ~1.1s (Table 1). The peak frequency of the entire call is ~13 kHz with a -20 dB frequency range spanning ~11-16 kHz, giving a bandwidth of ~5 kHz (Table 1). Pulses usually increase and then decrease in amplitude over the call (Fig. 25C, D). Pulse durations are short, and both pulse durations and pulse periods are consistent across the duration of the call (Ta- ble 21). The pulses in the call are all similar in their spectral properties (Table 21). This appears to be the first description of the call of this species. A Crm N ui nd > S) c 0) a oy ® — Le D Frequency (kHz) 80 60 40 20 0.00 0.10 0.20 0.30 Time (s) Fig. 25. Photographs and calling song spectrograms of Euceraia atryx. A. Male (photo credit: H. ter Hofstede); B. Female (photo credit: C. Kernan); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call (C) and three pulses from the same call (D). Table 21. Call pulse parameters of Euceraia atryx (3 individuals, 14 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (14) 2.2+0.5 12.84+1.4 Tie Oe) 15.64 1.9 4.5+1.0 2 (14) 2.7+0.2 91.0 +-19.1 1s 5 le ee NO >) 0.00 0.02 0.04 0.06 0.08 0.10 Time (s) Fig. 38. Photographs and calling song spectrograms of Pycnopalpa bicordata. A. Male; B. Female; C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Photo credit: H. ter Hofstede. Table 31. Call pulse parameters of Pycnopalpa bicordata (3 individuals, 14 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (14) 1.3+0.2 28.2 + 3.6 23.1+41.9 34.745.3 11.6 + 3.5 2 (14) 0.9 + 0.4 7.5 40.3 26.5 + 1.6 23M +711 31.9 +3.4 8.8 + 2.7 3 (14) 1.1+04 7.5 +0.6 26.1 41.2 23.8:+ 1.0 30.5 + 2.8 Gum hed: 4 (14) 1540.7 7.9 + 0.6 26.2 £058 23.3>451.3 30.3 + 1.8 7.0 + 0.6 5 (10) 1.8+0.1 7.8+1.8 26.3 + 0.6 23.4 + 0.7 30.5 + 1.0 vex Bes te JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Steirodon stalii (Brunner von Wattenwyl, 1878) Fig. 39 [MNHN-SO-2019-1803, -1804, -1805, -1806, -1807, -1808, -1809, -1810, -1811, -1812, -1813] Steirodon stalii is a very large (4.16 + 0.49 g, n = 22), green katydid with yellow-tipped ridges along the edge of the pronotum (Fig. 39A). This species is known from Nicaragua, Costa Rica, Pan- ama, Colombia, and Brazil (Nickle 1992, Cigliano et al. 2020). The call consists of a series of three pulses (Fig. 39B, C) with a total call duration ranging from 187-247 ms and having a mean A B Fig. 39. Photograph and calling song spectrograms of Steirodon stalii. A. Male; B. and C. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Photo credit: C. Wilson. @) Frequency (KHz) Frequency (kHz) 179 of ~209 ms (Table 1). The peak frequency of the entire call is »19 kHz with a -20 dB frequency range spanning ~13-24 kHz, giving a bandwidth of ~11 kHz (Table 1). Pulse amplitudes usually in- crease across the call (Fig. 39C). Pulse durations and pulse periods are quite consistent across the call (Table 32). The pulses in the call are all similar in their spectral properties (Table 32). This appears to be the first description of the call of this species. 80 60 40 20 ‘ii oe 0.0 0.1 0.2 Time (s) 0.3 0.4 Table 32. Call pulse parameters of Steirodon stalii (10 individuals, 92 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (92) 4.84+2.3 19.041.5 ISG 132. 24.2 +120 10.6 + 1.5 2 (92) 4.8+2.0 94.5 + 7.6 18.9+1.1 13.5+1.0 24,3 1.1 10.8 + 1.6 3 (92) 5.6 + 2.3 107.6 + 7.8 LO? hdl 7: 1322 4 TO) 24.441.3 Le 1? JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 180 Viadana brunneri Cadena-Castaneda, 2015 Fig. 40 [MNHN-SO-2019-1823, -1824, -1825, -1826, -1827, -1828, -1829, -1830, -1831, -1832, -1833] Viadana brunneri is a small (0.38 + 0.07 g, n = 70) and delicate green katydid with broad wings that give a strong impression of a single new leaf (Fig. 40A). This species was described by Gorochov and Cadena-Castafieda (2015), and they note that the species identified as V. zetterstedti in Panama by Hebard (1927, 1933) and Nickle (1992) corresponds with this species. This species is known from Panama and Colombia (Cigliano et al. 2020). A B & i SSR ee Se ee Fig. 40. Photograph and calling song spectrograms of Viadana brunneri. A. Male; B. and C. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Photo credit: C. Wilson. C H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE The call consists of a series of 2 pulses (Fig. 40B, C) with a total call duration ranging from 4-10 ms and having a mean of 8.6 ms (Table 1). The peak frequency of the entire call is ~16 kHz with a -20 dB frequency range spanning ~15-19 kHz, giving a bandwidth of ~4 kHz (Table 1). The second pulse is usually greater in ampli- tude than the first pulse (Fig. 40C). The two pulses in the call are similar in their temporal and spectral properties (Table 33). The calls of this species were previously described by Falk et al. (2015) and Symes et al. (2016) (identified as V. zetterstedti in these papers). 80 60 40 20 ) Frequency (KHz) 0.0 0.5 1.0 1.5 2.0 Frequency (kHz) 0.04 0.03 0.02 Time (s) Table 33. Call pulse parameters of Viadana brunneri (11 individuals, 195 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (195) 1.34+0.2 16.24 0.5 14.6 + 0.5 18.3 +0.8 3.7+0.5 2 (195) LO t 2. 6.9+0.5 16.24 0.5 14.9+0.5 18 .374.0)7 3.4 + 0.6 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE 181 Phaneropterinae gen. “Waxy sp.” Fig. 41 Phaneropterinae gen. “Waxy sp.” is a mid-sized (0.73 + 0.18 g, n = 73) katydid with very rounded and tough tegmina that have a waxy surface (Fig. 41A, B). We believe that this might be an unde- scribed species and provide the temporary name “Waxy sp.” due to the unusually waxy feel of the wings. The calls recorded from these individuals are all the same and can be readily distinguished from the other katydids we recorded in Panama. The call consists of a series of 6-8 pulses (mean: 6.5; Fig. 41C, D) produced in two groups with a total call duration A C ranging from 65-73 ms and having a mean of ~70 ms (Table 1). The peak frequency of the entire call is ~12 kHz with a -20 dB frequency range spanning ~10-18 kHz, giving a bandwidth of ~8 kHz (Table 1). Pulse amplitudes typically increase across each pulse group (Fig. 41D), but they can also be constant or decrease in amplitude. Pulse durations increase within each pulse group, whereas pulse periods within pulse groups are similar (Table 34). The peak frequency of each pulse decreases within each pulse group (Table 34). The low and high frequencies of each pulse also de- crease slightly within each pulse group, with a bandwidth of ~5-7 kHz (Table 34). 80 60 40 20 0 Z) Frequency (KH Frequency (kHz) 0.00 0.04 0.08 Time (s) 0.12 Fig. 41. Photographs and calling song spectrograms of “Waxy” sp. A. Male; B. Female; C. and D. Spectrogram (top panel) and oscil- logram (bottom panel) of one call at different time scales. Photo credit: C. Wilson. Table 34. Call pulse parameters of “Waxy sp.” (3 individuals, 13 calls; mean + SD); n = number of pulses measured. Pulse Group Pulse Number Pulse Duration Pulse Period (n) (ms) (ms) 1 1 (13) 0,7 #0.2 1 2 (13) 0.9 + 0.2 4.5 40.4 1 3 (13) 16+0.1 4.0 + 0.4 2 1 (13) 0.7 + 0.2 47.4 + 3.2 2 2 (13) 1140.5 Bier 2 3 (13) 16+0.1 4.6 + 0.2 Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (kHz) (kHz) (kHz) 15.5 40.3 12.5+0.8 18.0+0.5 5.5 + 0.6 Loe Oe? 10.6 + 1.0 17.0 + 0.4 6441.2 11.5 + 0.3 102-::0;,6 [ere = O57 5.14+1.2 15.1+0.3 11.6 +0.8 18.0 + 0.4 6.4 +4 1.0 12.94+0.5 10.6 + 0.9 16.7+0.9 (ored Les spl Pre 11.540.5 10.3 + 0.4 15.3 + 0.6 5.0+0.9 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 182 H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Pseudophyllinae Acanthodis curvidens (Stal, 1875) Fig. 42 [MNHN-SO-2019-209, -210, -211] Acanthodis curvidens is a very large (2.98 + 0.2 g, n= 6), brown and green mottled katydid with a blue and white face, purple markings on the ventral surface, and prominent hooked spines on the hind limbs (Fig. 42A, B). It is very well-camouflaged when rest- ing on lichen-covered bark. This species is known from Panama and Colombia (Cigliano et al. 2020). The call begins with a long, low amplitude sound, likely a wing opening sound, followed by 3-4 short pulses and ends with a long- er, higher amplitude pulse (Table 35; Fig. 42C, D). Wing-opening sounds are often also seen before each short pulse (Fig. 42D). The total call duration, not including the first wing-opening sound, ranges from 65-73 ms and has a mean of 64 ms (Table 1). The peak frequency of the call is ~16 kHz with a -20 dB range spanning ~10-22 kHz, giving a bandwidth of ~12 kHz (Table 1). The peak frequency and the amplitude of the pulses increase across the call (Table 35). The initial wing-opening sound is long, and the short pulses that follow the wing-opening sound tend to increase in both duration and peak frequency (Table 35). The fi- nal pulse is longer, greater in amplitude, and has a higher peak frequency than the preceding pulses. The pulse periods of the call are fairly consistent (Table 35). The calls of this species were previously described by Belwood (1988a) and Falk et al. (2015). In addition to acoustic signals, both males and females produce vibrational signals (described in Belwood 1988a). yA C — 80 N Frequency (kHz) 0.10 0.15 0.20 Time (s) Fig. 42. Photographs and calling song spectrograms of Acanthodis curvidens. A. Male (photo credit: T. Robillard); B. Face (photo credit: H. ter Hofstede); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Table 35. Call pulse parameters of Acanthodis curvidens (3 individuals, 38 calls; mean + SD); WO: wing-opening sound at start of each call; LP: last pulse, which is either pulse 4 or 5; n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) WO (38) 19.3. %:0.7 11.7+0.8 8.8+1.1 20.2 + 1.0 11.4+0.5 1 (38) 4.00+1.5 34.4445 12.6 +0.1 8.8+0.5 19.7 $1.3 10.9 + 0.9 2 (38) 4.8+1.8 15.0+1.5 De e052. i A as) Op 20.0 + 0.8 10.7 + 0.6 3 (38) 5.2 + 2.0 15.6 + 0.8 13.6 + 0.9 9.3+0.1 2052) + 055 10.9 + 0.7 4 (15) 5.9 + 0.2 16.0+1.8 14.7 + 0.6 9.6+0.1 19.6 + 2.3 10.0 + 2.4 LP (38) 12.9+ 4.3 13.9 + 0.7 16.14+0.9 11.24 1.4 20.2 + 2.0 8.9+2.8 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE 183 Balboana tibialis (Brunner von Wattenwyl, 1895) Fig. 43 [MNHN-SO-2019-298, -299, -300, -301] Balboana tibialis is a very large (4.76 + 0.75 g, n = 6) and robust dark brown katydid with black patches at the base of the forewings and bright green male cerci (Fig. 43A, B). This species is known from Nicaragua, Costa Rica, Panama, and Colombia (Cigliano et al. 2020). The call consists of a series of 5—8 pulses (mean: 7; Fig. 43C, D) with a total call duration ranging from 105-156 ms and having a mean of ~125 ms (Table 1). The peak frequency of the entire call is ~14 kHz with a -20 dB frequency range spanning ~9-18 kHz, giving a bandwidth of ~9 kHz (Table 1). Pulse amplitudes either A C Frequency (kHz) Frequency (kHz) consistently increase or they increase and then decrease across the call (Fig. 43D). Pulse durations, periods, and peak frequencies all increase across the call (Table 36). The low and high frequencies also in- crease across the call and the bandwidth of each pulse can vary from 5-9 kHz. Some calls have a conspicuous initial wing-open- ing sound with a duration of 11.6 + 0.9 (n = 11 calls). The calls of this species were previously described by Belwood and Morris (1987), Belwood (1988a), ter Hofstede et al. (2010), Jones et al. (2014), Falk et al. (2015), and Symes et al. (2016). In addition to acoustic signals, both males and females produce vi- brational signals (described in Belwood 1988a). 80 60 40 20 Time (s) Fig. 43. Photographs and calling song spectrograms of Balboana tibialis. A. Male (photo credit: H. ter Hofstede); B. Female (photo credit: C. Kernan); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. Table 36. Call pulse parameters of Balboana tibialis (4 individuals, 20 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (20) 3.8 + 0.8 12.44+1.8 8.4 + 1.0 16.8 + 1.6 8.542.2 2 (20) 6.141.1 19.54+2.3 13.041.5 8.8+0.5 Lee AL ne 7.4421 3 (20) 7.6 + 1.6 20.4 + 1.6 1322 43120 9541.2 16.74+1.3 food ea 4 (20) 8.641.5 21.0 + 2.0 14.14 1.6 10.74 1.8 Gs Lee 19 5.4 + 3.3 5 (20) 10.2 + 0.9 21.44 2.2 14.141.3 11.141.5 6.2418 5.0 + 3.0 6 (16) 9.8+41.8 21.8 + 1.6 14.44+1.4 Tle 21, 17.1+0.9 5.9+2.8 with) 9.2+0.4 23.8 + 1.0 14.9+2.2 9:5i4-1,9 17.5+1.2 ScOrt eG JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 184 H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Cocconotus wheeleri Hebard, 1927 Fig. 44 [MNHN-SO-2019-323, -324, -325, -326, -327, -328] Cocconotus wheeleri is a large (1.32 + 0.21 g, n = 18), cylindrical, tan katydid with green markings on the wings, significantly darker dorsal surface of pronotum (black to dark brown) compared to tan colored sides of pronotum, and five black vertical lines on the face (Fig. 44A, B). This species is only known from Panama (Cigli- ano et al. 2020). The call consists of a series of 4-16 (mean: 11) pulses (Fig. 44C, D) with a total call duration ranging from ~79-355 ms and having a mean of ~250 ms (Table 1). The peak frequency of the entire call is ~25 kHz with a -20 dB frequency range spanning A C — 80 60 40 20 Frequency (kHz) Frequency (kHz) 0.0 0.1 02 0.3 Time (s) ~20-27 kHz, giving a bandwidth of ~7 kHz (Table 1). The first 2-3 pulses are much lower in amplitude than the rest of the puls- es, which are usually quite constant in amplitude (Fig. 44C, D), although in some individuals the pulse amplitudes increase and then decrease over the call. The first two pulses are shorter in duration than the rest of the pulses (Table 37). Excluding the first two pulses, both pulse dura- tion and pulse period increase slightly across the call (Table 37). The pulses in the call are all similar in their spectral properties (Table 37). The calls of this species were previously described by Belwood and Morris (1987), Belwood (1988a), and Symes et al. (2016). In addition to acoustic signals, both males and females produce vibrational signals (described in Belwood 1988a). Fig. 44. Photographs and call- ing song spectrograms of Coc- conotus wheeleri. A. Male, inset showing striped face (one of the five black lines is obscured by white glare; photo credit: H. ter Hofstede); B. Female (photo credit: T. Robillard); C. and D. Spectrogram (top panel) and oscillogram (bot- tom panel) of one call at dif- ferent time scales. 04 0.5 Table 37. Call pulse parameters of Cocconotus wheeleri (6 individuals, 60 calls; mean + SD); n = number of pulses measured. Pulse Number (n) Pulse Duration Pulse Period(ms) Peak Frequency Low Frequency High Frequency Bandwidth (kHz) (ms) (kHz) (kHz) (kHz) 1 (60) 5.44 3.8 ie Oe Ga we BW 2072-42156 PLE hehe Wes ceed AS Pel TAL 2 (60) 7.542.9 eA AL eee 7 23.8 + 1.3 2037 5126 28.1 SHEL 2 7.5 41.7 3 (60) el Ne Pe 20.8 + 1.5 23.9414 21.0 + 1.7 27.8 + 1.0 6.9 + 1.6 4 (60) 9.842.1 21.8 + 1.0 24.7 + 0.9 20,2213 28:0 + 13 6.941.4 5 (60) 9441.8 23.8+1.5 24.8 + 0.8 21.441.5 27.9 4+1.2 6.541.3 6 (58) 8.8 + 2.2 23.4 + 0.6 24.9 + 0.8 21.441.4 28.2 4 1.5 6.8 + 1.6 7(55) yt ae bm 2 DE. 24.8 + 0.7 21.34+1.4 27.9 +0.9 6:74 1.2 8 (47) 9.6+0.9 22.8 + 156 24.7 + 1.0 21.441.1 Pits Be ol Ba) 6.5 4+ 1.3 9 (42) 8.9 + 2.1 22.94+1.5 24.6 + 0.8 20.9 + 0.9 28.3 41.2 734 1d 10 (37) 10.14+1.8 23.0.4 2.3 24.5 +0.7 20.9 + 0.8 28.2 41.5 7.241.4 11 (36) 9.541.7 24.7 + 0.7 24.3 + 0.6 21.0 + 1.0 28.2 41.5 7.241.4 12 (31) 11.0 + 2.6 23.7+1.5 24.3 + 0.8 21.0 + 1.1 28.3 41.7 7241.5 13 (27) 10.7 + 1.7 24.5+0.4 24.0 + 0.7 20.9 + 1.2 27.5 +0.7 6.54+1.4 14 (16) 10.4 4+ 2.2 25.3 + 1.0 24.2 + 0.8 20.7 + 1.0 A i lars was 0 Sh) Gi9:4e12 15 (9) 11.942.5 25.8 + 1.0 24.4+0.8 20.8 + 1.4 27.9 40.9 Telit 1h8 JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE Docidocercus gigliotosi (Griffini, 1896) Fig. 45 [MNHN-SO-2019-337, -338, -339, -340, -341, -342, -343] Docidocercus gigliotosi is a medium-sized (1.26 + 0.17 g,n=22), cylindrical, tan katydid with dark and light brown banding on the dorsal surface of the abdomen and a light blue face (Fig. 45A, B). Some individuals have shiny, greenish-yellow coloration on the dorsal surface of the pronotum. This species is only known from Panama (Cigliano et al. 2020). The call consists of 1-3 (mean: 1.6) identical pulses (Fig. 45C, D) with a total call duration ranging from ~13 ms (sin- gle pulse) to 376 ms (3 pulses) and having a mean of ~118 ms (Ta- A C Frequency (kHz) Frequency (kHz) 185 ble 1). Each pulse has a sinusoidal shape (Fig. 45D) and a mean duration of ~20 ms (Table 38). Of the 140 calls measured, 77 were a single pulse, 48 were two pulses, and 15 were three pulses. The peak frequency of each pulse (Table 38) and the entire call is ~24 kHz with a -20 dB frequency range spanning ~23.5-26 kHz, giving a narrow bandwidth of ~2.5 kHz (Table 1). Pulse amplitudes are similar in calls with more than one pulse. The calls of this species were previously described by Belwood and Morris (1987), Belwood (1988a), Morris et al. (1994), R6mer et al. (2010), ter Hofstede et al. (2010), and Falk et al. (2015). In addition to acoustic signals, both males and females produce vi- brational signals (described in Belwood 1988a, R6mer et al. 2010). Oo F&F ®D Ww JS ©C <2 0.00 0.02 0.04 0.06 0.08 0.10 Time (s) Fig. 51. Photographs and calling song spectrograms of Scopiorinus fragilis. A. Male (photo credit: C. Kernan); B. Female (photo credit: T. Robillard); C. and D. Spectrogram (top panel) and oscillogram (bottom panel) of one call at different time scales. JOURNAL OF ORTHOPTERA RESEARCH 2020, 29(2) 192 Thamnobates subfalcata Saussure & Pictet, 1898 Fig. 52 [|MNHN-SO-2019-1817, -1818, -1819] Thamnobates subfalcata is a mid-size (0.63 + 0.19 g, n = 24), brown, cylindrical katydid with a darkened stridulatory area in males (Fig. 52A, B). This species is only known from Panama (Cigliano et al. 2020). The call consists of 2 pulses (Fig. 52C, D) with a total call duration ranging from 21-33 ms and having a mean of 31 ms C A H.M. TER HOFSTEDE, L.B. SYMES, S.J. MARTINSON, T. ROBILLARD, P. FAURE, S$. MADHUSUDHANA AND R.A. PAGE (Table 1). The peak frequency of the entire call is ~19 kHz with a -20 dB frequency range spanning ~17.5-21 kHz, giving a band- width of ~3.5 kHz (Table 1). The two pulses are usually equal in amplitude (Fig. 52D). Wing-opening sounds are usually seen be- fore the first pulse. The first pulse is shorter in duration than the second pulse and the two pulses are similar in their spectral properties (Table 42). An oscillogram of the call of this species is given in Lang et al. (2005). 80 Frequency (kHz) mo F&F DD @W a SS Co