SYSTEMATICS
Reevaluation of the Diceroprocta texana Species Complex (Hemiptera: Cicadoidea: Cicadidae) ALLEN F. SANBORN1
AND
POLLY K. PHILLIPS2
Ann. Entomol. Soc. Am. 103(6): 860Ð865 (2010); DOI: 10.1603/AN10040
ABSTRACT The Diceroprocta texana species complex is currently composed of Diceroprocta texana texana (Davis, 1916) and Diceroprocta texana lata Davis, 1941. We analyzed physiological, morphological, 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 Davis, 1916 from southern Texas. He then described the variety Diceroprocta texana variety lata Davis, 1941 for a group of specimens from northern Mexico that showed afÞnities to D. texana but were larger and darker in coloration than the typical D. texana (Davis 1941). He also suggested that the songs of the members of the D. texana group would be a useful way to separate species within the group (Davis 1941). Song differences have been used as the basis of new species determination in the Cicadetta montana species complex in what are morphologically indistinguishable taxa (Schedl 1999; Puissant and Boulard 2000; Sueur and Puissant 2007; Gogala et al. 2008, 2009). Davis (1862Ð1945) described the majority of cicada species found in the United States. He often described varieties (which are now considered subspecies under the current Code of Zoological Nomenclature [ICZN 1999]) when specimens exhibited morphology similar to known taxa and he lacked biological information on the species. Several of his subspecies have since been elevated to species rank after their biology had been investigated further (Van Duzee 1916; Davis 1930, 1935; Simons 1953; Heath et al. 1971; Miller 1985; Sanborn and Phillips 2001; Sanborn 2009; our unpublished data). We have collected biogeographical, acoustic, thermal, and morphological data as part of larger investigations of North American cicadas. We were able to collect specimens of both the nominotypical species and subspecies in our expeditions. Consequently, we now have the ability to analyze and present here 1 Corresponding author: Department of Biology, Barry University, 11300 NE Second Ave., Miami Shores, FL 33161-6695 (e-mail:
[email protected]). 2 17446 SW 33rd Court, Miramar, FL 33029.
multiple types of data to determine whether the taxa represent two separate species instead subspecies. Materials and Methods Live specimens were collected during June 1985, 1991, and 1998; July 1993; and August 1996. D. texana were collected in Reeves, Travis, and Waller counties, TX. D. lata were collected in Hidalgo and Starr counties, TX. Biogeographical data also were collected from specimens in the collections of the American Museum of Natural History; Monte L. Bean Life Science Museum, Brigham Young University; Bohart Museum of Entomology, University of CaliforniaÐDavis; Cincinnati Museum of Natural History; Essig Museum of Entomology, University of CaliforniaÐBerkeley; Museum of Biodiversity at The Ohio State University; National Museum of Natural History (Smithsonian Institution); Philadelphia Academy of Natural Sciences; Snow Entomological Museum of the Kansas Natural History Survey; Staten Island Institute of Arts and Sciences; Texas A&M University Entomology Collection; University of Michigan Museum of Zoology; University of Mississippi; University of TennesseeÐKnoxville; and the Utah Museum of Natural History, University of Utah. Morphological measurements were made with Vernier calipers graduated to 0.05 mm. Live mass of the cicadas was determined with a Cent-O-Gram triple beam balance (OHaus Scale Corporation, Pine Brook, NJ) sensitive to ⫾5 mg. All mass measurements were recorded within 10 h of the specimens being collected. Calling songs were recorded using an Uher 4000 Report Monitor tape deck (Uher Werke, Munich, Germany) and a Sennheiser MKH 70 P 48 directional microphone (Sennheiser Electronic Corporation, Old Lyme, CT) with an MZW 70 wind screen. Frequency
0013-8746/10/0860Ð0865$04.00/0 䉷 2010 Entomological Society of America
November 2010
SANBORN AND PHILLIPS: D. texana SPECIES COMPLEX
response range of the recording equipment is 50 Ð 20,000 Hz. All calls were recorded on 1.9-cm audiotape at a tape speed of 19 cm s⫺1. The microphone was placed within 0.5 m from the calling animal in an effort to decrease background noise on the recording. Acoustic signals were analyzed with RavenPro 1.3 (Cornell Lab of Ornithology, Ithaca, NY) and a Macintosh computer. Recordings were digitized at a sampling rate of 40 kHz. Frequency spectra were analyzed using a narrow band Fast Fourier Transform. Specimens of D. texana were recorded 10 miles southeast of Pecos, Reeves Co., TX, whereas specimens of D. lata were recorded 10 miles north of Rio Grande City, Starr Co., TX. Although single populations were the source of the calls for the acoustic analysis, cicada calls show consistency in their characters over extended geographic ranges (e.g., Quartau et al. 2008). Pulse repetition rates were determined by counting the total number of pulses in individual syllables and dividing by the time for that syllable to occur. The pulse repetition rate and the pulse duration for ten syllables were counted, and the mean for each individual was calculated. The mean values for each individual were used to determine the mean for the species. Syllables from the middle of a calling bout were measured to eliminate any potential changes to pulse rate or syllable duration as an animal began or terminated a call. The calls were taken from a region of the song where the animal should have been producing constant call parameters during the calling bout. Peak frequency was determined by moving the cursor through a syllable and recording the frequency that had the greatest relative amplitude. Sound pressure levels (SPLs) were recorded using a 2235 SPL meter (Bru¨ el & Kjaer, Naerum, Denmark), a Type 4155 0.5-in. prepolarized condenser microphone, and an UA 0237 wind screen with a ßat response to 16 kHz (Bru¨ el & Kjaer). The peak time weighting setting (time constant of ⬍100 ms) was used to ensure that any rapid sound transients were measured. The instrument was oriented medially along the dorsal side of a singing cicada perpendicular to the long body axis at a distance of 50 cm (Sanborn and Phillips 1995). The distance was kept constant by placing a 6.35-mm (0.25-in.) dowel attached to the SPL meter near a calling animal. A reading was made only after the normal calling pattern had been reestablished if the cicada was disturbed by placement of the instrumentation. The alarm call was initiated in the laboratory by manipulating and rotating the insect in the laboratory at a distance of 50 cm from the microphone while the cicada was producing the alarm call for 30 s to 1 min. The measurement was made after the animal had reached an upper thermoregulatory body temperature that represents the body temperature of an animal active in the Þeld (see Sanborn and Phillips 1995). All intensity measurements are relative to 1 ⫻ 10⫺12 W m⫺2. Power output was determined using the equation Q ⫽ 4r2(I), where Q is sound power, r is distance from source in cm (⫽50 cm), and I is intensity reading for the individual. All SPL measurements were con-
861
verted to power levels (W) before calculating the statistics because intensity (dB) is measured on a logarithmic scale. Mean power output was used to calculate mean sound intensity at 50 cm reported for each species. The thermal responses (minimum ßight temperature, maximum voluntary tolerance temperature, and heat torpor temperature) were determined using the procedures outlined in previous cicada studies (Heath 1967, Heath and Wilkin 1970). A Physitemp model BAT-12 digital thermocouple thermometer (Physitemp Instruments Inc., Clifton, NJ) with a type MT29/1 copper/constantan 29-gauge hypodermic microprobe accurate to ⫾0.1⬚C that had been calibrated with a National Institute of Standards and Technology mercury thermometer was used to measure cicada body temperature (Tb) when a cicada exhibited a speciÞc behavior. Specimens were cooled into a torpid state and tossed 1Ð2 m into the air until they made a controlled ßight or landing. We then measured the Tb as the minimum ßight temperature or the lowest Tb of fully coordinated activity. Specimens were placed on a vertical towel under a heat lamp to determine the maximum voluntary tolerance or shade-seeking temperature, an upper thermoregulatory temperature representing a Tb when thermoregulation takes precedence over other behaviors (Heath 1970). Tb was measured when the animal walked or ßew from the heat source. Heat torpor temperature was determined by heating a cicada with a heat lamp within a paper container. Tb was measured when movement stopped. The heat torpor procedure is not lethal to the specimens as cicadas recover within a few minutes as they cool. Heat torpor temperature is the upper limit of activity and represents an ecologically lethal Tb because animals are no longer able to avoid continued increases in Tb. The Tb range within which cicadas are fully active is delineated by the minimum ßight temperature and heat torpor temperatures. Specimens were handled only by the wings for insertion of the thermocouple to prevent conductive heat transfer with the insect. All Tb measurements were recorded within 5 s of the insect performing individual behaviors. All statistics are reported as mean ⫾ SD. A twotailed t-test was performed to indicate differences in the population means. Analyses were performed using InStat 3.1a (GraphPad Software Inc., San Diego, CA). Results and Discussion Our analyses show there are signiÞcant biogeographical, morphological, acoustic, and thermal differences between D. texana and D. t. lata. Therefore, we propose that D. t. lata be elevated to species rank and should be known as D. lata Davis, 1941 n. stat. We found D. lata in southern Texas. These specimens represent the Þrst deÞnitive records for the species in the United States as it was originally described from Mexico (Davis 1941). The label data for these specimens are: “TEXAS Starr County, U.S. 83 4 miles E of Rio Grande City, 25 June 1994, P. Phillips
862
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Table 1.
Vol. 103, no. 6
Morphological, acoustic, and thermal parameters of the D. texana species group
Variable
D. texana 关mean ⫾ SD (n)兴
D. lata 关mean ⫾ SD (n)兴
Live mass (mg) Body length (mm) Length of forewing (mm) Forewing width (mm) Length of head (mm) Width across eyes (mm) Width of pronotum (mm) Width of mesonotum (mm) Wing span (mm) Peak song frequency (kHz) Pulse repetition rate (Hz) Syllable duration (ms) Intersyllable interval (ms) Calling song power (mW) Alarm call power (mW) Mean calling song SPL (dB) Mean alarm call SPL (dB) Min. controlled ßight (oC) Max voluntary tolerance (oC) Heat torpor (oC)
428 ⫾ 85 (25) 22.19 ⫾ 0.97 (18) 27.35 ⫾ 1.44 (18) 9.65 ⫾ 0.45 (18) 3.63 ⫾ 0.18 (18) 9.20 ⫾ 0.44 (18) 8.87 ⫾ 0.39 (18) 7.58 ⫾ 0.32 (18) 62.28 ⫾ 3.09 (18) 9.36 ⫾ 0.269 (5) 93.38 ⫾ 3.62 (5) 140.38 ⫾ 5.95 (5) 103.00 ⫾ 29.44 (5) 10.85 ⫾ 3.71 (7) 26.19 ⫾ 16.41 (8) 95.38 99.21 19.18 ⫾ 1.98 (24) 38.33 ⫾ 3.44 (21) 46.42 ⫾ 1.99 (25)
487 ⫾ 75 (12) 23.85 ⫾ 1.04 (10) 30.18 ⫾ 0.80 (10) 10.33 ⫾ 0.40 (10) 3.91 ⫾ 0.21 (10) 10.08 ⫾ 0.30 (10) 9.62 ⫾ 0.45 (10) 8.29 ⫾ 0.31 (10) 68.63 ⫾ 1.70 (10) 8.53 ⫾ 0.495 (5) 124.30 ⫾ 9.55 (5) 104.38 ⫾ 5.95 (5) 69.33 ⫾ 9.33 (5) 62.22 ⫾ 28.67 (4) 39.72 ⫾ 24.12 (7) 102.97 101.02 20.89 ⫾ 0.96 (7) 35.07 ⫾ 4.61 (11) 44.43 ⫾ 2.13 (12)
t (df)
P
2.049 (35) 4.218 (26) 5.709 (26) 3.981 (26) 3.625 (26) 5.555 (26) 4.612 (26) 5.562 (26) 5.991 (26) 3.302 (8) 6.770 (8) 8.801 (8) 2.438 (8) 4.871 (9) 1.285 (13)
0.0480 0.0003 ⬍0.0001 0.0005 0.0012 ⬍0.0001 ⬍0.0001 ⬍0.0001 ⬍0.0001 0.0108 0.0001 ⬍0.0001 0.0407 0.0009 0.2212
2.192 (29) 2.264 (30) 2.800 (35)
0.0366 0.0310 0.0083
Measurements were obtained from male specimens. Intensity measurements are relative to 1 ⫻ 10⫺12 W m⫺2 (⫽2 ⫻ 10⫺4 dynes cm⫺2) 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 E of Rio Grande City, 25 June 1994, A. Sanborn coll.” (2 males); “TEXAS Starr County, FM 755 20 miles SW of La Gloria, 25 June 1994, A. Sanborn, P. Phillips coll.” (1 male); “TEXAS Starr County, FM 755 20 miles SW of La Gloria, 25 June 1994, P. Phillips coll.” (1 male); “TEXAS Hidalgo County, Hidalgo, 26o 05⬘ 89⬙ N 78o 14⬘ 32⬙ W, 29 August 1996, A. Sanborn coll.” (1 male). Davis (1941) mentioned specimens near D. lata (but failed to designate the specimens deÞnitively as D. lata) in his collection from Starr and Jim Hogg counties, TX, that were smaller than the typical D. lata but had similar coloration. The specimens we collected are also smaller than the typical D. lata but match the 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 measurements 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 analyses, all nine of the morphological variable measured show statistically signiÞcant differences. When comparing 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 greenish brown rather than greenish yellow, and the infuscation on the radial, radio-medial crossveins and on the apex of the ambient vein are much thicker than in D. texana. The uncus of D. lata when viewed from the side has an almost straight posterior margin between
the dorsal projection and the tip that extends into a curved point (Fig. 2). The uncus of D. texana has a smoothly arching posterior margin between the dorsal projection and the tips do not extend into curved points (Fig. 2). From a posterior perspective, the uncus of D. lata has a wishbone shape with the tips recurving medially so the wishbone is widest in the middle of its length, whereas the uncus of D. texana has the tips of the wishbone remaining laterally and is widest at the terminus (Fig. 2). There is a small region of overlap in southern Texas in the distribution of D. lata and D. texana, but otherwise they are not sympatric (Fig. 3). This area of sympatry is found in the northern limits to D. lata and
Fig. 1. Habitus of D. texana holotype (top) and D. lata holotype (bottom). Specimens are deposited in the collection of the American Museum of Natural History. (Online Þgure in color.)
November 2010
SANBORN AND PHILLIPS: D. texana SPECIES COMPLEX
Fig. 2. Male genitalia of D. texana (A and B) and D. lata (C and D). The arrow illustrates the widening of the uncal lobes being widest at the terminus in D. texana (A), whereas it is widest in the middle of the wishbone shape in D. lata (C). The lateral views show the difference in the shape of the recurved uncus in D. texana (B) and D. lata (D). (Online Þgure in color.)
the southeastern limits of the distribution of D. texana. Individual host plants could not be determined as we observed both species active in various natural and disturbed habitats. D. texana exhibited a tendency to call from Larrea tridentata (DC.) Colville in the west-
Fig. 3. Biogeography of D. texana (circles) and D. lata (squares). (Online Þgure in color.)
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 museum collections with one exception. Davis (1928) suggested that some specimens from Cuernavaca, Mexico, were D. texana. As we did not see this specimen 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 distribution. 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 (Christensen 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 and Phillips 2001). The calling song parameters and sound pressure levels also indicate signiÞcant differences between the species (Table 1). The songs of both species are composed of a series of syllables (Fig. 4). Each syllable is 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 as the number of ribs buckling with each muscle contraction varies. However, the pulse repetition rate (P ⫽ 0.0001), syllable duration (P ⬍ 0.0001), and intersyllable interval (P ⫽ 0.0407) all differ signiÞcantly. 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 differences 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 ⫽ 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 ⫽ 0.2212) for the alarm call power data may be related to our inability to stimulate maximum sound energy production during manipulation in the laboratory. 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, whereas three of the seven D. lata alarm calls were less 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 predicted. Multiple analyses have demonstrated the differences 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
Fig. 4. A. Oscillogram (top trace) and sonagram (bottom trace) of the calling song produced by D. texana. The call is a series of syllables of constant frequency and amplitude with a peak frequency of ⬇9.4 kHz. Syllable duration is ⬇140 ms, with individual sound pulses being produced at a rate of ⬇93 Hz. B. Oscillogram (top trace) and sonagram (bottom trace) of the calling song produced by D. lata. The call is a series of syllables of constant frequency and amplitude with a peak frequency of ⬇8.5 kHz. Syllable duration is ⬇104 ms, with individual sound pulses being produced at a rate of ⬇124 Hz.
ability in the laboratory responses seems to have inßuenced the signiÞcance of the results. The thermal responses determined for each group are summarized in Table 1. All three thermal responses show statistical differences between the species. Although minimum ßight temperature has been suggested to be related to habitat (Heath et al. 1971, 1972), further investigations have shown minimum ßight temperature does not relate only to the environment of a species but is inßuenced by the morphology of the ßight system (Sanborn et al. 2001). The higher maximum voluntary tolerance and heat torpor temperatures for D. texana probably relate to the potentially warmer environments in which the species is found. The values of the thermal responses determined for D. texana and D. lata are similar to the other species that inhabit have been collected with D. texana and D. lata like the Diceroprocta cinctifera (Uhler) species group (Sanborn and Phillips 1996), Cacama valvata (Uhler) (Heath et al. 1972), D. aurantiaca and Diceroprocta delicata (Osborn) (Sanborn and Phillips 2001). There is an overlap of the distributions of D. texana and D. lata with the distributions of all of these
We thank the C. Barr (Essig Museum of Entomology, University of California Berkeley), R. Baumann and S. Clark (Monte L. Bean Life Science Museum, Brigham Young University), 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 Natural 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 Mississippi), M. OÕBrien (University of Michigan Museum of Zoology), E. Riley (Texas A&M University Entomology Collection), R. Schuh and S. Oygur (American Museum of Natural History), O. Schwarz and C. Boake (University of TennesseeÐKnoxville), A. Sharkov (Museum of Biodiversity at The Ohio State University), and J. Weintraub and D. Azuma (Philadelphia Academy of Natural Sciences) for assistance and access to their collections. Two anonymous reviewers made suggestions that improved the manuscript. A.F.S. was supported by Sister John Karen Frei and Research MiniGrants and a sabbatical leave from Barry University.
References Cited Bennet-Clark, H. C. 1995. Insect sound production: transduction mechanisms and impedance matching. Symp. Soc. Exp. Biol. 49: 383Ð 409. Bennet-Clark, H. C., and D. Young. 1994. The scaling of song frequencies in cicadas. J. Exp. Biol. 191: 291Ð294. Christensen, N. L. 1988. Vegetation of the Southeastern coastal plain, pp. 317Ð363. In M. G. Barbour and W. D. Billings (eds.), North American terrestrial vegetation. Cambridge University Press, Cambridge, MA. Daniel, H. J., C. Knight, T. M. Charles, and A. L. Larkins. 1993. Predicting species by call in three species of North Carolina cicadas. J. Elisha Mitchell Sci. Soc. 109: 67Ð76. Davis, W. T. 1916. Notes on cicadas from the United States with descriptions of several new species. J. NY Entomol. Soc. 24: 42Ð 65. Davis, W. T. 1921. Records of cicadas from North America with descriptions of new species. J. NY Entomol. Soc. 29: 1Ð16. Davis, W. T. 1928. Cicadas belonging to the genus Diceroprocta with descriptions of a new species. J. NY Entomol. Soc. 36: 439 Ð 458. Davis, W. T. 1930. The distribution of cicadas in the United States with descriptions of new species. J. NY Entomol. Soc. 38: 53Ð73. Davis, W. T. 1935. Six new cicadas from the western United States. J. NY Entomol. Soc. 43: 299 Ð310. Davis, W. T. 1941. New cicadas from North America with notes. J. NY Entomol. Soc. 49: 85Ð99. Doolan, J. M., and D. Young. 1989. Relative importance of song parameters during ßight phonotaxis and courtship in
November 2010
SANBORN AND PHILLIPS: D. texana SPECIES COMPLEX
the bladder cicada Cystosoma saundersii. J. Exp. Biol. 141: 113Ð131. Gogala, M., S. Drosopoulos, and T. Trilar. 2008. Cicadetta montana complex (Hemiptera, Cicadidae) in GreeceÑa new species and new records based on bioacoustics. Dtsch. Entomol. Z. 55: 91Ð100. Gogala, M., S. Drosopoulos, and T. Trilar. 2009. Two mountains, two species: new taxa of the Cicadetta montana species complex in Greece (Hemiptera: Cicadidae). Acta Entomol. Slov. 17: 13Ð28. Heath, J. E. 1967. Temperature responses of the periodical “17-year” cicada, Magicicada cassini (Homoptera, Cicadidae). Am. Midl. Nat. 77: 64 Ð 67. Heath, J. E. 1970. Behavioral regulation of body temperature in poikilotherms. Physiologist 13: 399 Ð 410. Heath, J. E., J. L. Hanagan, P. J. Wilkin, and M. S. Heath. 1971. Adaptation of the thermal responses of insects. Am. Zool. 11: 147Ð158. Heath, J. E., and P. J. Wilkin. 1970. Temperature responses of the desert cicada, Diceroprocta apache (Homoptera, Cicadidae). Physiol. Zool. 43: 145Ð154. Heath, J. E., P. J. Wilkin, and M. S. Heath. 1972. Temperature responses of the cactus dodger, Cacama valvata (Homoptera, Cicadidae). Physiol. Zool. 45: 238 Ð246. [ICZN] International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomenclature, 4th ed. International Trust for Zoological Nomenclature, London, United Kingdom. MacMahon, J. A. 1988. Warm deserts, pp. 231Ð264. In M. G. Barbour and W. D. Billings (eds.), North American terrestrial vegetation. Cambridge University Press, Cambridge, MA. Miller, S. E. 1985. The California Channel IslandsÑpast, present, and future: an entomological perspective, pp. 3Ð27. In A. S. Menke and D. R. Miller (eds.), Entomology of the California Channel Islands. Proceedings of the Þrst Symposium. Santa Barbara Museum of Natural History, Santa Barbara, CA. Pringle, J.W.S. 1954. A physiological analysis of cicada song. J. Exp. Biol. 31: 525Ð560. Puissant, S., and M. Boulard. 2000. Cicadetta cerdaniensis, espe` ce jumelle de Cicadetta montana de´ crypte´ e par
865
lÕacoustique (Auchenorhyncha, Cicadidae, Tibicininae). EPHE, Trav. Lab. Biol. Evol. Ins. Hemipt. 13: 111Ð117. Quartau, J. A., G. Andre´, G. Pinto-Juma, S. G. Seabra, and P. C. Simo˜ es. 2008. Geographic variation in the calling song of Cicada orni L. (Hemiptera: Cicadoidea) in the Mediterranean area. Bol. Mus. Mun. Funchal, Sup. 14: 127Ð130. Sanborn, A. F. 2007. New species, new records and checklist of cicadas from Mexico (Hemiptera: Cicadoidea: Cicadidae). Zootaxa 1651: 1Ð 42. Sanborn, A. F. 2009. Checklist, new species and key to the cicadas of Cuba (Hemiptera: Cicadoidea: Cicadidae). Dtsch. Entomol. Z. 59: 85Ð92. Sanborn, A. F., and P. K. Phillips. 1995. Scaling of sound pressure level and body size in cicadas (Homoptera: Cicadidae; Tibicinidae). Ann. Entomol. Soc. Am. 88: 479 Ð 484. Sanborn, A. F., and P. K. Phillips. 1996. Thermal responses of the Diceroprocta cinctifera species group (Homoptera: Cicadidae). Southwest. Nat. 41: 136 Ð139. Sanborn, A. F., and P. K. Phillips. 2001. Re-evaluation of the Diceroprocta delicata species complex (Homoptera: Cicadidae). Ann. Entomol. Soc. Am. 94: 159 Ð165. Sanborn, A. F., L. M. Perez, C. G. Valdes, and A. K. Seepersaud. 2001. Wing morphology and minimum ßight temperature in cicadas (Insecta: Homoptera: Cicadoidea). FASEB J. 15: A1106. Schedl, W. 1999. Eine neue Unterart der Bergsingzikade im Balkan, Cicadetta montana macedonica ssp. n. (Hemiptera: Auchenorrhyncha: Cicadomorpha: Tibicinidae). Reichenbachia 33: 87Ð90. Sueur, J., and S. Puissant. 2007. Similar look but different song: a new Cicadetta species in the montana complex (Insecta, Hemiptera, Cicadidae). Zootaxa 1442: 55Ð 68. Simons, J. N. 1953. New California cicadas with taxonomic notes on other species. Pan-Pac. Entomol. 29: 191Ð198. Sims, P. L. 1988. Grasslands, pp. 266 Ð2286. In M. G. Barbour and W. D. Billings (eds.), North American terrestrial vegetation. Cambridge University Press, Cambridge, MA. Received 9 March 2010; accepted 27 August 2010.
