SYSTEMATICS Reevaluation of the Diceroprocta texana Species Complex (: Cicadoidea: )

1 2 ALLEN F. SANBORN AND POLLY K. PHILLIPS

Ann. Entomol. Soc. Am. 103(6): 860Ð865 (2010); DOI: 10.1603/AN10040 ABSTRACT The Diceroprocta texana species complex is currently composed of Diceroprocta texana Downloaded from https://academic.oup.com/aesa/article/103/6/860/116669 by guest on 30 September 2021 texana (Davis, 1916) and Diceroprocta texana lata Davis, 1941. We analyzed physiological, morpho- logical, and biogeography to determine whether these taxa in fact represent two distinct species rather than subspecies. There are statistically signiÞcant differences in morphological, acoustic, and thermal parameters as well as the biogeographic patterns of the two taxa. From these data, we suggest that the two taxa actually represent two species and that D. texana lata be elevated to species full species rank with the name Diceroprocta lata Davis, 1941 n. stat. We provide the Þrst records of D. lata collected in the United States.

KEY WORDS systematics, morphometrics, acoustic behavior, thermal adaptation, cicadas

Davis (1916) described the cicada Diceroprocta texana multiple types of data to determine whether the taxa Davis, 1916 from southern Texas. He then described represent two separate species instead subspecies. the variety Diceroprocta texana variety lata Davis, 1941 for a group of specimens from northern Mexico that Materials and Methods showed afÞnities to D. texana but were larger and darker in coloration than the typical D. texana (Davis Live specimens were collected during June 1985, 1941). He also suggested that the songs of the mem- 1991, and 1998; July 1993; and August 1996. D. texana bers of the D. texana group would be a useful way to were collected in Reeves, Travis, and Waller counties, separate species within the group (Davis 1941). Song TX. D. lata were collected in Hidalgo and Starr coun- differences have been used as the basis of new species ties, TX. Biogeographical data also were collected determination in the Cicadetta montana species com- from specimens in the collections of the American plex in what are morphologically indistinguishable Museum of Natural History; Monte L. Bean Life Sci- taxa (Schedl 1999; Puissant and Boulard 2000; Sueur ence Museum, Brigham Young University; Bohart Mu- and Puissant 2007; Gogala et al. 2008, 2009). seum of Entomology, University of CaliforniaÐDavis; Davis (1862Ð1945) described the majority of cicada Cincinnati Museum of Natural History; Essig Museum species found in the United States. He often described of Entomology, University of CaliforniaÐBerkeley; varieties (which are now considered subspecies under Museum of Biodiversity at The Ohio State University; the current Code of Zoological Nomenclature [ICZN National Museum of Natural History (Smithsonian 1999]) when specimens exhibited morphology similar Institution); Philadelphia Academy of Natural Scienc- to known taxa and he lacked biological information on es; Snow Entomological Museum of the Kansas Nat- the species. Several of his subspecies have since been ural History Survey; Staten Island Institute of Arts and elevated to species rank after their biology had been Sciences; Texas A&M University Entomology Collec- investigated further (Van Duzee 1916; Davis 1930, tion; University of Michigan Museum of Zoology; Uni- 1935; Simons 1953; Heath et al. 1971; Miller 1985; versity of Mississippi; University of TennesseeÐKnox- Sanborn and Phillips 2001; Sanborn 2009; our unpub- ville; and the Utah Museum of Natural History, lished data). University of Utah. We have collected biogeographical, acoustic, ther- Morphological measurements were made with Ver- mal, and morphological data as part of larger investi- nier calipers graduated to 0.05 mm. Live mass of the gations of North American cicadas. We were able to cicadas was determined with a Cent-O-Gram triple collect specimens of both the nominotypical species beam balance (OHaus Scale Corporation, Pine Brook, and subspecies in our expeditions. Consequently, we NJ) sensitive to Ϯ5 mg. All mass measurements were now have the ability to analyze and present here recorded within 10 h of the specimens being collected. Calling songs were recorded using an Uher 4000 Report Monitor tape deck (Uher Werke, Munich, 1 Corresponding author: Department of Biology, Barry University, Germany) and a Sennheiser MKH 70 P 48 directional 11300 NE Second Ave., Miami Shores, FL 33161-6695 (e-mail: [email protected]). microphone (Sennheiser Electronic Corporation, Old 2 17446 SW 33rd Court, Miramar, FL 33029. Lyme, CT) with an MZW 70 wind screen. Frequency

0013-8746/10/0860Ð0865$04.00/0 ᭧ 2010 Entomological Society of America November 2010 SANBORN AND PHILLIPS: D. texana SPECIES COMPLEX 861 response range of the recording equipment is 50Ð verted to power levels (W) before calculating the 20,000 Hz. All calls were recorded on 1.9-cm audiotape statistics because intensity (dB) is measured on a at a tape speed of 19 cm sϪ1. The microphone was logarithmic scale. Mean power output was used to placed within 0.5 m from the calling in an effort calculate mean sound intensity at 50 cm reported for to decrease background noise on the recording. each species. Acoustic signals were analyzed with RavenPro 1.3 The thermal responses (minimum ßight tempera- (Cornell Lab of Ornithology, Ithaca, NY) and a Macin- ture, maximum voluntary tolerance temperature, and tosh computer. Recordings were digitized at a sam- heat torpor temperature) were determined using the pling rate of 40 kHz. Frequency spectra were analyzed procedures outlined in previous cicada studies (Heath using a narrow band Fast Fourier Transform. Speci- 1967, Heath and Wilkin 1970). A Physitemp model mens of D. texana were recorded 10 miles southeast of BAT-12 digital thermocouple thermometer (Physi- Pecos, Reeves Co., TX, whereas specimens of D. lata temp Instruments Inc., Clifton, NJ) with a type MT- were recorded 10 miles north of Rio Grande City, Starr 29/1 copper/constantan 29-gauge hypodermic micro- Downloaded from https://academic.oup.com/aesa/article/103/6/860/116669 by guest on 30 September 2021 Co., TX. Although single populations were the source probe accurate to Ϯ0.1ЊC that had been calibrated of the calls for the acoustic analysis, cicada calls show with a National Institute of Standards and Technology consistency in their characters over extended geo- mercury thermometer was used to measure cicada graphic ranges (e.g., Quartau et al. 2008). body temperature (Tb) when a cicada exhibited a Pulse repetition rates were determined by counting speciÞc behavior. Specimens were cooled into a torpid the total number of pulses in individual syllables and state and tossed 1Ð2 m into the air until they made a dividing by the time for that syllable to occur. The controlled ßight or landing. We then measured the Tb pulse repetition rate and the pulse duration for ten as the minimum ßight temperature or the lowest Tb of syllables were counted, and the mean for each indi- fully coordinated activity. Specimens were placed on vidual was calculated. The mean values for each in- a vertical towel under a heat lamp to determine the dividual were used to determine the mean for the maximum voluntary tolerance or shade-seeking tem- species. Syllables from the middle of a calling bout perature, an upper thermoregulatory temperature were measured to eliminate any potential changes to representing a Tb when thermoregulation takes pre- pulse rate or syllable duration as an animal began or cedence over other behaviors (Heath 1970). Tb was terminated a call. The calls were taken from a region measured when the animal walked or ßew from the of the song where the animal should have been pro- heat source. Heat torpor temperature was determined ducing constant call parameters during the calling by heating a cicada with a heat lamp within a paper bout. Peak frequency was determined by moving the container. Tb was measured when movement stopped. cursor through a syllable and recording the frequency The heat torpor procedure is not lethal to the speci- that had the greatest relative amplitude. mens as cicadas recover within a few minutes as they Sound pressure levels (SPLs) were recorded using cool. Heat torpor temperature is the upper limit of a 2235 SPL meter (Bru¨ el & Kjaer, Naerum, Denmark), activity and represents an ecologically lethal Tb be- a Type 4155 0.5-in. prepolarized condenser micro- cause are no longer able to avoid continued phone, and an UA 0237 wind screen with a ßat re- increases in Tb. The Tb range within which cicadas are sponse to 16 kHz (Bru¨ el & Kjaer). The peak time fully active is delineated by the minimum ßight tem- weighting setting (time constant of Ͻ100 ms) was perature and heat torpor temperatures. Specimens used to ensure that any rapid sound transients were were handled only by the wings for insertion of the measured. The instrument was oriented medially thermocouple to prevent conductive heat transfer along the dorsal side of a singing cicada perpendicular with the . All Tb measurements were recorded to the long body axis at a distance of 50 cm (Sanborn within5softheinsect performing individual be- and Phillips 1995). The distance was kept constant by haviors. placing a 6.35-mm (0.25-in.) dowel attached to the All statistics are reported as mean Ϯ SD. A two- SPL meter near a calling animal. A reading was made tailed t-test was performed to indicate differences in only after the normal calling pattern had been rees- the population means. Analyses were performed using tablished if the cicada was disturbed by placement of InStat 3.1a (GraphPad Software Inc., San Diego, CA). the instrumentation. The alarm call was initiated in the laboratory by manipulating and rotating the insect in Results and Discussion the laboratory at a distance of 50 cm from the micro- phone while the cicada was producing the alarm call Our analyses show there are signiÞcant biogeo- for 30 s to 1 min. The measurement was made after the graphical, morphological, acoustic, and thermal dif- animal had reached an upper thermoregulatory body ferences between D. texana and D. t. lata. Therefore, temperature that represents the body temperature of we propose that D. t. lata be elevated to species rank an animal active in the Þeld (see Sanborn and Phillips and should be known as D. lata Davis, 1941 n. stat. 1995). All intensity measurements are relative to 1 ϫ We found D. lata in southern Texas. These speci- 10Ϫ12 WmϪ2. mens represent the Þrst deÞnitive records for the Power output was determined using the equation species in the United States as it was originally de- Q ϭ 4␲r2(I), where Q is sound power, r is distance scribed from Mexico (Davis 1941). The label data for from source in cm (ϭ50 cm), and I is intensity reading these specimens are: “TEXAS Starr County, U.S. 83 4 for the individual. All SPL measurements were con- miles E of Rio Grande City, 25 June 1994, P. Phillips 862 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 6

Table 1. Morphological, acoustic, and thermal parameters of the D. texana species group

D. texana D. lata Variable t (df) P ͓mean Ϯ SD (n)͔ ͓mean Ϯ SD (n)͔ Live mass (mg) 428 Ϯ 85 (25) 487 Ϯ 75 (12) 2.049 (35) 0.0480 Body length (mm) 22.19 Ϯ 0.97 (18) 23.85 Ϯ 1.04 (10) 4.218 (26) 0.0003 Length of forewing (mm) 27.35 Ϯ 1.44 (18) 30.18 Ϯ 0.80 (10) 5.709 (26) Ͻ0.0001 Forewing width (mm) 9.65 Ϯ 0.45 (18) 10.33 Ϯ 0.40 (10) 3.981 (26) 0.0005 Length of head (mm) 3.63 Ϯ 0.18 (18) 3.91 Ϯ 0.21 (10) 3.625 (26) 0.0012 Width across eyes (mm) 9.20 Ϯ 0.44 (18) 10.08 Ϯ 0.30 (10) 5.555 (26) Ͻ0.0001 Width of pronotum (mm) 8.87 Ϯ 0.39 (18) 9.62 Ϯ 0.45 (10) 4.612 (26) Ͻ0.0001 Width of mesonotum (mm) 7.58 Ϯ 0.32 (18) 8.29 Ϯ 0.31 (10) 5.562 (26) Ͻ0.0001 Wing span (mm) 62.28 Ϯ 3.09 (18) 68.63 Ϯ 1.70 (10) 5.991 (26) Ͻ0.0001 Peak song frequency (kHz) 9.36 Ϯ 0.269 (5) 8.53 Ϯ 0.495 (5) 3.302 (8) 0.0108

Pulse repetition rate (Hz) 93.38 Ϯ 3.62 (5) 124.30 Ϯ 9.55 (5) 6.770 (8) 0.0001 Downloaded from https://academic.oup.com/aesa/article/103/6/860/116669 by guest on 30 September 2021 Syllable duration (ms) 140.38 Ϯ 5.95 (5) 104.38 Ϯ 5.95 (5) 8.801 (8) Ͻ0.0001 Intersyllable interval (ms) 103.00 Ϯ 29.44 (5) 69.33 Ϯ 9.33 (5) 2.438 (8) 0.0407 Calling song power (mW) 10.85 Ϯ 3.71 (7) 62.22 Ϯ 28.67 (4) 4.871 (9) 0.0009 Alarm call power (mW) 26.19 Ϯ 16.41 (8) 39.72 Ϯ 24.12 (7) 1.285 (13) 0.2212 Mean calling song SPL (dB) 95.38 102.97 Mean alarm call SPL (dB) 99.21 101.02 Min. controlled ßight (oC) 19.18 Ϯ 1.98 (24) 20.89 Ϯ 0.96 (7) 2.192 (29) 0.0366 Max voluntary tolerance (oC) 38.33 Ϯ 3.44 (21) 35.07 Ϯ 4.61 (11) 2.264 (30) 0.0310 Heat torpor (oC) 46.42 Ϯ 1.99 (25) 44.43 Ϯ 2.13 (12) 2.800 (35) 0.0083

Measurements were obtained from male specimens. Intensity measurements are relative to 1 ϫ 10Ϫ12 WmϪ2 (ϭ2 ϫ 10Ϫ4 dynes cmϪ2)at a distance of 50 cm. SPL values were calculated from the mean power values. coll.” (4 males); “TEXAS Starr County, U.S. 83 4 miles the dorsal projection and the tip that extends into a E of Rio Grande City, 25 June 1994, A. Sanborn coll.” curved point (Fig. 2). The uncus of D. texana has a (2 males); “TEXAS Starr County, FM 755 20 miles SW smoothly arching posterior margin between the dorsal of La Gloria, 25 June 1994, A. Sanborn, P. Phillips coll.” projection and the tips do not extend into curved (1 male); “TEXAS Starr County, FM 755 20 miles SW points (Fig. 2). From a posterior perspective, the un- of La Gloria, 25 June 1994, P. Phillips coll.” (1 male); cus of D. lata has a wishbone shape with the tips “TEXAS Hidalgo County, Hidalgo, 26o 05Ј 89Љ N78o 14Ј recurving medially so the wishbone is widest in the 32Љ W, 29 August 1996, A. Sanborn coll.” (1 male). middle of its length, whereas the uncus of D. texana has Davis (1941) mentioned specimens near D. lata (but the tips of the wishbone remaining laterally and is failed to designate the specimens deÞnitively as D. widest at the terminus (Fig. 2). lata) in his collection from Starr and Jim Hogg coun- There is a small region of overlap in southern Texas ties, TX, that were smaller than the typical D. lata but in the distribution of D. lata and D. texana, but oth- had similar coloration. The specimens we collected erwise they are not sympatric (Fig. 3). This area of are also smaller than the typical D. lata but match the sympatry is found in the northern limits to D. lata and description and genitalia of D. lata as illustrated in Davis (1941), whereas the genitalia differ from true D. texana. The description of D. lata provided by Davis (1941) is sufÞcient to separate D. lata from D. texana. The species can be separated by the morphological mea- surements given in Table 1 remembering that the differences in size will be larger in specimens from Mexico because the holotype of D. lata is larger than the specimens used for the comparison (Fig. 1). Even with the smaller northern population used in the anal- yses, all nine of the morphological variable measured show statistically signiÞcant differences. When com- paring specimens directly, D. lata is larger, the head is more truncated, the pronotum is wider with a thicker pronotal collar, the coloration is dark brown with large areas of black rather than light brown with small black markings, the basal membrane of the forewing is dark gray rather than light gray, the costal margin is green- ish brown rather than greenish yellow, and the infus- cation on the radial, radio-medial crossveins and on Fig. 1. Habitus of D. texana holotype (top) and D. lata the apex of the ambient vein are much thicker than in holotype (bottom). Specimens are deposited in the collec- D. texana. The uncus of D. lata when viewed from the tion of the American Museum of Natural History. (Online side has an almost straight posterior margin between Þgure in color.) November 2010 SANBORN AND PHILLIPS: D. texana SPECIES COMPLEX 863

ern portion of its range, whereas both species were found calling from Prosopis glandulosa Torr. in the sympatric region in southern Texas. All the locations identiÞed in the literature (Davis 1916, 1921, 1928, 1941; Sanborn 2007) were found in our visits to mu- seum collections with one exception. Davis (1928) suggested that some specimens from Cuernavaca, Mexico, were D. texana. As we did not see this spec- imen and it is signiÞcantly out of range for the species, we presume that it represents one of the Mexican species of Diceroprocta Davis would later describe and was not included in the map or analysis of the distri- bution. Downloaded from https://academic.oup.com/aesa/article/103/6/860/116669 by guest on 30 September 2021 There are some differences in the habitats used by D. texana and D. lata as well. We found D. texana in a plant community that is characterized as the northern portion of the Chihuahuan Desert in New Mexico and western Texas (MacMahon 1988), the desert grassland of the south central United States (Sims 1988), and the vegetation of the southeastern coastal plain (Chris- tensen 1988) as the species expands south and east. The plant communities for D. lata can be described as primarily desert grassland (Sims 1988) and the eastern edges of the Chihuahuan Desert (MacMahon 1988). A similar differentiation in habitat use was found when elevating D. aurantiaca Davis to species rank (Sanborn Fig. 2. Male genitalia of D. texana (A and B) and D. lata and Phillips 2001). (C and D). The arrow illustrates the widening of the uncal The calling song parameters and sound pressure lobes being widest at the terminus in D. texana (A), whereas levels also indicate signiÞcant differences between the it is widest in the middle of the wishbone shape in D. lata (C). species (Table 1). The songs of both species are com- The lateral views show the difference in the shape of the posed of a series of syllables (Fig. 4). Each syllable is recurved uncus in D. texana (B) and D. lata (D). (Online Þgure in color.) produced as a series of sound pulses produced as the timbal plate and timbal ribs buckle. There is a variable number of pulses produced within an individual song the southeastern limits of the distribution of D. texana. as the number of ribs buckling with each muscle con- Individual host plants could not be determined as we traction varies. However, the pulse repetition rate observed both species active in various natural and (P ϭ 0.0001), syllable duration (P Ͻ 0.0001), and disturbed habitats. D. texana exhibited a tendency to intersyllable interval (P ϭ 0.0407) all differ signiÞ- call from Larrea tridentata (DC.) Colville in the west- cantly. The song of D. texana has a signiÞcantly higher peak song frequency than the song of D. lata that would be expected based on weight or body size (Bennet-Clark and Young 1994, Daniel et al. 1993). Because the frequency characteristics of a cicada call are determined by the physics of the sound production system (Pringle 1954, Bennet-Clark 1995), there is a physical basis to the observed differences. These dif- ferences are used by female cicadas to select mates (e.g., Doolan and Young 1989). The song power and SPL levels also differ signiÞcantly for D. texana and D. lata (P ϭ 0.0009). This relationship is expected based on the differences in body mass of the two species (Sanborn and Phillips 1995). The lack of signiÞcance (P ϭ 0.2212) for the alarm call power data may be related to our inability to stimulate maximum sound energy production during manipulation in the labo- ratory. Most (Þve of eight) of the D. texana alarm calls were more powerful than the loudest animal recorded in the Þeld with one producing more than twice the power output of the loudest animal in the Þeld, Fig. 3. Biogeography of D. texana (circles) and D. lata whereas three of the seven D. lata alarm calls were less (squares). (Online Þgure in color.) than the lowest value reported in the Þeld. The vari- 864 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 6

species so similar thermal responses would be pre- dicted. Multiple analyses have demonstrated the differ- ences between D. texana and D. lata. Morphological, acoustic, thermal and biogeographical data all strongly support the hypothesis that D. lata be elevated to species rank, and thus D. texana lata should now be known as D. lata n. stat.

Acknowledgments We thank the C. Barr (Essig Museum of Entomology, University of California Berkeley), R. Baumann and S. Clark Downloaded from https://academic.oup.com/aesa/article/103/6/860/116669 by guest on 30 September 2021 (Monte L. Bean Life Science Museum, Brigham Young Uni- versity), C. Bills and E. Rickart (Utah Museum of Natural History, University of Utah), R. Brooks, G. Byers and K. Segelquist (Snow Entomological Museum of the Kansas Nat- ural History Survey), G. Dahlem (Cincinnati Museum of Natural History), M. Epstien and R. Froeschner (National Museum of Natural History, Smithsonian Institution), M. S. and J. E. Heath, S. Heydon (Bohart Museum of Entomology, University of CaliforniaÐDavis), E. Johnson (Staten Island Institute of Arts and Sciences), P. Lago (University of Mis- sissippi), M. OÕBrien (University of Michigan Museum of Zoology), E. Riley (Texas A&M University Entomology Col- lection), R. Schuh and S. Oygur (American Museum of Nat- ural History), O. Schwarz and C. Boake (University of Ten- nesseeÐKnoxville), A. Sharkov (Museum of Biodiversity at The Ohio State University), and J. Weintraub and D. Azuma (Philadelphia Academy of Natural Sciences) for assistance Fig. 4. A. Oscillogram (top trace) and sonagram (bottom and access to their collections. Two anonymous reviewers trace) of the calling song produced by D. texana. The call is made suggestions that improved the manuscript. A.F.S. was a series of syllables of constant frequency and amplitude with supported by Sister John Karen Frei and Research Mini- a peak frequency of Ϸ9.4 kHz. Syllable duration is Ϸ140 ms, Grants and a sabbatical leave from Barry University. with individual sound pulses being produced at a rate of Ϸ93 Hz. B. Oscillogram (top trace) and sonagram (bottom trace) References Cited of the calling song produced by D. lata. The call is a series of syllables of constant frequency and amplitude with a peak Bennet-Clark, H. C. 1995. Insect sound production: trans- frequency of Ϸ8.5 kHz. Syllable duration is Ϸ104 ms, with duction mechanisms and impedance matching. Symp. individual sound pulses being produced at a rate of Ϸ124 Hz. Soc. Exp. Biol. 49: 383Ð409. Bennet-Clark, H. C., and D. Young. 1994. The scaling of song frequencies in cicadas. J. Exp. Biol. 191: 291Ð294. ability in the laboratory responses seems to have in- Christensen, N. L. 1988. Vegetation of the Southeastern ßuenced the signiÞcance of the results. coastal plain, pp. 317Ð363. In M. G. Barbour and W. D. The thermal responses determined for each group Billings (eds.), North American terrestrial vegetation. are summarized in Table 1. All three thermal re- Cambridge University Press, Cambridge, MA. sponses show statistical differences between the spe- Daniel, H. J., C. Knight, T. M. Charles, and A. L. Larkins. cies. Although minimum ßight temperature has been 1993. Predicting species by call in three species of North suggested to be related to habitat (Heath et al. 1971, Carolina cicadas. J. Elisha Mitchell Sci. Soc. 109: 67Ð76. 1972), further investigations have shown minimum Davis, W. T. 1916. Notes on cicadas from the United States ßight temperature does not relate only to the envi- with descriptions of several new species. J. NY Entomol. Soc. 24: 42Ð65. ronment of a species but is inßuenced by the mor- Davis, W. T. 1921. Records of cicadas from North America phology of the ßight system (Sanborn et al. 2001). The with descriptions of new species. J. NY Entomol. Soc. 29: higher maximum voluntary tolerance and heat torpor 1Ð16. temperatures for D. texana probably relate to the po- Davis, W. T. 1928. Cicadas belonging to the genus Dicero- tentially warmer environments in which the species is procta with descriptions of a new species. J. NY Entomol. found. The values of the thermal responses deter- Soc. 36: 439Ð458. mined for D. texana and D. lata are similar to the other Davis, W. T. 1930. The distribution of cicadas in the United species that inhabit have been collected with D. texana States with descriptions of new species. J. NY Entomol. and D. lata like the Diceroprocta cinctifera (Uhler) Soc. 38: 53Ð73. Davis, W. T. 1935. Six new cicadas from the western United species group (Sanborn and Phillips 1996), Cacama States. J. NY Entomol. Soc. 43: 299Ð310. valvata (Uhler) (Heath et al. 1972), D. aurantiaca and Davis, W. T. 1941. New cicadas from North America with Diceroprocta delicata (Osborn) (Sanborn and Phillips notes. J. NY Entomol. Soc. 49: 85Ð99. 2001). There is an overlap of the distributions of D. Doolan, J. M., and D. Young. 1989. Relative importance of texana and D. lata with the distributions of all of these song parameters during ßight phonotaxis and courtship in November 2010 SANBORN AND PHILLIPS: D. texana SPECIES COMPLEX 865

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