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Reevaluation of the Diceroprocta Texana Species Complex (Hemiptera: Cicadoidea: Cicadidae)

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Reevaluation of the Diceroprocta Texana Species Complex (Hemiptera: Cicadoidea: Cicadidae) SYSTEMATICS Reevaluation of the Diceroprocta texana Species Complex (Hemiptera: Cicadoidea: Cicadidae) 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 animal 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 animals 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 insect. 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.
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