Diversity 2013, 5(2), 166-239; doi:10.3390/d5020166

Article
Biogeography of the Cicadas (Hemiptera: Cicadidae) of North America, North of Mexico
Allen F. Sanborn 1,* and Polly K. Phillips 2
1
Department of Biology, Barry University, 11300 NE Second Avenue, Miami Shores, FL 33161-6695, USA
2
17446 SW 33rd Ct, Miramar, FL 33029, USA; E-Mail: pkmcp31@illinoisalumni.org
*
Author to whom correspondence should be addressed; E-Mail: asanborn@mail.barry.edu; Tel.: +1-305-899-3219; Fax: +1-305-899-3225.
Received: 28 January 2013; in revised form: 2 March 2013 / Accepted: 22 March 2013 /
Published: 9 April 2013

Abstract

: We describe and illustrate the biogeography of the cicadas inhabiting continental North America, north of Mexico. Species distributions were determined through our collecting efforts as well as label data from more than 110 institutional collections. The status of subspecies is discussed with respect to their distributions. As we have shown over limited geographic areas, the distribution of individual species is related to the habitat in which they are found. We discuss the biogeography of the genera with respect to their phylogenetic relationships. California is the state with the greatest alpha diversity (89 species, 46.6% of taxa) and unique species (35 species, 18.3% of taxa). Texas, Arizona, Colorado and Utah are the states with the next greatest alpha diversity with Texas, Arizona and Utah being next for unique species diversity. Maine, New Hampshire and Rhode Island are the states with the least amount of cicada diversity. Diversity is greatest in states and areas where there is a diversity of plant communities and habitats within these communities. Mountainous terrain also coincides with increases in diversity. Several regions of the focus area require additional collection efforts to fill in the distributions of several species.
Keywords:
cicada; distribution; Diceroprocta; Tibicen; Okanagana; Okanagodes; Cacama; Magicicada; Platypedia; Cicadetta

1. Introduction

Cicadas are large, generally conspicuous insects in their environments. The calling songs of the males are thought of as a sign of summer and cicadas are part of the popular culture in multiple regions of the world. Even though the presence of cicadas is generally easy to ascertain and they are popular with amateur entomologists, there is still much to be determined with respect to the diversity and distributions of cicada species in the world. For example, over the past decade we and our colleagues have expanded our knowledge of the ranges of cicada species throughout the New World while simultaneously increasing our understanding of the known cicada diversity in individual countries by as much as 500% [1,2,3,4,5,6,7,8,9,10,11,12,13,14,15,16,17,18,19,20,21,22,23,24,25] including the first records of cicadas in El Salvador [1]. The uncertainty in the composition of the New World cicada fauna included, until recently, North America, north of Mexico where 17 species were removed from the fauna in the first comprehensive study of the region [26].

The diversity of the cicadas of North America, north of Mexico recently has been determined [26]. The biogeography of individual species has been investigated as part of studies on the biology of specific taxa [19,23,24,27,28,29,30,31,32] but a complete review of the biogeography of the North American cicadas has not been produced. The distributions of cicadas for a limited number of states have been published but more recent analyses have adding to the often dated and incomplete cicada diversity within those states [21,33,34,35,36,37,38,39,40,41,42,43,44,45,46,47,48,49,50].

This study is the summary of our field research and the data obtained from the labels of cicada specimens stored in more than 110 institutional collections. We illustrate the distribution of each species currently known to inhabit North America, north of Mexico. We attempt to associate the distributions with phytogeographic regions if possible and host plants if the data exist since cicadas are often limited in their distribution by the phytogeographic region they inhabit [18,21,27,29,30,31,49,51,52,53,54,55,56,57]. We have also elevated subspecies to species rank in part based on geographic separations [27,29,30]. These data will provide an historical perspective on the distribution of cicadas from the region so changes due to human activity can be monitored and species that might be considered for conservation efforts can be identified.

2. Results and Discussion

The cicada fauna of North America, north of Mexico currently consists of 170 species and 21 subspecies [26]. The taxa are distributed among 16 genera, eight tribes, and three subfamilies within the family Cicadidae. The taxonomy and a list of the species currently known to inhabit North America, north of Mexico are summarized in Table 1. The table also contains a summary of the states and/or provinces for which we have records for each species as illustrated in the figures. The genera are discussed following the higher taxonomy listed in Table 1.

Table Table 1. Diversity and distribution of the cicadas of North America, north of Mexico. All species are members of the family Cicadidae. State and provincial abbreviations follow the postal code system for the USA and Canada.

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Table 1. Diversity and distribution of the cicadas of North America, north of Mexico. All species are members of the family Cicadidae. State and provincial abbreviations follow the postal code system for the USA and Canada.
SubfamilyTribeGenusSpeciesDistribution
CicadinaeCryptotympaniniCacamaC. californica DavisCA, NV
C. collinaplaga Sanborn & HeathTX
C. crepitans (Van Duzee)CA
C. moorei Sanborn & HeathAZ, NV
C. valvata (Uhler)AZ, CA, CO, KS, NV, NM, OK, TX, UT
C. variegata DavisTX
DiceroproctaD. apache (Davis)AZ, CA, CO, NV, UT
D. arizona (Davis)AZ
D. aurantiaca DavisNM, TX
D. averyi DavisTX
D. azteca (Kirkaldy)KS, OK, TX
D. bequaerti (Davis)AL, AR, LA, MS, OK, TX
D. bibbyi DavisTX
D. biconica (Walker)FL
D. canescens DavisTX
D. cinctifera cinctifera (Uhler)NM, TX
D. cinctifera limpia DavisTX
D. cinctifera viridicosta DavisTX
D. delicata (Osborn)LA, TX
D. eugraphica (Davis)AZ, CO, KS, NM, OK, TX
D. knighti (Davis)AZ
D. lata DavisTX
D. marevagans DavisTX
D. olympusa (Walker)AL, FL, GA, MS, NC, SC
D. semicincta (Davis)AZ, NM
DiceroproctaD. swalei swalei (Distant)AZ
D. swalei davisi DavisAZ
D. texana (Davis)TX, NM
D. viridifascia (Walker)AL, FL, GA, LA, MS, NC, SC, VA
D. vitripennis (Say)AL, AR, IL, IN, KS, KY, LA, MI, MS, MO, NE, OK, TN, TX, WI
Tibicen (eastern species)T. auriferus (Say)AR, KS, MO, NE, NM, OK, TX
T. canicularis (Harris)AR, CT, DC, IL, IN, IA, KS, ME, MB, MD, MA, MI, MN, MO, NE, NB, NH, NJ, NY, NC, ND, NS, OH, ON, PA, PE, QC, RI, SC, SD, TN, VT, VA, WV, WI
T. davisi davisi (Smith & Grossbeck)AL, DE, DC, FL, GA, LA, MD, MA, MS, NJ, NY, NC, PA, SC, TN, TX, VA, WV
T. davisi hardeni DavisAR, MS
T. latifasciatus (Davis)MD, NJ, NC, VA
T. linnei (Smith & Grossbeck)AL, AR, CT, DE, DC, FL, GA, IL, IN, IA, KS, KY, LA, ME, MD, MA, MI, MN, MS, MO, NE, NJ, NY, NC, OH, ON, PA, SC, TN, VT, VA, WV, WI
T. lyricen lyricen (Degeer)AL, AR, CT, DE, DC, FL, GA, IL, IN, IA, KS, KY, LA, MD, MA, MI, MS, MO, NE, NH, NJ, NY, NC, OH, OK, ON, PA, RI, SC, TN, TX, VA, WV, WI
T. lyricen engelhardti (Davis)AL, CT, DE, DC, FL, GA, IN, KY, MD, MA, MS, NJ, NY, NC, OH, PA, RI, SC, TN, VA, WV
TibicenT. lyricen virescens DavisFL, GA, NC, SC
T. pruinosus pruinosus (Say)AL, AR, CO, FL, GA, IL, IN, IA, KS, KY, LA, MA, MD, MI, MN, MS, MO, NE, NJ, NY, NC, OH, OK, PA, SC, SD, TN, TX, VA, WV, WI
T. pruinosus fulvus BeamerKS, OK
T. robinsonianus DavisAL, AR, DC, FL, GA, IN, KS, MD, MS, MO, NC, OH, PA, TN, TX, VA
T. similaris (Smith & Grossbeck)AL, FL, GA, LA, MS, NC, SC
T. superbus (Fitch)AR, KS, LA, MO, NM, OK, TX
T. tibicen tibicen (Linnaeus)AL, AR, CT, DE, DC, FL, GA, IL, IN, IA, KS, KY, LA, MD, MA, MI, MS, MO, NE, NJ, NY, NC, OH, OK, PA, RI, SC, SD, TN, TX, VT, VA, WV, WI
T. tibicen australis (Davis)AL, FL, GA, MS, NC, SC, TN, VA
T. winnemanna (Davis)AL, AR, DE, DC, GA, IN, KY, LA, MD, MS, NC, OH, OK, PA, SC, TN, TX, VA, WV
Tibicen (western species)T. bifidus (Davis)AZ, CO, KS, NM, OK, TX, UT
T. chiricahua DavisAZ, NM
T. chisosensis DavisTX
T. duryi DavisAZ, CO, NM, OK, TX, UT
T. inauditus DavisAZ, CO, NM, OK, TX
T. longiperculus DavisAZ
T. parallelus DavisAZ
T. simplex DavisAZ
T. texanus MetcalfNM, OK, TX
T. townsendii (Uhler)AZ, NM, TX
Tibicen (large species)T. auletes (Germar)AL, AR, CT, DE, DC, FL, GA, IL, IN, IA, KS, KY, LA, MD, MA, MI, MS, MO, NE, NJ, NY, NC, OH, OK, PA, SC, TN, TX, VA, WV, WI
T. cultriformis (Davis)AZ, NM
T. dealbatus (Davis)CO, IA, KS, MT, NE, NM, ND, OK, SD, TX, WY
T. dorsatus (Say)AR, CO, ID, IL, IA, KS, MO, MT, NE, NM, OK, SD, TX, WY
T. figuratus (Walker)AL, AR, FL, GA, LA, MS, NC, SC, TN, TX, VA
T. pronotalis pronotalis DavisIA, MO, NE, ND, OK, SD
T. pronotalis walkeri MetcalfAL, AR, FL, GA, IL, IN, IA, KS, KY, LA, MD, MI, MN, MS, MO, NE, NC, ND, OH, OK, SD, TN, TX, VA, WV, WI, WY
T. resh (Haldeman)AR, KS, LA, MS, NE, OK, SC, TN, TX
T. resonans (Walker)AL, FL, GA, LA, MS, NC, SC, TN, TX, VA
T. tremulus ColeCO, KS, NE, OK, TX
CornupluraC. nigroalbata DavisAZ
FidicininiBeameriaB. ansercollis Sanborn & HeathUT
B. venosa (Uhler)AZ, AR, CO, IA, KS, MO, NE, NM, OK, TX, UT
B. wheeleri DavisAZ, NM
PacarinaP. puella DavisAZ, LA, OK, TX
P. shoemakeri Sanborn & HeathAZ, CO, NM, TX
HyantiiniQuesadaQ. gigas (Olivier)TX
CicadiniNeocicadaN. chisos (Davis)TX
N. hieroglyphica hieroglyphica (Say)AL, AR, DE, FL, GA, IL, IN, KS, KY, LA, MD, MS, MO, NJ, NY, NC, OH, OK, SC, TN, TX, VA
N. hieroglyphica johannis (Walker)AL, FL, GA, NC, SC
CicadettinaeTaphuriniMagicicadaM. cassinii (Fisher)GA, IA, IL, IN, KS, KY, MD, MO, NC, NE, NJ, NY, OH, OK, PA, TN, TX, VA, WI, WV
M. neotredecim Marshall & CooleyAR, IA, IL, IN, KY, MO, TN
M. septendecim (Linnaeus)CT, DC, DE, GA, IA, IL, IN, KS, KY, MA, MD, MI, MO, NC, NE, NJ, NY, OH, PA, RI, SC, TN, VA, WI, WV
M. septendecula Alexander & MooreGA, IA, IL, IN, KS, KY, MO, NC, NJ, NY, OH, PA, TN, VA, WV
M. tredecassini Alexander & MooreAL, AR, GA, IA, IL, IN, KY, MD, MO, MS, NC, OK, SC, TN, VA
M. tredecim (Walsh & Riley)AL, AR, GA, IL, IN, KY, LA, MD, MO, MS, NC, OK, SC, TN, VA
M. tredecula Alexander & MooreAL, AR, GA, IA, IL, IN, KY, LA, MO, MS, NC, OK, SC, TN, VA
CicadettiniCicadettaC. calliope calliope (Walker)AL, AR, CO, FL, GA, IL, IN, IA, KS, KY, LA, MD, MS, MO, NE, NC, OH, OK, SC, SD, TN, TX, VA
C. calliope floridensis (Davis)AL, FL, GA
C. camerona (Davis)TX
C. kansa (Davis)CO, KS, LA, NE, NM, OK, TX
C. texana (Davis)TX
TibicininaePlatypediiniPlatypediaP. affinis DavisUT
P. aperta DavisCA
P. areolata (Uhler)BC, CA, ID, MT, NV, OR, UT, WA, WY
P. balli DavisAZ
PlatypediaP. barbata DavisCA
P. bernardinoensis DavisAZ, CA, CO, NM
P. falcata DavisAZ, CA, OR, TX, WA
P. intermedia Van DuzeeCA, NV
P. laticapitata DavisCA
P. mariposa DavisCA
P. middlekauffi SimonCA
P. minor UhlerCA, OR
P. mohavensis mohavensis DavisAZ, CA, CO, NV, NM, UT
P. mohavensis rufescens DavisAZ, CA, CO, NV, NM, UT
P. putnami putnami (Uhler)AZ, BC, CA, CO, ID, MT, NE, NV, NM, OR, SD, UT, WA, WY
P. putnami keddiensis DavisCA
P. putnami lutea DavisAZ, CA, CO, ID, MT, NV, NM, OR, SD, UT, WA, WY
P. putnami occidentalis DavisCA
P. rufipes DavisCA
P. scotti DavisCA
P. similis DavisCA, OR
P. sylvesteri SimonsCA
P. tomentosa DavisCA
P. usingeri SimonsCA
P. vanduzeei DavisCA
NeoplatypediaN. ampliata (Van Duzee)CA, OR
N. constricta DavisAZ, CA, CO, ID, NV, NM OR, UT
TibicininiClidophlepsC. beameri DavisCA
C. blaisdelli (Uhler)AZ, CA
ClidophlepsC. distanti distanti (Van Duzee)CA, NV
C. distanti truncata (Van Duzee)CA
C. rotundifrons (Davis)AZ, CA
C. tenuis DavisCA, NV
C. vagans DavisAZ, CA, NV
C. wrighti DavisCA
OkanaganaO. annulata DavisCA, CO
O. arboraria WymoreCA
O. arctostaphylae Van DuzeeCA
O. aurora DavisCA, NV
O. balli DavisIL, IA, KS, MN, NE, ND, SD, WI
O. bella DavisAB, AZ, BC, CA, CO, ID, MT, NV, NM, OR, SD, UT, WA, WY
O. canadensis (Provancher)AB, BC, CA, CO, ID, ME, MB, MI, MN, MT, NB, NH, NY, NT, OH, ON, OR, PA, QC, SK, SD, UT, VT, WI
O. canescens Van DuzeeCA, WA
O. catalina DavisCA
O. cruentifera (Uhler)CA, ID, NV, OR, UT, WY
O. ferrugomaculata DavisCA, CO, ID, OR, WA
O. formosa DavisNV, UT
O. fratercula DavisAB, BC, CA, CO, ID, MT, NV, OR, WA, WY
O. fumipennis DavisAZ, CO, NM, UT
O. georgi Heath & SanbornAZ
O. gibbera DavisCA, CO, ID, MT, NV, NM, OR, UT, WA
O. hesperia (Uhler)AZ, CA, CO, KS, MT, NE NV, NM, OK, SD, TX, UT, WY
OkanaganaO. hirsuta DavisCA
O. lurida DavisBC, ID, OR, WA
O. luteobasalis DavisAB, BC, CA, CO, ID, MB, MT, NV, ND, OR, UT, WA, WY
O. magnifica DavisAZ, CA, CO, NV, NM, OR, UT, WY
O. mariposa mariposa DavisAZ, CA, NM, OR, UT
O. mariposa oregonensis DavisOR
O. napa DavisCA, ID, WA
O. nigriviridis DavisCA
O. nigrodorsata DavisCA, NV, WA
O. noveboracensis (Emmons)NY, ON
O. occidentalis (Walker)AZ, BC, AB, CA, CO, ID, MT, NV, NM, OR, UT, WA
O. opacipennis DavisCA
O. oregona DavisCA, ID, MT, OR, WA
O. orithyia BlivenCA
O. ornata Van DuzeeBC, CA, ID, NV, OR
O. pallidula DavisCA, NV, OR
O. pernix BlivenCA
O. rhadine BlivenCA
O. rimosa rimosa (Say)AB, BC, CA, CT, ID, IL, IN, IA, ME, MB, MD, MA, MI, MN, MT, NV, NB, NH, NJ, NY, ND, OH, ON, OR, PA, QC, SD, UT, VT, VA, WA, WI, WY
O. rimosa ohioensis DavisOH
O. rubrovenosa DavisAZ, CA, OR, UT
O. salicicola BlivenCA
O. salicicola BlivenCA
O. schaefferi DavisAZ, CO, NV, NM, UT
OkanaganaO. sequoiae BlivenCA
O. simulata DavisCA, OR
O. sperata Van DuzeeCA, NV
O. striatipes (Haldeman)AZ, CA, CO, ID, KS, MT, NE, NV, NM, OR, SD, UT, WY
O. sugdeni DavisUT
O. synodica synodica (Say)AB, AZ, CA, CO, IA, KS, MO, MT, NE, NM, ND, OR, SD, TX, UT, WY
O. synodica nigra DavisAZ, NM, UT
O. tanneri DavisAZ, CO, UT
O. triangulata DavisCA, NV
O. tristis tristis Van DuzeeAZ, CA, NV, OR, WA
O. tristis rubrobasalis DavisCA, NV
O. uncinata Van DuzeeCA
O. utahensis DavisAZ, BC, CA, CO, ID, MT, NV, NM, OR, UT, WA
O. vanduzeei DistantBC, CA, ID, NV, OR, UT, WA
O. vandykei Van DuzeeCA, OR
O. venusta DavisCA
O. villosa DavisCA
O. viridis DavisAR, LA, MS, TX
O. vocalis BlivenCA
O. wymorei DavisCA
O. yakimaensis DavisWA
OkanagodesO. gracilis gracilis DavisAZ, CA, NV, UT
O. gracilis viridis DavisAZ, CA
O. terlingua DavisTX
TibicinoidesT. cupreosparsa (Uhler)CA
T. mercedita (Davis)CA
T. minuta (Davis)CA

The North America species are all representatives of the family Cicadidae [26]. The extant Tettigarctidae are only found in Australia. The species are split into the three subfamilies that comprise the Cicadidae. Half of the genera are found in the Cicadinae which is also the most diverse subfamily worldwide. The Cicadettinae are represented by two genera in North America but are found in greater diversity in Australia, Asia, Europe and Africa. The Tibicininae has its greatest diversity in the New World with additional representatives found in Europe, Africa and Asia. Tribal affiliations of the North American genera illustrate phylogenetic relationships to other regions of the world. The Cryptotympanini (including the genera Cacama, Diceroprocta, Tibicen and Cornuplura from North America, north of Mexico) extends across Australia, Asia, Madagascar, Europe, North America, Central America and South America. The Fidicinini (including the genera Beameria and Pacarina from North America, north of Mexico) contains taxa that are primarily found in South and Central America. The Hyantiini (including the genus Quesada from North America, north of Mexico) extends across South and Central America extending into North America but has more diversity in South America. The Cicadini (including the genus Neocicada from North America, north of Mexico) is primarily composed of Asian species with some genera extending into Europe and North Africa and Central America. The Taphurini (including the genus Magicicada from North America, north of Mexico) includes primarily species from South and Central America, Australia, Asia and Africa. The Cicadettini (including the genus Cicadetta from North America, north of Mexico) is found in Europe, Africa, Asia, and is very diverse in Australia with only a single questionable species from South America. The Platypediini (including the genera Platypedia and Neoplatypedia from North America, north of Mexico) is primarily found in North America, north of Mexico. The Tibicinini (including the genera Clidophleps, Okanagana, Okanagodes and Tibicinoides from North America, north of Mexico) has its greatest species diversity in North America, north of Mexico but also contains species inhabiting Europe, North Africa and Asia.

The species distributions are presented in the figures not only to illustrate their ranges but also to provide additional information with respect to their taxonomic or phylogenetic relationships. Subspecies are placed on the same map in order to illustrate the distribution of each subspecies providing the opportunity to differentiate their ranges. Species that were originally described as varieties were elevated to subspecies rank with the publication of the last edition of the Code [58]. By placing the subspecies for a taxon on the same map, we hope to facilitate the separation of true species from subspecies as has occurred in several cicada species [10,27,29,30,59,60,61,62,63]. We have used distributional patterns to show geographic isolation along with morphological, acoustical and physiological data to elevate cicada subspecies to species rank in the past [10,27,29,30]. Species that form a complex of related species are also illustrated on a single map when possible. All remaining species are grouped together based on the ability to demonstrate the distributions of the collected species without interference. What we call host plants are plants that we have observed cicadas using as food and/or oviposition sites.

2.1. Biogeography of the Cacama Species

Half of the species of the genus Cacama are found in North America, north of Mexico [24]. All but one of the recently described species are represented in Mexico with the range of the genus extending into Central America [6,8,24,26]. The species of Cacama are associated with Opuntia Miller [24,36,49,56,64,65,66,67,68,69] and are often called “cactus dodgers” based on their habits of flying from cactus pad to cactus pad as they perch for a calling bout. Cacama crepitans has also been associated with California sage (Artemisia californica Lessing) [69].

Cacama valvata is the most widespread member of the genus in North America, north of Mexico found in nine states (Figure 1). In contrast, C. crepitans (Figure 1), C. variegata (Figure 1), and C. collinaplaga (Figure 2) are found in a single state with C. collinaplaga being the only species not extending into Mexico. Cacama californica and C. moorei are each found in two states and Mexico (Figure 2).

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Figure 1. Distribution of Cacama crepitans (green triangles), C. valvata (blue circles) and C. variegata (red diamonds).

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Figure 1. Distribution of Cacama crepitans (green triangles), C. valvata (blue circles) and C. variegata (red diamonds).
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Figure 2. Distribution of Cacama californica (blue circles), C. moorei (red squares) and C. collinaplaga (green triangles).

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Figure 2. Distribution of Cacama californica (blue circles), C. moorei (red squares) and C. collinaplaga (green triangles).
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Cacama collinaplaga is found in the Hill Country of central Texas [24] in the desert grasslands phytogeographic region [70]. Cacama moorei is restricted to the Sonoran Desert [24]. Cacama crepitans is found in the California upland forests and woodlands showing a strong affiliation with the Peninsular and Baja Range [71]. Cacama variegata is found in the southern portions of the southeastern coastal plain vegetation [72] and desert grasslands [70]. Cacama californica is found in several phytogeographic regions with specimens collected with the California upland forests and woodlands as well as portions of the Sonoran and Mojave deserts [71]. Cacama valvata has the greatest distribution being found in several plant communities. It has been collected in desert grasslands [70], Great Basin, Chihuahuan, Mojave and Sonoran deserts [56,73], chaparral [74,75], and intermountain deserts [76]. The success of Opuntia colonizing various habitats appears to have facilitated the spread of the species into a wide variety of environments.

2.2. Biogeography of the Diceroprocta species

There are 21 species and three subspecies of the genus Diceroprocta in North America, north of Mexico [26]. The genus appears to have originated in Mexico as this is where the genus exhibits its greatest diversity [8]. The genus is most closely related to Tibicen with many of the species being assigned to or originally described as part of the genus Tibicen [26]. The species of Diceroprocta can be separated into two groups, those whose males have rounded opercula and are primarily found in the central and eastern United States and those whose males have pointed opercula and are found in the southern and western United States and the Florida Keys.

The D. cinctifera species complex is associated with riparian habitats of the Rio Grande River system (Figure 3) with Prosopis L. being a primary host [77]. It has also been associated with cottonwoods (Populus spp.) [66]. Diceroprocta cinctifera limpia is found in the Limpa creek tributary system [77,78] whereas D. cinctifera cinctifera and D. cinctifera viridicosta are found on the Rio Grande proper with D. cinctifera viridicosta found on the southern and eastern portion of the river system [77]. Both D. cinctifera viridicosta [8] and D. cinctifera cinctifera [14] have had their ranges extended into Mexico. Diceroprocta cinctifera cinctifera has been associated with mesquite (Prosopis grandulosa Torrey) and desert willow (Chilopsis linearis (Cav.) Sweet) [56] and we have collected specimens calling from Acacia spp., creosote bush (Larrea tridentate (D.C.) Coville), ocotillo (Fouquieria splendens Engelmann) and Fremont cottonwood (Populus fremontii Watson). We found D. cinctifera limpia calling in cottonwood (Populus spp.), Acacia spp. and both D. cinctifera limpia and D. cinctifera viridicosta from mesquite (Prosopis spp.).

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Figure 3. Distribution of Dicerprocta cinctifera cinctifera (red circles), D. cinctifera limpia (blue diamonds), and D. cinctifera viridicosta (green triangles).

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Figure 3. Distribution of Dicerprocta cinctifera cinctifera (red circles), D. cinctifera limpia (blue diamonds), and D. cinctifera viridicosta (green triangles).
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The other current subspecies pair of the genus is D. swalei swalei and D. swalei castanea (Figure 4). Both subspecies are primarily found in Arizona with the known range of D. swalei swalei recently being extended into Mexico [8]. Both subspecies are found in the Mexican Highlands Section of the Basin and Range Province [73]. We found the nominotypical subspecies calling from Acacia spp. and mesquite (Prosopis spp.) and it has also been associated with ocotillo (Fouquieria splendens), Baccharus spp. and catclaw (Acacia greggii Gray). The significant overlap of the subspecies and nominotypical subspecies suggests the subspecies is a color variation and further analyses should be performed to determine the status of subspecies. The known range of D. canescens (Figure 4) is more limited being found in the drainage plain of Tornillo Creek and south of the Chisos Mountains in Texas. We found the species associated with Larrea tridentata and Prosopis grandulosa [56]. Diceroprocta azteca expands from Mexico across Texas and Oklahoma and has recently been found in Kansas [79]. It inhabits several habitats but is primarily associated with desert grasslands [70]. We found it calling from mesquite (Prosopis spp.) and grasses and it has been associated with various trees, weeds, and bushes [80]. Diceroprocta viridifascia (Figure 4) is restricted to primarily coastal environments (Figure 4) but can be found inland along sandy ridges [21]. It has been reported from the Southern Coastal Plains and Southern Florida Coastal Plain ecoregions [81] within Florida [21]. The species has been associated with grasses [82,83], sea oats (Uniola spp.) [37,84] as well as oaks (Quercus spp.) and other small trees [21]. We found it most often in sand pine (Pinus clausa (Chapm. Ex Engelm.)Sargent) and scrub live oak (Quercus germinata Small).

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Figure 4. Distribution of Dicerprocta swalei swalei (blue circles), D. swalei castanea (red diamonds), D. canescens (light blue stars), D. azteca (orange triangles) and D. viridifascia (green squares).

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Figure 4. Distribution of Dicerprocta swalei swalei (blue circles), D. swalei castanea (red diamonds), D. canescens (light blue stars), D. azteca (orange triangles) and D. viridifascia (green squares).
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Diceroprocta aurantiaca was originally described as a variety of D. delicata [85]. It was elevated to species status in part by the different range of the species from D. delicata [27]. Diceroprocta aurantiaca is found in the desert grasslands and Chihuahuan Desert (Figure 5) [27,70,73]. We found the species associated with Acacia spp. and creosote bush (Larrea tridentata). In contrast, D. delicata inhabits southeastern coastal plain vegetation (Figure 5) [21,72] and is associated with Iva frutescens L. [86]. We collected the species from a variety of plants including mesquite (Prosopis spp.), scrubby plants and grasses. Diceroprocta semicincta is restricted to the Sonoran Desert (Figure 5) [73] being found primarily in the Mexican Highland Section of the Basin and Range Province. It has significant overlap with D. apache (Figure 6) but is generally found in more elevated environments. We found D. semicincta calling from mesquite (Prosopis spp.), catclaw (Acacia greggii), Emory oak (Quercus emoryi Torrey), desert willow (Chilopsis linearis), Utah juniper (Juniperus osteosperma (Torr.)Little) and creosote bush (Larrea tridentata) but has also been associated with ocotillo (Fouquieria splendens) [87].

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Figure 5. Distribution of Dicerprocta aurantiaca (gold circles), D. delicata (blue triangles) and D. semicincta (red diamonds).

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Figure 5. Distribution of Dicerprocta aurantiaca (gold circles), D. delicata (blue triangles) and D. semicincta (red diamonds).
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Figure 6. Distribution of Dicerprocta apache (red circles), D. bibbyi (blue triangles) and D. bequaerti (green squares).

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Figure 6. Distribution of Dicerprocta apache (red circles), D. bibbyi (blue triangles) and D. bequaerti (green squares).
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Diceroprocta apache is found in the Sonoran desert (Figure 6) [52,88] in riparian habitats e.g., [89,90,91] following river systems into Nevada and Utah from Arizona and California. It has been associated with Fremont cottonwood (Populus fremontii) and Goodding willow (Salix gooddingii Ball) [89,91], introduced tamarix (Tamarix chinensis Loureiro), Baccharis spp., Acacia spp., velvet mesquite (Prosopis velutina) [91], undetermined mesquite (Prosopis spp.) [67,92], and palo verde (Cercidium spp.) [66,91,93]. Diceroprocta bibbyi (Figure 6) is found in creosote flats (Larrea tridentata) [56] of western Texas. It was recently described from Mexico [14] and appears to have a more extensive distribution in the Chihuahuan Desert [73]. Diceroprocta bequaerti (Figure 6) is found primarily in the southwestern region of the eastern deciduous forests [94] but extends into the grasslands [70] of the southern central United States. We found it associated with willow (Salix spp.) and grasses but it has also been reported in poplars (Populus deltoids W. Bartram ex Humphrey Marshall), weeds (Dysphania ambrosioides (L.) Mosyakin & Clemants) [95], and black mangroves (Avicennia germinans (L.) L.) [80].

Diceroprocta lata was originally described as a variety of D. texana [96]. It was recently reported from the United States and elevated to species status in part by the different range of the species from D. texana [29]. Diceroprocta texana is associated with Larrea tridentata and Prosopis grandulosa [29] in the Chihuahua desert [73] and desert grasslands [70] phytogeographic regions. Diceroprocta lata is also found in Prosopis glandulosa [29] in desert grasslands [70] but on the edges of the Chihuahuan Desert [73]. Diceroprocta knighti is restricted to the Sonoran Desert (Figure 7) [73] being found primarily in the Mexican Highland Section of the Basin and Range Province. We found it associated with creosote bush (Larrea tridentata), palo verde (Cercidium spp.), mesquite (Prosopis spp.), and ocotillo (Fouguieria splendens).

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Figure 7. Distribution of Dicerprocta lata (blue diamonds), D. texana (green circles) and D. knighti (red squares).

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Figure 7. Distribution of Dicerprocta lata (blue diamonds), D. texana (green circles) and D. knighti (red squares).
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Diceroprocta eugraphica has an extensive distribution (Figure 8) across several phytogeographic regions including grasslands [70], intermountain deserts, and the Chihuahuan and Sonoran deserts [73]. The species has been reported from creosote flats (Larrea tridentata) [56] and Ponderosa pine (Pinus ponderosa Douglas ex C. Lawson)-Pinyon (Pinus edulis Engelmann)-Juniper (Juniperus spp.) woodlands [49] as well as mesquite (Prosopis spp.) [80] and we found it associated with four-wing saltbush (Atriplex canescens (Pursh)Nuttall), sand sage (Artemisia filifolia Torrey), and Acacia spp. in some habitats. The known range of D. eugraphica was recently extended into Mexico [6]. Diceroprocta marevagans is found in coastal environments (Figure 8) of the southeastern coastal plain vegetation [72]. We found it calling from grasses and it has been associated with Amaranthus spp. [41,92]. Diceroprocta olympusa has the most extensive distribution of any cicada in Florida and is found in all Florida ecoregions except the Everglades [21]. The distribution extends with much less frequency to Mississippi and North Carolina (Figure 8). It often calls from trees but uses grasses as a host plant [21,97].

Diceroprocta arizona is another species restricted to the Sonoran Desert (Figure 9) [73] being found primarily in the Mexican Highland Section of the Basin and Range Province. We found it associated with gray oak (Quercus grisea Liebmann). Diceroprocta averyi has two separate populations in desert grasslands [70]. Diceroprocta vitripennis expands further north than any other species of Diceroprocta with specimens being collected as far north as Michigan and Wisconsin (Figure 9). It is found in grassland habitats [70] as well as the western portion of the eastern deciduous forest phytogeographic region [94]. We collected the species in willow (Salix spp.) and grasses and it has also been reported to use cottonwoods (Populus spp.) [98]. Diceroprocta biconica has one of the more limited distributions of cicadas in the United States (Figure 9) but is also known from Cuba [10]. It is restricted to the Southern Florida Coastal Plain ecoregion [81], and is found only in the Florida Keys associated with black mangrove (Avicennia germinans) [21,53], but has also been observed and collected using multiple ornamental tree species [21]. It shows greater morphological similarities to species of Diceroprocta in Cuba and the Bahamas than to the other species of Diceroprocta in Florida [2,10,21].

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Figure 8. Distribution of Dicerprocta eugraphica (blue circles), D. marevagans (green triangles) and D. olympusa (red diamonds).

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Figure 8. Distribution of Dicerprocta eugraphica (blue circles), D. marevagans (green triangles) and D. olympusa (red diamonds).
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Figure 9. Distribution of Dicerprocta arizona (red squares), D. averyi (blue diamonds), D. vitripennis (green circles) and D. biconica (orange triangles).

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Figure 9. Distribution of Dicerprocta arizona (red squares), D. averyi (blue diamonds), D. vitripennis (green circles) and D. biconica (orange triangles).
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2.3. Biogeography of the Tibicen species

The current status of the genus Tibicen is being debated. Boulard [99,100,101,102,103] has argued for the suppression of the genus in favor of Lyristes Horvath. The last proposal to suppress the genus was submitted to the International Commission of Zoological Nomenclature was by Melville and Sims [104]. Further comments were made from several scientists [105,106,107], but no ruling of the commission has occurred. We retain the use of the genus Tibicen since Latrielle [108] specifically mentioned Cicada plebeia (sic) as the only example of the genus and thus unambiguously designated a type species under Article 12.2.5 [58] regardless of the ambiguities that may have occurred since that designation [99,101,102,103,104,105,106,107].

The genus Tibicen is found in the Nearctic, Palaearctic, Oriental, and Neotropical biogeographical regions with the greatest diversity in the Nearctic region [26]. There are 31 species and six subspecies currently in the genus Tibicen in North America, north of Mexico [26]. These species can be split into three groups based on size and the shape of the abdomen [26,60] with preliminary molecular data supporting the three groups [109].

2.3.1. Biogeography of the Eastern Tibicen species

Davis [60] characterized the eastern Tibicen species as large, usually green and black cicadas. These animals appear to be more closely related morphologically to members of the genus in the Oriental region. They inhabit areas in the eastern half of the United States and southern Canada east of the 100th meridian [26]. There are 12 species and five subspecies within this group.

The name T. tibicen was shown to be a senior synonym of a common cicada known as T. chloromerus [110]. The species is known as the swamp cicada as it is the only eastern Tibicen species to be found in low vegetation in swampy or marshy habitats as well as being found in upland meadows and overgrown fields [111]. The nominotypical subspecies expands across much of the eastern United States (Figure 10) with the subspecies T. tibicen australis being found primarily in Florida and Georgia (Figure 10). Tibicen tibicen tibicen and T. tibicen australis are found in forested environments of the Southern Coastal Plains and Southeastern Plains ecoregions [81] within Florida but was surprisingly absent from the Everglades [21]. The nominotypical subspecies has been observed to oviposit in golden rod (Solidago spp.), sumac (Rhus spp.), asters (Aster spp.), and willow shrubs (Salix spp.) with females also observed ovipositing in dead twigs [38,40,43,98]. The significant overlap between the subspecies suggests further analysis is required to determine their status.

The species T. lyricen currently has three described subspecies [26]. All are found in the eastern deciduous forest ecosystems [94]. The nominotypical subspecies has the greatest range (Figure 11) with T. lyricen engelhardti being surrounded by the nominotypical subspecies along the Appalachian Mountains and areas west of the mountains and T. lyricen virescens (Figure 11) found in southeastern states with greatest concentration of collection localities in Florida (Figure 11). Tibicen lyricen lyricen and T. lyricen virescens are found in all Florida ecoregions with representatives collected in the Miami Ridge/Atlantic Coastal Strip of the Southern Florida Coastal Plain ecoregion [81] within Florida [21]. The nominotypical subspecies has been associated with oaks (Quercus spp.), forests that contain maple (Acer spp.) and beeches (Fagus grandifolia Ehrhart) [45], persimmon (Diospyros virginiana L.) [38,112], oaks and peach (Prunus persica (L.) Stokes) [112]. The significant overlap in the nominotypical subspecies and T. lyricen virescens suggests further work should be performed to determine the status of the subspecies.

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Figure 10. Distribution of Tibicen tibicen tibicen (blue circles) and T. tibicen australis (red diamonds).

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Figure 10. Distribution of Tibicen tibicen tibicen (blue circles) and T. tibicen australis (red diamonds).
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Figure 11. Distribution of Tibicen lyricen lyricen (orange circles), T. lyricen engelhardti (blue diamonds) and T. lyricen virescens (green triangles).

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Figure 11. Distribution of Tibicen lyricen lyricen (orange circles), T. lyricen engelhardti (blue diamonds) and T. lyricen virescens (green triangles).
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Tibicen davisi is another species that also has two subspecies. The distribution of the nominotypical subspecies is across much of the southeast extending north into New England and west into Texas (Figure 12). The subspecies T. davisi hardeni is restricted to Mississippi and one collection site in Arkansas (Figure 12). Further analyses should be performed to determine the status of the subspecies. Tibicen davisi davisi is found in all Florida ecoregions except the Everglades and insular regions with representatives collected in the Miami Ridge/Atlantic Coastal Strip of the Southern Florida Coastal Plain ecoregion [81] within the state of Florida [21]. We have found it a variety of trees including mahogany (Swietenia mahagoni (L.) Jacquin) and it has been reported from pine forests (Pinus spp.) [38]. Tibicen auriferus is found in the grasslands of the central plains (Figure 12). We have collected it from grasses, shrubs and small trees in various localities. It has been associated with grassy areas with shrubs including the prairie grass Panicum virgatum L., wood grass (Sorghastrum nutans (L.) Nash) and sumac (Rhus glabra L.).

The final species of Tibicen in North America, north of Mexico to have subspecies is Tibicen pruinosus. The nominotypical subspecies expands across most of the eastern United States in both deciduous forests [94] and grassland [70] ecosystems (Figure 13). The specimen in western Nebraska extends the known range westward beyond what was a recent western expansion of the known range [113]. The species is always associated with trees in the grassland environments, often being found in riparian habitats and is found in high concentration in the towns of the Great Plains regions where trees are common [114]. On a more local scale, T. pruinosus pruinosus is found in the forested interior of the Southern Coastal Plains and Southeastern Plains ecoregions [81] within Florida [21]. The species is associated with hardwood environments and has been observed ovipositing in ash (Fraxinus americana L.), walnut (Juglans nigra L.), elm (Ulmus spp.), maple (Acer spp.), and sumac (Rhus spp.) [98,115]. We also found the species in hackberry (Celtis occidentalis L.) and willows (Salix spp.). The subspecies T. pruinosus fulvus is localized to the southeastern corner of Kansas and eastern Oklahoma (Figure 13) and may represent a local color variation. It is also associated with hardwood environments and has been associated with oak (Quercus spp.), walnut (Juglans nigra), elm (Ulmus spp.), maple (Acer spp.), and sumac (Rhus spp.) [112,116]. Tibicen latifasciatus has been variously classified as a species or subspecies of T. pruinosus [26]. Tibicen latifasciatus is found in coastal environments in the mid-Atlantic region (Figure 13). The geographic separation from T. pruinosus supports the species status for T. latifasciatus through geographic isolation. Tibicen latifasciatus has been associated with cedar (probably Chamaecyparis thyoides (L.)Britton, Sterns & Poggenburg), pines (Pinus spp.), oaks (Quercus spp.) and hollies (Ilex opaca Aiton) [40].

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Figure 12. Distribution of Tibicen davisi davisi (blue circles), T. davisi hardeni (red triangles) and T. auriferus (green diamonds).

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Figure 12. Distribution of Tibicen davisi davisi (blue circles), T. davisi hardeni (red triangles) and T. auriferus (green diamonds).
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Figure 13. Distribution of Tibicen pruinosus pruinosus (blue circles), T. pruinosus fulva (orange diamonds) and T. latifasciatus (green triangles).

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Figure 13. Distribution of Tibicen pruinosus pruinosus (blue circles), T. pruinosus fulva (orange diamonds) and T. latifasciatus (green triangles).
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Tibicen winnemanna is another species that has been classified as a variety of T. pruinosus [26]. It is generally found in areas of the southeast and south (Figure 14) within the eastern deciduous forests [94] where T. pruinosus pruinosus is absent (Figure 13) although there is some overlap. We have seen the species using several species of hardwoods as hosts including maple (Acer spp.), oak (Quercus spp.) and sycamore (Platanus occidentalis L.). Tibicen linnei is one of the more common species being found in both eastern deciduous forests [94] and grassland [70] ecosystems. Similar to T. pruinosus pruinosus, the species is found in trees within the grassland environments. On a more local scale, T. linnei is found in forested environments of the Southern Coastal Plains ecoregion [81] within Florida [21]. The species has been associated with slippery elm (Ulmus rubra Muhlenberg) [117], maple (Acer spp.) [118] and other trees [21,40,45,46,98] but has been reported to avoid pines (Pinus spp.) [40].

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Figure 14. Distribution of Tibicen winnemanna (blue circles) and T. linnei (green squares).

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Figure 14. Distribution of Tibicen winnemanna (blue circles) and T. linnei (green squares).
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The species Tibicen canicularis is named for the dog days of summer and has a wide distribution across the northern United States and southern Canada (Figure 15). Like Tibicen linnei and T. pruinosus pruinosus it is a common species being found in both eastern deciduous forests [94] and grassland [70] ecosystems. We have seen T. canicularis use multiple species of hardwoods as hosts including oaks (Quercus spp.) and elm (Ulmus spp.). No specific tree species seems to be identifiable as a host and they have been reported to use various trees [119] including oak (Quercus spp.) [38] and pine (Pinus spp,) forests [40]. Tibicen similaris is found in the vegetation of the southeastern coastal plain [72]. On a more local scale, T. similaris is found in forested environments of the Southern Coastal Plains and Southeastern Plains ecoregions [81] within Florida [21]. We found it calling from multiple large deciduous trees and it has been associated with turkey oak (Quercus laevis Walter) [34].

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Figure 15. Distribution of Tibicen canicularis (gold circles) and T. similaris (blue diamonds).

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Figure 15. Distribution of Tibicen canicularis (gold circles) and T. similaris (blue diamonds).
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Tibicen robinsonianus is found across the central region of the eastern deciduous forests [94] (Figure 16). The habits of singing high in trees exhibited by a species with low population densities may explain the spacing seen in the distribution. Tibicen robinsonianus inhabits the Southeastern Plains ecoregion [21] and, on a more local scale, the Gulf Coast Flatwoods division of the Southern Coastal Plains ecoregion [81] within Florida. The species has been associated with oaks (Quercus spp.), locusts (Robinia spp.), and cedar (probably Juniperus virginiana L.) [38,78,87,120]. We found them associated with oaks and various other deciduous trees. The final eastern Tibicen species, T. superbus, is found in trees within grassland environments primarily in eastern Texas and Oklahoma but spreads into neighboring states. We found it in a variety of trees, both native and ornamentals [28] including elm (Ulmus spp.), oak (Quercus spp.), juniper (Juniperus spp.) and mesquite (Prosopis spp.).

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Figure 16. Distribution of Tibicen robinsonianus (blue squares) and T. superbus (green circles).

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Figure 16. Distribution of Tibicen robinsonianus (blue squares) and T. superbus (green circles).
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2.3.2. Biogeography of the Western Tibicen Species

The western Tibicen species differ from the other two groups of the genus in that the sides of the abdomen are conspicuously parallel to each other instead of gently tapering. Their coloration is also generally black with orange or reddish markings. Davis [60] suggested that this group more closely resembled Tibicen plebejus and thus is more closely related to Palearctic members of the genus. These animals inhabit areas in the southwestern United States [26]. There are 10 species within this group.

Tibicen bifidus is found in desert grasslands [30,70] and intermountain deserts [76] (Figure 17). It has been associated with yucca (no species given) [98], sage (Artemisia L.) [66,98], fineleaf hymenopappus (Hymenopappus filifolius Hooker) [121], Atriplex L., willows (Salicaceae) [49], Acacia Miller and Juniperus L. [30]. We also found it associated with Mexican Mormon tea (Ephedra trifurcata Torrey ex S. Watson). Tibicen simplex was originally described as a variety of T. bifidus [96] but was elevated to species status in part based on the distribution of the species [30]. Tibicen simplex is also found in desert grasslands [30,70] but in the Mexican Highland Section of the Basin and Range Province of the Sonoran Desert [73] (Figure 17). Tibicen simplex is associated with soaptree yucca (Yucca elata Engelmann) and females were observed to oviposit in Mexican Mormon tea (Ephedra trifurca) [30]. Tibicen texanus is found in trees within the grassland habitats [70] (Figure 17). It has been associated with junipers (Juniperus spp.) and oaks (Quercus spp.) [78]. We collected the species in these trees as well as mesquite (Prosopis spp.). Tibicen chisosensis is found only in the high Chisos Mountains of Texas at elevations above 1980 m (6000 ft) in Mexican piñon pine (Pinus cembroides Zuccarini) [56] but would also call from piñon pine (Pinus edulis) and Texas madrone (Arbutus xalapensis Kunth). It would have the most restricted range of any species in North America, north of Mexico except that the known range of the species was recently extended into Mexico [8].

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Figure 17. Distribution of Tibicen bifidus (blue squares), T. simplex (red circles), T. texanus (orange diamonds) and T. chisosensis (green triangles).

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Figure 17. Distribution of Tibicen bifidus (blue squares), T. simplex (red circles), T. texanus (orange diamonds) and T. chisosensis (green triangles).
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Tibicen chiricahua is found in the Mexican Highland Section of the Basin and Range Province of the Sonoran Desert [73] and intermountain woodlands [76] (Figure 18). The known range of T. chiricahua was recently extended into Mexico [8]. It has been associated with pines (Pinus spp.), juniper (Juniperus spp.) and grasses [87] and we found it in mountain mahogany (Cercocarpus spp.) and one seed juniper (Juniperus monosperma (Engelmann) Sargent). Tibicen townsendii is found in desert grasslands [70], intermountain deserts [76], and the Chihuahuan Desert [73]. Tibicen townsendii was reported from lower desert mountain vegetation associated with soaptree yucca (Yucca elata) in the Chisos Mountains of Texas [56].

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Figure 18. Distribution of Tibicen chiricahua (gold circles) and T. townsendii (blue diamonds).

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Figure 18. Distribution of Tibicen chiricahua (gold circles) and T. townsendii (blue diamonds).
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Tibicen duryi occurs across much of the southwest in a variety of habitats. It has been recorded from chaparral [74,75], Chihuahuan Desert, Mexican Highland Section of the Sonoran Desert [73], and intermountain woodlands [76]. The species is only found with piñon pine (Pinus edulis) [49,67,122] or Mexican piñon pine (Pinus cembroides) [56]. Tibicen longioperculus is found in the Mexican Highland Section of the Basin and Range Province of the Sonoran Desert [73] (Figure 19). It has been collected only in two locations in southern Arizona. We collected it only in the area of Sunnyside Canyon where alligator juniper (Juniperus deppeana Steudel) is found even though there are other species of juniper in other parts of the canyon.

Tibicen parallelus is found in the Mexican Highland Section of the Basin and Range Province of the Sonoran Desert [73] (Figure 20) with the known range of T. parallelus recently being extended into Mexico [6]. The extra limital point on the map at Albuquerque, New Mexico is the type locality for the species. It may be that the individual who sent the specimens to Davis was from Albuquerque and this mistakenly became the type locality even though this location is distant from the remaining collection sites. We found the species associated with oaks (Quercus spp.) as has been reported previously [87] but also collected specimens from alligator juniper (Juniperus deppeana). Tibicen inauditus is found in trees in desert grasslands [70], intermountain deserts [76], and the Chihuahuan Desert [73]. It has been associated with skunk bush (Rhus trilobata Nuttall), soapweed (Yucca glauca Nuttall), bear grass (Xerophyllum tenax (Pursh)Nuttall) [80], oak (Quercus spp.) in various habitats [56,78,123]. We also collected specimens in mesquite (Prosopis spp.), juniper (Juniperus spp.) and mountain mahogany (Cercocarpus spp.).

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Figure 19. Distribution of Tibicen duryi (red circles) and T. longioperculus (blue diamonds).

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Figure 19. Distribution of Tibicen duryi (red circles) and T. longioperculus (blue diamonds).
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Figure 20. Distribution of Tibicen parallelus (green circles) and T. inauditus (purple squares).

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Figure 20. Distribution of Tibicen parallelus (green circles) and T. inauditus (purple squares).
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2.3.3. Biogeography of the large Tibicen species

The large Tibicen species are the largest of the genus. The mesonotum is generally decorated with two pipe shaped marks that are similar to the Hebrew letter resh [60]. These species inhabit areas in most of the United States except the states of the Pacific coast, northwestern Rocky Mountains and Northeast. There are nine species and two subspecies within this group.

The final subspecies within the genus Tibicen is found in this group. The nomenclature has been confused [26] but was recently analyzed and reorganized based on the Code [50]. Tibicen pronotalis pronotalis is found in the northern Great Plains (Figure 21). We found the nominotypical subspecies associated with various isolated trees and riparian habitats including willows (Salix spp.). Tibicen pronotalis walkeri is found in the grassland habitats [70] and eastern deciduous forests [94] west of the Appalachians (Figure 21). We found this subspecies singing in tall trees, including cottonwoods (Populus spp.), and in riparian habitats. It has been reported to be associated with willow (Salix spp.), cottonwoods (Populus spp.), birch (Betula spp.) and maple (Acer spp.) [98,112].

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Figure 21. Distribution of Tibicen pronotalis pronotalis (blue diamonds) and T. pronotalis walkeri (red circles).

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Figure 21. Distribution of Tibicen pronotalis pronotalis (blue diamonds) and T. pronotalis walkeri (red circles).
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Tibicen dealbatus occurs across the grasslands ecosystem [70] across the central United States (Figure 22). The species is always associated with trees in the grassland environments often being found in riparian habitats [49,124]. We found it associated with willow (Salix spp.) and cottonwoods (Populus spp.) as previously reported [98,124] but also with ornamentals and nut trees in orchards. It has also been reported in high concentrations in introduced salt cedar (Tamarix ramosissima Ledebour) [49]. Tibicen figuratus is found in the vegetation of the southeastern coastal plain [72]. At a more local scale, T. figuratus is found in forested environments of the Southern Coastal Plains and Southeastern Plains ecoregions with representatives collected in the Miami Ridge/Atlantic Coastal Strip of the Southern Florida Coastal Plain ecoregion [81] within Florida [21].

Tibicen dorsatus extends across the grasslands [70] ecosystem of the central United States (Figure 23). The species is usually associated with shrubs in the grassland environments. We found it associated with sage (Artemisia spp.), squawbush sumac (Rhus trilobata) and various shrubs with the occasional hardwood used as a calling perch. It has also been associated with four-wing saltbush (Atriplex canescens), and rabbitbrush (Ericameria nauseosa (Pallas ex. Pursh)G.L. Nesom & Baird) [49], cottonwoods (Populus spp.) [125], willows (Salix spp.), elms (Ulmus spp.) [112], bluestem grass (Andropogon spp.) [126], sumac (Rhus spp.) and goldenrod (Solidago spp.) [43]. Tibicen resonans is found in the vegetation of the southeastern coastal plain [72]. On a more local scale, T. resonans is found in forested environments in all Florida ecoregions except the Everglades with representatives collected in the Miami Ridge/Atlantic Coastal Strip of the Southern Florida Coastal Plain ecoregion [81] within Florida [21]. We collected specimens from longleaf pine (Pinus palustris Miller) and slash pine (Pinus elliottii Engelmann) in the panhandle of Florida.

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Figure 22. Distribution of Tibicen dealbatus (orange circles) and T. figuratus (blue triangles).

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Figure 22. Distribution of Tibicen dealbatus (orange circles) and T. figuratus (blue triangles).
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Figure 23. Distribution of Tibicen dorsatus (green circles) and T. resonans (red squares).

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Figure 23. Distribution of Tibicen dorsatus (green circles) and T. resonans (red squares).
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Tibicen auletes has the most eastern distribution of the large Tibicen species (Figure 24) being found over most of the eastern deciduous forest ecosystem [94]. Within Florida [21], it has been reported from the Southern Coastal Plains and Southeastern Plains ecoregions [81]. The species has been associated with oaks in general (Quercus spp.), post oak (Quercus stellata Wangenheim), black oak (Quercus velutina Lambert), white oak (Quercus alba L.), and willows (Salix spp.), and other hardwood habitats [38,46,98,112,126]. Tibicen tremulus is found in the Great Plains [70]. We found it associated with sage (Artemisia spp.).

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Figure 24. Distribution of Tibicen auletes (blue circles) and T. tremulus (red triangles).

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Figure 24. Distribution of Tibicen auletes (blue circles) and T. tremulus (red triangles).
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Tibicen cultriformis is found in the Mexican Highland Section of the Basin and Range Province of the Sonoran Desert [73] (Figure 25) with its range recently being extended into Mexico [8]. We found T. cultriformis in riparian habitats congregating in cottonwood (Populus spp.) and willow (Salix spp.) [127]. It also has been associated with black willow (Salix nigra Marshall), ash (Fraxinus spp.), sycamore (Platanus wrightii) and walnut (Juglans major) [78]. Tibicen resh is found in the grassland habitats [70] and western regions of the eastern deciduous forests [94] (Figure 25). It overlaps the extremes of the ranges of and fills the gap between the generally more western T. dorsatus and T. dealbatus and generally the more eastern T. auletes (see Figure 22, Figure 23, Figure 24, Figure 25). We found the species singing in trees, including white ash (Fraxinus americana) and willows (Salix spp.), and in riparian habitats. Tibicen resh has been associated with oaks (Quercus spp.) in general [128] and blackjack oak (Quercus marilandica Muenchhausen) [112] specifically. A specimen of Tibicen resh from South Carolina was verified during this work. However, it is separated from the other specimens by several hundred km and may have been mislabeled.

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Figure 25. Distribution of cultriformis (orange squares) and T. resh (green circles).

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Figure 25. Distribution of cultriformis (orange squares) and T. resh (green circles).
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2.4. Biogeography of the Cornuplura species

The genus Cornuplura is primarily distributed through Mexico [8,26,31]. Davis [129] placed the genus between Diceroprocta and Tibicen. The species are larger than but most similar to the western Tibicen species [26]. Cornuplura species are distributed in central and southern Mexico with the known range of one species extending into the region of interest for this work [8,31].

A single species, C. nigroalbata, inhabits North America, north of Mexico. The species was originally described from southern Arizona [129] but the known range has been extended to southern Sonora, Mexico [8]. The distribution of the species has its northern limit in the southern portion of Arizona (Figure 26) and is known only from Santa Cruz and Cochise counties [31]. The species is associated with Emory oak (Quercus emoryi) and Mexican blue oak (Quercus oblongifolia Torrey) [31] in the oak-juniper woodland [130] within the Mexican Highlands Desert Section of the Basin and Range floristic province of North America [73].

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Figure 26. Distribution of Cornuplura nigroalbata (blue triangles) and Quesada gigas (red squares).

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Figure 26. Distribution of Cornuplura nigroalbata (blue triangles) and Quesada gigas (red squares).
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2.5. Biogeography of the Beameria species

The genus Beameria was erected for two species of grass dwelling cicadas of the central and western United States [26] with a third species being described recently [23]. The genus is primarily found in North America, north of Mexico but two of the three species have been reported from Mexico [6,8]. The genus represents the northernmost limit of the genera within the tribe Fidicinini that extends southward through Central and South America to Argentina [4,5,7,8,9,13,14,15,17,20,22,131,132].

All species of Beameria are associated with grasses [23,36,56,80]. Beameria venosa has the greatest distribution being found throughout much of 11 states and Mexico (Figure 27). This species can be found in intermountain deserts [76], Chihuahuan Desert [56,73], chapparal [74,75], and grasslands [70]. Beameria ansercollis is known from San Juan County, Utah (Figure 27) in the intermountain desert and steppe habitat [76] where we found it associated with Atriplex spp. and Indian rice grass (Achnatherum hymenoides Ricker ex Piper). Beameria wheeleri is found in the Sonoran Desert [73] of southern New Mexico, southeastern Arizona and northern Mexico (Figure 27). Only B. wheeleri can be associated with a definitive host plant. The distribution of B. wheeleri is dependent on the presence of grasses of the genus Sporobolus R. Brown. Two species of grass with which we found B. wheeleri to be associated were S. wrightii Munro ex Scribner and S. airoides (Torrey)Torrey.

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Figure 27. Distribution of Beameria venosa (blue circles), B. wheeleri (green diamonds) and B. ansercollis (red triangles).

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Figure 27. Distribution of Beameria venosa (blue circles), B. wheeleri (green diamonds) and B. ansercollis (red triangles).
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2.6. Biogeography of the Pacarina Species

The genus Pacarina is found primarily in Central America and Mexico [5,8,14,32] and is related to genera that inhabit Central and South America. Two species of the genus inhabit North America, north of Mexico [26,32]. References to Pacarina championi (Distant) in the literature from the region are considered erroneous [32].

The two species of Pacarina in North America, north of Mexico were first distinguished based on host plants differences [32,133]. We found P. puella associated with mesquite (Prosopis spp.) and it has been recorded from Arizona, Louisiana, Oklahoma and Texas extending into Central America [5,8,14,32] (Figure 28). We also found the species calling extensively from Acacia spp. Pacarina shoemakeri is associated with juniper (Juniperus spp.) and has been recorded from Arizona, Colorado, New Mexico and Texas in chaparral [49,74,75], intermountain woodlands [76] and elevated regions of the Chihuhuan desert [73]. It is likely that the species will also be found in Mexico. We found P. shoemakeri in Juniperus ashei Buchholz in Arizona.

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Figure 28. Distribution of Pacarina puella (blue circles) and P. shoemakeri (red diamonds).

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Figure 28. Distribution of Pacarina puella (blue circles) and P. shoemakeri (red diamonds).
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2.7. Biogeography of the Quesada Species

The genus Quesada has an extensive distribution within the New World. Although only two species occur in the genus, the distribution extends from central Argentina to Texas [26,129]. It is the only representative of the tribe Hyantiini in North America, north of Mexico. The remaining genera of the tribe are found in Central and South America [17,134].

Quesada gigas is the only species of the genus Quesada found in North America, north of Mexico. The species may exhibit the greatest latitudinal gradient of any cicada. It has been collected from central Texas (Figure 26) to central Argentina [22,26,129,135]. It appears to have a generalist approach to host plants that facilitates its use of multiple plant communities [22]. For example, in a single park in Brazil, the species was found to use 17 different families as host plants [136]. In Argentina we found Q. gigas in the tropical cloud forests of the Yunga and tropical rainforests of the Paranense floristic provinces as well as the temperate Chaco and Espinal floristic provinces [22]. The species disappears in Argentina when potential host trees disappear from the floristic provinces. It is also an endothermic species [135]. The endothermy may permit Q. gigas to colonize a greater variety of habitats. The species is associated with the vegetation of the southeastern coastal plain in Texas [72]. The northern limit of the distribution extends to the grasslands in central Texas [70], similar to that of Argentina.

2.8. Biogeography of the Neocicada Species

The genus Neocicada has its greatest diversity in Mexico and Central America [19]. It was reassigned to the subtribe Leptopsaltriina based on molecular evidence [137]. The other genera of this subtribe are distributed in Asia and India so further work appears to be necessary to determine its phylogenetic relationships. Two species and one subspecies inhabit North America, north of Mexico [19,26].

All species of Neocicada inhabiting North America, north of Mexico are associated with oaks (Quercus spp.) [19,43,56,78,82,87,98]. The general coloration patterns of the species make them difficult to see on oaks. Neocicada hieroglyphica hieroglyphica and N. hieroglyphica johannis inhabit much of the eastern United States with N. hieroglyphica johannis restricted to the southeastern United States (Figure 29). They inhabit the eastern deciduous forests [94] and vegetation of the southeastern coastal plains [72] and we found both subspecies associated with live oak (Quercus virginiana Miller) in Florida and the nominotypical subspecies in post oak (Quercus stellata) and black oak (Quercus velutina) with additional records from blackjack oak (Quercus marilandica) [98]. The nominotypical subspecies has been collected in all Florida ecoregions [81], N. hieroglyphica johannis is found in the Tallahasee Hills/Volusia Limesink of the Southeastern Plain ecoregion, and both subspecies are found in the Miami Ridge/Atlantic Coastal Strip of the Southern Florida Coastal Plain in southern Florida [21]. Further analysis of songs and morphology should be performed to determine the status of the subspecies since there is significant overlap in Alabama, Florida, Georgia, and the Carolinas [21].

Neocicada chisos has its northern range limit in southwestern Texas with the majority of its distribution in Mexico [19,56] (Figure 29). It is restricted to mountains in Texas extending to lower elevation Chihuahuan Desert areas of Mexico [19,56,73] and is often found in oaks (Quercus gravesii Sudworth and Q. graciliformis Muller) in riparian habitats [56] but we also found the species calling from Texas madrone (Arbutus spp.).

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Figure 29. Distribution of Neocicada hieroglyphica hieroglyphica (red circles), N. hieroglyphica johannis (blue diamonds) and N. chisos (green triangles).

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Figure 29. Distribution of Neocicada hieroglyphica hieroglyphica (red circles), N. hieroglyphica johannis (blue diamonds) and N. chisos (green triangles).
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2.9. Biogeography of the Magicicada Species

The species of the genus Magicicada are also known as periodical cicadas due to the long, predictable life cycle they exhibit. The genus was moved into the Cicadettinae and Taphurini from the Tibicinini of the Tibicininae by Moulds [138]. The seven species are restricted to North America, north of Mexico [26]. Davis [87] compared the genus to Tibicina Kolenati but a more detailed analysis placed the genus in the Taphurini [138].

The analysis of the distribution of the Magicicada species began with the formation of the year classes or broods by Marlatt [139]. The original brood maps have since been refined [140,141,142] with more detailed analyses currently being performed e.g., [143,144,145,146,147,148].

The Magicicada species maps presented here (Figure 30, Figure 31, Figure 32) were produced and kindly provided by John Cooley. The maps represent collection records from the database of specimens collected by him and or one of his colleagues that are members of the periodical cicada research group. Unverified historical records are not included as these can be incorrect as shown in the significant modifications of the original brood maps produced by Marlatt [139]. Literature records are included in the distribution listed in Table 1.

All species are primarily found in the eastern deciduous forests [94] but extend into the forested areas of eastern tall grass prairie habitats [70] (Figure 30, Figure 31, Figure 32). The database used to construct these maps is incomplete for the –cassinii and –decula species with points representing collection locations of J. Cooley or his co-workers in the Magicicada mapping project and do not include potentially incorrect historical or literature records (J. Cooley, personal communication).

There is some differentiation in the densities of the individual species across the generic range illustrated in Figure 30, Figure 31. The northernmost populations are almost entirely M. septendecim, the southwestern populations are primarily M. cassinii and M. tredecassini with the greatest density of M. septendecula and M. tredecula in the southeast (J. Cooley, personal communication).

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Figure 30. Distribution of the 17-year Magicicada species (brown circles). All species are represented collectively with each location being a single or multiple species. The distribution of individual species follows the patterns outlined in the text.

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Figure 30. Distribution of the 17-year Magicicada species (brown circles). All species are represented collectively with each location being a single or multiple species. The distribution of individual species follows the patterns outlined in the text.
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Figure 31. Distribution of the 13-year Magicicada species (gold circles). All species are represented collectively with each location being a single or multiple species. The distribution of individual species follows the patterns outlined in the text.

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Figure 31. Distribution of the 13-year Magicicada species (gold circles). All species are represented collectively with each location being a single or multiple species. The distribution of individual species follows the patterns outlined in the text.
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Figure 32. Distribution of Magicicada septendecim (brown circles), M. tredecim (gold circles) and M. neotredecim (green circles).

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Figure 32. Distribution of Magicicada septendecim (brown circles), M. tredecim (gold circles) and M. neotredecim (green circles).
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The species use different portions of their habitats [149,150] and have been shown to use a significant number of trees as host plants e.g., [151,152,153,154] primarily ovipositing in deciduous trees.

2.10. Biogeography of the Cicadetta Species

The status of the Cicadettini has long been confused (e.g., [155]) and the tribe continues to be significantly reorganized (e.g., [156]). The four species North American species have been placed in either Melampsalta and Cicadetta but are now considered to be members of Cicadetta [26]. The genus is widespread with members found in the Nearctic, Palaearctic, Oriental and Ethiopian biogeographic regions [26].

The nominotypical subspecies C. calliope calliope is the most widespread in North America (Figure 33) extending from the mid-Atlantic states to the Rocky Mountains. It is found in grasses and weeds and has been associated with meadow environments on grasses and weeds [98] and Stipa spartea Trinius-Andropogon scoparius (Michaux)Nash habitats [157]. The subspecies C. calliope floridensis was erected for Florida specimens that were completely green without black markings [21]. We have also recorded the subspecies from southern Alabama and Georgia (Figure 33). Both subspecies have been associated with the Southeastern Plains, Southern Coastal Plains and Miami Ridge/Atlantic Coastal Strip of the Southern Florida Coastal Plains ecoregions [81] in Florida [21]. The significant overlap of the subspecies suggests the validity of the subspecies needs to be investigated further.

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Figure 33. Distribution of Cicadetta calliope calliope (blue circles) and C. calliope floridensis (red diamonds).

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Figure 33. Distribution of Cicadetta calliope calliope (blue circles) and C. calliope floridensis (red diamonds).
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The remaining species of Cicadetta in North America are found in the grasslands [70] of the central United States (Figure 34). Cicadetta kansa has the most extensive distribution of these species being found from Nebraska to Texas and the distribution is coincident with the shortgrass prairie, mixed grass prairie and desert grassland ecotypes [70]. Cicadetta texana is found in the southern portion of the mixed grass prairie [70]. Cicadetta camerona has the most limited known range being found only in southern Texas (Figure 34). It currently inhabits vegetation in agriculturally disturbed regions of the vegetation of the southeastern coastal plain [72].

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Figure 34. Distribution of Cicadetta kansa (blue circles), C. texana (orange diamonds) and C. camerona (green triangles).

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Figure 34. Distribution of Cicadetta kansa (blue circles), C. texana (orange diamonds) and C. camerona (green triangles).
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2.11. Biogeography of the Platypedia Species

The species of the Platypediini were moved into the Tibicininae from their own family based on an extensive cladistic analysis [138]. The species of the Platypediini differ from the other genera of North American cicadas in that they lack timbal organs and communicate through crepitation (e.g., [158,159]).

There are 21 species and four subspecies of Platypedia in North America, north of Mexico [26]. A single species is found only in Mexico with the remaining species of the genus found in the western United States and Canada [26].

Three of the four subspecies of Platypedia are subspecies of P. putnami. The nominotypical subspecies and P. putnami lutea are found over much of the western United States (Figure 35) and have been collected in forested areas of the western portion of the central grassland [70], Rocky Mountain forests [160], Mexican Highlands of the Sonoran Desert [73], intermountain woodlands [76], California forest [71], chaparral [74,75], and Pacific coastal/Cascadian forests [161] ecosystems. Platypedia putnami putnami has been associated with skunkbrush (Rhus trilobata), wild plum (Prunus Americana Marshall), green ash (Fraxinus pennsylvanica Marshall), American elm (Ulmus Americana L.) and silver maple (Acer saccharinum L.) in shrublands [49], aspen (Populus tremaloides Michaux) [162], scrub oak (Quercus turbinella Greene) [66], mountain mahogany (Cercocarpus spp.) [158], various trees, shrubs and deciduous fruit trees in California [69] and we found it in New Mexico locust (Robinia neomexicana Gray). Platypedia putnami lutea has been associated with pointleaf manzanita (Arctostaphylos pungens Kunth) and silverleaf oak (Quercus hypoleucoides Camus) [163] as well as piñon pine (Pinus edulis) [164] and greasewood (Sarcobatus spp.) [165]. Platypedia putnami occidentalis and P. putnmai keddiensis are found in northern and central California (Figure 35). The single record for P. putnami occidentalis in southern California may be a misidentified P. putnami putnami which was collected in the same location. Platypedia putnami occidentalis and P. putnami keddiensis are associated with California forest [71], chaparral [74], and Pacific coastal/Cascadian forests [161] ecosystems.

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Figure 35. Distribution of Platypedia putnami putnami (green squares), P. putnami lutea (gold diamonds), P. putnami occidentalis (red circles) and P. putnami keddiensis (purple triangles).

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Figure 35. Distribution of Platypedia putnami putnami (green squares), P. putnami lutea (gold diamonds), P. putnami occidentalis (red circles) and P. putnami keddiensis (purple triangles).
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Platypedia mohavensis mphavensis and P. mohavensis rufescens are found over much of the western United States (Figure 36) and have been collected in Rocky Mountain forests [160], Mexican Highlands of the Sonoran Desert [73], intermountain woodlands [76], California forest [71], and chaparral [74,75] ecosystems. The nominotypical subspecies has been associated with scrub pine (Pinus spp.), juniper (Juniperus spp.) [158], Russian thistle (Echinops exaltatus Schrader) [61], serviceberry (Amelanchier spp.) and to a lesser extent rabbitbrush (Ericameria nauseosa) [49]. The subspecies P. mohavensis rufescens has been associated with scrub pine (Pinus spp.), junipers (Juniperus spp.), black locust (Robinia pseudoacacia L.) and oaks (Quercus spp.) [78]. We found it associated with one-seed juniper (Juniperus monosperma) and Utah juniper (Juniperus osteosperma). Platypedia barbata is found in primarily coastal regions of California in what are chaparral [74], and California forest [161] ecosystems.

Platypedia areolata extends over much of the northwestern United States and southern British Columbia with one specimen having been collected almost at the southern Alaska border along the coast of British Columbia (Figure 37). It is found in several ecosystems including the Pacific coastal/Cascadian forests [161], intermountain forests [76], Rocky Mountain forests [160] and California forests [71] and chaparral [74]. It has been reported with alder (Alnus spp.), sumac (Rhus spp.), balsam trees (Populus spp.) and manzanita (Arctostaphylos spp.) [158,166] and to oviposit in many deciduous fruit and olive (Olea europaea L.) trees [44,69]. Platypedia aperta is found in southern California and was recently described from Mexico [7]. It is associated with the California chaparral and woodlands [74]. Platypedia balli is found in chaparral [74,75] in Arizona (Figure 37). We found it associated with Arizona white oak (Quercus arizonica L.) and scrub oak (Quercus turbinella).

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Figure 36. Distribution of Platypedia mohavensis mohavensis (blue circles), P. mohavensis rufescens (red diamonds) and P. barbata (green triangles).

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Figure 36. Distribution of Platypedia mohavensis mohavensis (blue circles), P. mohavensis rufescens (red diamonds) and P. barbata (green triangles).
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Figure 37. Distribution of Platypedia areolata (blue circles), P. aperta (orange diamonds), and P. balli (red triangles).

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Figure 37. Distribution of Platypedia areolata (blue circles), P. aperta (orange diamonds), and P. balli (red triangles).
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Platypedia falcata has a broad yet patchy distribution from Texas to Oregon (Figure 38). The type location is in far west Texas [166] and has since been found in Arizona, California and Oregon. It is another species found in several ecosystems including forests of the Chihuahuan Desert [73], the Pacific coastal/Cascadian forests [161], intermountain forests [76], California forests [71] and chaparral [74]. Platypedia mariposa is associated with the California forests [71] ecosystem with one location in the intermountain forests [76]. Platypedia bernardinoensis has a fairly broad distribution, found in chaparral [74] and Rocky Mountain forests [160] with populations separated by low elevation desert environments (Figure 38). The species has been associated with willows (Salix spp.), birch (Betula spp.) and maple (Acer spp.) [78].

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Figure 38. Distribution of Platypedia falcata (blue diamonds), P. mariposa (red circles), and P. bernardinoensis (green squares).

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Figure 38. Distribution of Platypedia falcata (blue diamonds), P. mariposa (red circles), and P. bernardinoensis (green squares).
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Platypedia similis is found in several ecosystems (Figure 39) including California forests [71] and chaparral [74]. Platypedia affinis is found in the intermountain forests [76] of Utah (Figure 39). We found it associated with Gambel’s oak (Quercus gambelii Nuttall), manzanita (Arctostaphylos spp.) and Utah juniper (Juniperus osteosperma).

Platypedia minor is found across California with extensions into Oregon (Figure 40) and Mexico [8]. It is found in a variety of ecosystems including intermountain forests [76], Rocky Mountain forests [160], California forests [71] and chaparral [74] and has been associated with oaks (Quercus spp.), baccharis (Baccharis spp.), Pacific madrone (Arbutus menziesii Pursh), willow (Salix spp.) and fruit trees [44,69]. Platypedia rufipes is found in California forests [71], chaparral and woodlands [74]. Platypedia sylvestri is known from only two locations in the California forests [71] (Figure 40).

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Figure 39. Distribution of Platypedia similis (blue circles) and P. affinis (green squares).

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Figure 39. Distribution of Platypedia similis (blue circles) and P. affinis (green squares).
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Figure 40. Distribution of Platypedia minor (green circles), P. rufipes (red diamonds), and P. sylvestri (purple squares).

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Figure 40. Distribution of Platypedia minor (green circles), P. rufipes (red diamonds), and P. sylvestri (purple squares).
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The remaining species of Platypedia are restricted to California. Platypedia tomentosa, P. scotti, P. vanduzeei (Figure 41), P. laticapitata, and P. intermedia (Figure 42) are found in California forests [71] and chaparral [74,75] ecosystems. Platypedia tomentosa has been associated with California juniper (Juniperus californica Carrière) [93], P. scotti with thick brush [158] and P. vanduzeei with grassy hillsides and sagebrush (Artemisia spp.) [64,158,166]. We collected P. intermedia in Pacific madrone (Arbutus menziesii). Platypedia middlekauffi and P. usingeri are found in California forests [71] (Figure 42).

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Figure 41. Distribution of Platypedia tomentosa (blue triangles), P. scotti (red diamonds), and P. vanduzeei (green squares).

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Figure 41. Distribution of Platypedia tomentosa (blue triangles), P. scotti (red diamonds), and P. vanduzeei (green squares).
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Figure 42. Distribution of Platypedia laticapitata (green squares), P. intermedia (orange circles), P. middlekauffi (red triangles), and P. usingeri (blue diamonds).

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Figure 42. Distribution of Platypedia laticapitata (green squares), P. intermedia (orange circles), P. middlekauffi (red triangles), and P. usingeri (blue diamonds).
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2.12. Biogeography of the Neoplatypedia Species

The genus Neoplatypedia is related to Platypedia differing primarily in the costal margin and number of apical cells in the fore wings and is the only other member of the Platypediini [26]. Both species are restricted to North America, north of Mexico.

Noeplatypedia ampliata is found in California and Oregon (Figure 43). Neoplatypedia constricta has a more extensive distribution being found in 8 western states (Figure 43). The species of Neoplatypedia are associated with the California upland forests and woodlands habitat [71], Pacific Northwest forests [161] and N. constricta also is found in the forests of the Rocky Mountains [160] and intermountain woodlands [76]. Neoplatypedia constricta has been associated with sagebrush (Artemisia spp.) [49,165,167] and rubber rabbitbush (Ericameria nauseosa) [49].

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Figure 43. Distribution of Neoplatypedia ampliata (green diamonds) and N. constricta (orange circles).

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Figure 43. Distribution of Neoplatypedia ampliata (green diamonds) and N. constricta (orange circles).
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2.13. Biogeography of the Clidophleps Species

There are seven species and two subspecies of Clidophleps in North America, north of Mexico [26]. Clidophleps is related to Okanagana, Okanagodes, and Tibicinoides with these genera classified together in the Tibicinini. Other members of the tribe are found in Europe, Asia, India and northern Africa (the South American genus Ahomana Distant needs to be reexamined for its tribal placement).

The majority of Clidophleps species are found in California. Clidophleps distanti distanti and C. distanti truncatus are found in the California upland forests and woodlands [71] (Figure 44). The nominotypical subspecies has been reported from chaparral as well [66,74,75] and we found it associated with blue oak (Quercus douglasii Hooker & Arnott) but it has also been associated with live oak (Quercus agrifolia Née) [66]. The single record in Nevada for C. distanti distanti is suspect. The type locality of C. vagans (Figure 46) is unknown because the type specimen was collected off an automobile [87] and perhaps something similar happened with the C. distanti distanti specimen collected in Nevada as the location is between high population centers in southern California and southeastern Nevada. The overlap in the distributions of the subspecies suggests the status of the subspecies should be examined further using acoustic methods. The known range of C. distanti distanti and C. distanti truncata was recently extended into Mexico [8]. Clidophleps rotundifrons is currently known from Arizona (Figure 44), where the collection sites are found in the creosote bush-bur sage habitats [130] of the Sonoran Desert [73]. The location in Merced County, CA is questionable because even though the specimen was checked and has been properly identified, this record is in a different habitat and may have been mislabeled.

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Figure 44. Distribution of Clidophleps distanti distanti (blue circles), C. distanti truncata (red triangles) and C. rotundifrons (orange diamonds).

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Figure 44. Distribution of Clidophleps distanti distanti (blue circles), C. distanti truncata (red triangles) and C. rotundifrons (orange diamonds).
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Clidophleps wrighti, C. tenuis (Figure 45) and C. blaisdelli (Figure 46) are also found in the California upland forests and woodlands [70] with C. blaisdelli also reported from chaparral [66,168]. Clidophleps blaisdelli and C. wrighti were reported to be restricted to chamise (Adenostoma fasciculatum Hooker & Arnott) in a southern California habitat [169]. There are also some suspect locations for C. tenuis in central California and southern Nevada and C. blaisdelli in northern California and southwestern Arizona. The known range of C. wrighti was recently extended into Mexico [8]. Clidophleps beameri has a more restricted distribution (Figure 45) and has been reported from chaparral [74,75] where we found it associated with an Atriplex-like plant (probably Adenostoma spp. based on appearance and distribution of the genus in California) on which it is cryptically colored and difficult to see.

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Figure 45. Distribution of Clidophleps beameri (red triangles), C. wrighti (green squares) and C. tenuis (blue diamonds).

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Figure 45. Distribution of Clidophleps beameri (red triangles), C. wrighti (green squares) and C. tenuis (blue diamonds).
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Figure 46. Distribution of Clidophleps vagans (red squares) and C. blaisdelli (green diamonds).

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Figure 46. Distribution of Clidophleps vagans (red squares) and C. blaisdelli (green diamonds).
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The most extensive range for a Clidophleps species is that of C. vagans (Figure 46). Specimens have been collected in California, Nevada and Arizona. Collection sites are found in the California upland forest and woodlands [71], chaparral [74], and intermountain deserts and shrub steppes [76].

2.14. Biogeography of the Okanagana Species

The genus Okanagana is the most diverse member of the Tibicinini found in North America. There are 57 species and four subspecies of Okanagana in North America, north of Mexico [26]. It is related structurally to Okanagodes, Clidophleps and Tibicinoides with Okanagana probably being the phylogenetic origin for these genera. The species of Okanagana are distributed over most of North America, north of Mexico except for southeastern states.

The first group of Okanagana to be discussed is a complex of species related to Okanagana rimosa. Several species and subspecies at one time or another were classified related to O. rimosa [26]. Many specimens also have been misidentified historically (e.g., [49]). Okanagana rimosa rimosa is found in northern regions of North America extending from New Brunswick to California (Figure 47) with the first documented records for New Jersey [170] being published recently. The species is found in a variety of habitats including eastern deciduous forests [94], central plains [70], Rocky Mountain forests [160], intermountain forests [76], Pacific coastal/Cascadian forests [161], and California forests [71]. The nominotypical subspecies has been reported from aspen trees (Populus spp.) [162], white birch (Betula papyrifera Marshall), maple (Acer spp.) [170,171], pine (Pinus spp.) [66], blueberry barrens (Vaccinium spp.) [85,96], bigtooth aspen (Populus grandidentata Michaux) and blackberries (Rubus alleghaniensis Porter) [172]. The subspecies O. rimosa ohioensis is only known from northeastern Ohio (Figure 47) and may simply be a localized color variation. It has been associated with white pines (Pinus strobus L.) and hemlocks (Tsuga spp.) [93]. The validity of O. noveboracensis was recently confirmed [12] with the known distribution in western New York and southeastern Ontario (Figure 47). Okanagana occidentalis has a more western distribution (Figure 47) with the type location determined to be Chilliwack, British Columbia [173]. It can be found in large numbers in sagebrush shrublands in Colorado [49]. It has been associated with cottonwoods (Populus spp.), trees, underbrush and flowers [174]. We found it associated with several bushy plants in a wash. It is restricted to mountainous environments and is associated with the Rocky Mountain forests [160], intermountain forests [76], Pacific coastal/Cascadian forests [161], and California forests [71].

Okanagana canadensis has a similar range to O. rimosa rimosa; it is found in northern regions of North America extending from New Brunswick to California (Figure 48) in a variety of habitats including eastern deciduous forests [94], central plains [70], Rocky Mountain forests [160], intermountain forests [76], Pacific coastal/Cascadian forests [161], and California forests [71]. A specimen collected in Sambaa Deh Falls Territorial Park, Northwest Territory is the northernmost (61°N, 120°W) cicada we have recorded (Figure 48). It has been reported to use sagebrush (Artemisia spp.) [49,175], cedar trees (no species given), willows (Salix spp.) [174], cherry (Prunus spp.), Scots pine (Pinus sylvestris L.) and northern white cedar (Thuja occidentalis L.) [172] and we collected it is ponderosa pine (Pinus ponderosa). Okanagana bella has been reported to use various tree species [44,52,66] including ponderosa pine, piñon pines (Pinus edulis) [67], junipers (Juniperus spp.), aspens (Populus spp.), and willows (Salix spp.) [49] in the Rocky Mountain forests [160], intermountain forests [76], Pacific coastal/Cascadian forests [161], California forests [71] and chaparral [74]. We also found it in quaking aspen (Populus tremuloides), lodgepole pine (Pinus contorta Douglas) and mountain mahogany (Cercocarpus spp.). Okanagana annulata has a distribution similar to that of O. bella (Figure 48) and has been associated with the Rocky Mountain forests [160], intermountain forests [76], Pacific coastal/Cascadian forests [161], California forests [71] and chaparral [74,75]. We found it in oak (Quercus spp.), pine (Pinus spp.) and sequoia (Sequoiadendron giganteum (Lindley)J. Buchholz).

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Figure 47. Distribution of Okanagana rimosa rimosa (red squares), O. rimosa ohioensis (green triangles), O. noveboracensis (light blue stars) and O. occidentalis (blue circles).

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Figure 47. Distribution of Okanagana rimosa rimosa (red squares), O. rimosa ohioensis (green triangles), O. noveboracensis (light blue stars) and O. occidentalis (blue circles).
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Figure 48. Distribution of Okanagana canadensis (green squares), O. bella (red circles) and O. annulata (blue diamonds).

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Figure 48. Distribution of Okanagana canadensis (green squares), O. bella (red circles) and O. annulata (blue diamonds).
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Okanagana synodica synodica is found in grasses [36,92,129,162,174] in a wide variety of habitats but has also been reported to use sand sage (Artemisia filifolia) and low shrubs [49]. The host plants are probably grasses because we found specimens in habitats that had no vegetation other than short clumps of grass. The nominotypical subspecies is found in grasslands [70] and Rocky Mountain vegetation [160], intermountain areas [76] and California uplands [71] (Figure 49). Okanagana synodica nigra is also found in grasses but is more restricted in its distribution in southern areas (Figure 49). Okanagana balli is also a grass species that is distributed in the grasslands [70] of the northern Great Plains (Figure 49). It has been associated with weeds and grasses, lead plant (Amorpha canescens Pursh) [176] and stands of bluestem grass (Andropogon gerardii Vitman) [157]. We found specimens in remnants of prairie along railroad tracks in otherwise agricultural environments and on a remnant of tallgrass prairie in southern Minnesota. The only species of Okanagana to extend into the southern United States is O. viridis (Figure 49). We found it associated with hardwoods in the southeastern coastal plains vegetation [72] where the green coloration provided crypsis. It has also been reported to use blackjack (Quercus marilandica) and post oaks (Quercus stellata) [87].

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Figure 49. Distribution of Okanagana synodica synodica (blue squares), O. synodica nigra (red triangles), O. balli (orange circles) and O. viridis (green diamonds).

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Figure 49. Distribution of Okanagana synodica synodica (blue squares), O. synodica nigra (red triangles), O. balli (orange circles) and O. viridis (green diamonds).
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Another species of Okanagana to have subspecies is O. mariposa (Figure 50). The nominotypical subspecies has been associated with chaparral [74,75,92] and California forests [71]. It has been specifically associated with oak (Quercus spp.) [66] and we found it associated with Arizona cypress (Cupressus arizonica Greene), mountain mahogany (Cercocarpus spp.) and scrub oak (Quercus turbinella). Okanagana mariposa oregonensis is restricted to a single location in Oregon (Figure 50) near the northern border of the California forests [71] ecosystem. It is completely surrounded by the nominotypical subspecies which suggests it is highly localized color variation and further analyses to assess the status of the subspecies should be performed. A very common species of Okanagana in the west is O. utahensis (Figure 50). It is found in Great Basin sagebush (Artemisia tridentate Nuttall) [177] and sagebrush shrublands [49] in the Rocky Mountain forests [160], intermountain steppes [76], California forests [71], chaparral [74] and Palouse prairie [70].

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Figure 50. Distribution of Okanagana mariposa mariposa (red circles), O. mariposa oregonensis (green star) and O. utahensis (blue squares).

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Figure 50. Distribution of Okanagana mariposa mariposa (red circles), O. mariposa oregonensis (green star) and O. utahensis (blue squares).
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Figure 51. Distribution of Okanagana tristis tristis (green circles) and O. tristis rubrobasalis (red triangels).

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Figure 51. Distribution of Okanagana tristis tristis (green circles) and O. tristis rubrobasalis (red triangels).
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Okanagana tristis tristis (Figure 51) is found using trees in a variety of habitats including chaparral [74,75], California forests [71], intermountain forests [76] and forests of the Pacific coasts [161]. Okanagana tristis rubrobasalis is restricted to California (Figure 51) in the California forests [71] and chaparral [74]. The significant overlap of the subspecies suggests it is a color variation and further analyses to assess the status of the subspecies should be performed.

Okanagana tanneri was originally described as a subspecies of O. schaefferi [60]. Both species (Figure 52) are found in intermountain forests [76] and Rocky Mountain forests [160]. Okanagana schaefferi has been reported in sagebrush (Artemisia spp.) [174,178] and sagebrush shrublands [49]. We found O. schaefferi in Great Basin sagebrush (Artemisia tridentata) and juniper (Juniperus spp.) and O. tanneri in sage (Artemisia tridentata) and oaks (Quercus spp.). Okanagana sugdeni (Figure 52) is found in intermountain forests [76] in Utah. We found it only on to piñon pine (Pinus edulis).

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Figure 52. Distribution of Okanagana schaefferi (blue squares), O. tanneri (green circles) and O. sugdeni (red diamonds).

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Figure 52. Distribution of Okanagana schaefferi (blue squares), O. tanneri (green circles) and O. sugdeni (red diamonds).
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Another common species of Okanagana in the west is O. striatipes (Figure 53). It is found in Great Basin sagebush (Artemisia tridentata) [78,177] and sagebrush shrublands [49] in the Rocky Mountain forests [160], intermountain steppes [76], California forests [71], chaparral [74], grasslands and Palouse prairies [70]. Okanagana oregona is found in trees in the California forests [71], chaparral [74,75], intermountain [76] and rocky Mountain forests [160] (Figure 53). We found it associated with white fir (Abies concolor (Gordon)Lindley ex Hildebrand) and also calling from grasses. Okanagana nigrodorsata is restricted to California upland forests [71] (Figure 53).

Okanagana hesperia (Figure 54) is a common grass species that has been associated with chaparral [74,75] and the Great Plains [49,70] but is also found in intermountain areas [76]. We found it associated with grasses and weedy plants such as rabbit bush (Chlysothamnus spp.) but it would also call from trees like ponderosa pine (Pinus ponderosa) and one-seed juniper (Juniperus monosperma). It has previously (as O. striatipes beameri Davis) been reported to oviposit in weeds [66]. Okanagana yakimaensis is restricted to the Palouse prairie [70] of central Washington (Figure 54). Okanagana ornata is primarily found in California forests [71] with a few records in chaparral [74] and forests of the Rocky Mountains [160]. It has been associated with oak bushes (Quercus spp.) [59].

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Figure 53. Distribution of Okanagana oregona (green circles), O. nigrodorsata (blue triangles) and O. striatipes (orange squares).

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Figure 53. Distribution of Okanagana oregona (green circles), O. nigrodorsata (blue triangles) and O. striatipes (orange squares).
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Figure 54. Distribution of Okanagana yakimaensis (red star), O. ornata (blue diamonds) and O. hesperia (orange circles).

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Figure 54. Distribution of Okanagana yakimaensis (red star), O. ornata (blue diamonds) and O. hesperia (orange circles).
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One of the most common species of Okanagana in collections is O. vanduzeei (Figure 55). It has been reported to inhabit chaparral [64,74,75], California forests [71], intermountain areas [76], Rocky Mountain vegetation [160] and the Palouse prairie [70]. As previously reported [64,66], we collected it from a variety grasses and sage (Artemisia spp.) over a wide range of California and it has been associated with dry grasses in fields and hillsides of California [69]. It has also been known to use wild buckwheat (Eriogonum spp.), sage, Baccharis spp., goldenrod (Solidago spp.) and pines (Pinus spp.) [44,64] but is rarely found in trees [174]. The known range of O. vanduzeei was recently extended into Mexico [8]. Okanagana fratercula is found in intermountain regions [76], forests of the Rocky Mountains [160] and grasslands [70]. We found it in small bushes and it has been associated with sagebrush (Artemisia spp.) [178]. Perhaps the most restricted distribution of any species in North America, north of Mexico is that of O. georgi (Figure 55) which is currently known from a restricted section of chaparral [18,74,75] in northern Arizona. We found it associated with Arizona cypress (Cupressus arizonica), and Lowell ash (Fraxinus anomala var. lowellii (Sargent) Little).

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Figure 55. Distribution of Okanagana vanduzeei (blue squares), O. fratercula (red diamonds) and O. georgi (green circle).

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Figure 55. Distribution of Okanagana vanduzeei (blue squares), O. fratercula (red diamonds) and O. georgi (green circle).
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Okanagana luteobasalis (Figure 56) has a wide distribution across much of the western states and southern Canada east of the Sierra Nevada Mountains. It is associated with several ecosystems including the intermountain vegetation [76], Rocky Mountain vegetation [160] and grassland environments [70]. We collected the species on sagebrush (Artemisia spp.) as has been reported previously [61]. Okanagana rubrovenosa is found in chaparral [74,75] and California forests [71] (Figure 56) on manzanita (Arctostaphylos spp.) [38,44,66,78,174] and was reported to be restricted to Arctostaphylos pungens in a southern California habitat [168] where their red coloration provides crypsis. Okanagana vandykei is associated with the California forests [71] (Figure 56) and has been reported to be arboreal [44].

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Figure 56. Distribution of Okanagana luteobasalis (gold circles), O. rubrovenosa (red squares) and O. vandykei (blue triangles).

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Figure 56. Distribution of Okanagana luteobasalis (gold circles), O. rubrovenosa (red squares) and O. vandykei (blue triangles).
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Okangana gibbera (Figure 57) is found in intermountain areas [76], Rocky Mountain vegetation [160] and grassland environments [70]. Okanagana fumipennis has been associated with sagebrush-greasewood shrublands and juniper-sagebrush woodlands [49] and emergence holes have been found under big sagebrush (Artemisia tridentata) [78] and greasewood (Sarcobatus vermiculatus (Hooker)Torrey) [49]. We also collected the species on yellow menodora (Menodora scabra Gray). It is found primarily in the southern portions of the intermountain shrub steppes [76]. Okanagana cruentifera is found in the California forests [71] with some extensions into the intermountain forests [76] (Figure 57).

Okanagana ferrugomaculata is found in several forested areas of the northwest (Figure 58) and has been collected in chaparral [74,75], California forests [71], intermountain forests [76] and Pacific coastal forests [161]. We found it associated with lodgepole pine (Pinus contorta) and oaks (Quercus spp.). Okanagana pallidula is found in chaparral [74,75,179] primarily along the central valley of California (Figure 58) and has been associated with bushy weeds and grasses [66] and we also found it in grasses. Okanagana lurida is found in the Palouse prairies [70] and intermountain forests [76] and we found it associated with grasses and weeds. Okanagana formosa (Figure 58) is more restricted in its distribution and is found in trees in the intermountain forest ecosystem [76]. We found it associated with juniper (Juniperus spp.).

Okanagana magnifica is a large species that extends across much of the southwest (Figure 59). It has been reported in chaparral [75], pinyon-juniper woodlands [66,67,174], juniper (Juniperus spp.) [66,174], piñon pine (Pinus edulis) [67,164], sagebrush (Artemisia spp.), Gambel oak (Quercus gambelii, and cottonwood (Populus spp.) [36,174]. We found it in piñon pine and single leaf piñon pine (Pinus monophylla Torrey & Frémont). It is inhabits the forests of the Rocky Mountains [160], chaparral [74,75], intermountain forests [76] and California forests [71]. Okanagana simulata (Figure 59) is also found in the California forests [71]. Okanagana napa has a scattered distribution and is found in California forests [71], chaparral [74] and intermountain forests [76] and has been associated with oak (Quercus spp.), other trees and shrubs in the hillsides coastal California mountain range [69].

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Figure 57. Distribution of Okanagana gibbera (blue circles), O. cruentifera (orange triangles) and O. fumipennis (green diamonds).

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Figure 57. Distribution of Okanagana gibbera (blue circles), O. cruentifera (orange triangles) and O. fumipennis (green diamonds).
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Figure 58. Distribution of Okanagana formosa (blue squares), O. pallidula (orange triangles), O. ferrugomaculata (red circles) and O. lurida (green diamonds).

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Figure 58. Distribution of Okanagana formosa (blue squares), O. pallidula (orange triangles), O. ferrugomaculata (red circles) and O. lurida (green diamonds).
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Figure 59. Distribution of Okanagana magnifica (orange circles), O. simulata (blue diamonds) and O. napa (green triangles).

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Figure 59. Distribution of Okanagana magnifica (orange circles), O. simulata (blue diamonds) and O. napa (green triangles).
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Okanagana triangulata is found in chaparral [74,75] and was recently reported from Mexico [8]. It has been associated with meadows and grasses [44,174] and we found it associated with large species of grasses. Okanagana canescens is found in chaparral [74,75] and California forests [71] (Figure 60) where we collected it on blue oak (Quercus douglasii). Okanagana aurora is found in California forests [71] and western regions of the intermountain forests [76]. Okanagana sperata (Figure 60) has a scattered distribution within the known range of the California upland forests [71].

Bliven [180] described several species of cicadas from northern California (Figure 61). Okanagana orithya, O. vocalis, O. salicicola, O. sequoia and O. rhadine are found at the southern end of the Pacific coastal forests [161]. Okanagana pernix is found in the northern end of the California upland forests [71]. No further information on the biology of these species is available.

Another series of California cicadas related to chaparral [74] is O. arctostaphylae, O. venusta, O. opacipennis and O. uncinata [75] (Figure 62). Okanagana arctostaphylae was named for its affinity for manzanita (Arctostaphylos spp.) [181] and was reported to be restricted to Arctostaphylos pungens in a southern California habitat [168] in which the red coloration provides crypsis. We found only two locations for the species, one of which is Cajon Pass, a common collecting site in California. Okanagana opacipennis has also been associated with manzanita (Arctostaphylos spp.) [40,44,166,175] where its coloration would also provide crypsis. Okanagana villosa is restricted to high elevations in the California upland forests [71] (Figure 62).

The remaining species of Okanagana are also restricted to California (Figure 63). Okanagana wymorei is found in chaparral [74,75] and California forests [71] as is O. arboraria. We found O. wymorei in grasses. Okanagana nigriviridis is found in elevated chaparral [74] associated with shrubs [66] and was reported to be restricted to chamise (Adenostoma fasciculatum) in a southern California habitat [169]. Finally, Okanagana hirsuta is found in coastal chaparral and offshore islands and O. catalina is geographically isolated in the chaparral [74] of the Catalina Islands.

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Figure 60. Distribution of Okanagana triangulata (green diamonds), O. sperata (orange circles), O. canescens (red squares) and O. aurora (blue diamonds).

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Figure 60. Distribution of Okanagana triangulata (green diamonds), O. sperata (orange circles), O. canescens (red squares) and O. aurora (blue diamonds).
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Figure 61. Distribution of Okanagana orythia (dark blue squares), O. vocalis (light blue circles), O. rhadine (orange triangle), O. salicola (red diamond), O. pernix (purple crosses) and O. sequoia (green stars).

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Figure 61. Distribution of Okanagana orythia (dark blue squares), O. vocalis (light blue circles), O. rhadine (orange triangle), O. salicola (red diamond), O. pernix (purple crosses) and O. sequoia (green stars).
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Figure 62. Distribution of Okanagana arctostaphylae (light blue stars), O. opacipennis (orange diamonds), O. uncinata (red circles), O. venusta (dark blue squares) and O. villosa (green triangles).

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Figure 62. Distribution of Okanagana arctostaphylae (light blue stars), O. opacipennis (orange diamonds), O. uncinata (red circles), O. venusta (dark blue squares) and O. villosa (green triangles).
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Figure 63. Distribution of Okanagana arboraria (blue squares), O. hirsuta (orange triangles), O. wymorei (red circles), O. catalina (purple stars) and O. nigriviridis (green diamonds).

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Figure 63. Distribution of Okanagana arboraria (blue squares), O. hirsuta (orange triangles), O. wymorei (red circles), O. catalina (purple stars) and O. nigriviridis (green diamonds).
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2.15. Biogeography of the Okanagodes Species

The genus Okanagodes is named for its relationship to Okanagana [92] but is most similar to Tibicinoides or Clidophleps in certain structural characteristics [26]. Although primarily distributed in North America, north of Mexico, one species of the genus have also been reported from Mexico [6,8].

Two species of Okanagodes are known, one of which has two subspecies. All species are associated with Atriplex spp. [52] on which they are cryptically colored and difficult to see. Okanagodes gracilis gracilis and O. gracilis viridis are found in salt flats of the Sonoran, Mohave, Colorado, and Great Basin deserts [52,73,88] (Figure 64). The nominotypical subspecies extends over four states and Mexico with O. gracilis viridis being found primarily in the southeastern portion of the species distribution with a few specimens recorded from California. The subspecies may in fact be a local variation in color. Further analysis of songs and morphology should be performed to determine the status of the subspecies.

Okanagodes terlingua is currently only known from the Big Bend region of Texas [52,56] (Figure 64) but should be found in Mexico with continued collecting. It is also restricted to the salt flats but is found in the Chihuahuan Desert [52,56,73].

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Figure 64. Distribution of Okanagodes gracilis gracilis (blue circles), O. gracilis viridis (green diamonds) and O. terlingua (orange triangles).

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Figure 64. Distribution of Okanagodes gracilis gracilis (blue circles), O. gracilis viridis (green diamonds) and O. terlingua (orange triangles).
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2.16. Biogeography of the Tibicinoides Species

The genus Tibicinoides is related to Okanagana and Okanagodes [26] and all three species are restricted to California. We collected T. mercedita in grasses and expect the other species to be associated with grass as well.

Tibicinoides cupreosparsa has the southernmost distribution extending into northern Mexico (Figure 65). Tibcicinoides mercedita extends across the central two thirds of western California (Figure 65) in chaparral [74,75]. Tibicinoides minuta is generally more concentrated in central California (Figure 65). All three species are associated with the California upland forests and woodlands habitat [71] and their distributions coincide with this floristic region. All species have been associated with grasses [44,70,175] with T. minuta associated with tarweed (probably Holocarpha virgata (A. Gray)D.D. Keck) [167] and T. cupreosparsa with tufted grasses (probably Poa spp.) [64].

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Figure 65. Distribution of Tibicinoides minuta (green squares), T. mercedita (red diamonds) and T. cupreosparsus (blue triangles).

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Figure 65. Distribution of Tibicinoides minuta (green squares), T. mercedita (red diamonds) and T. cupreosparsus (blue triangles).
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The North American cicada fauna show distributional patterns that are related to their phylogeny. The representatives of Cacama, Diceroprocta, Cornuplura, Pacarina, Quesada and Neocicada in North America, north of Mexico are the northernmost of their respective genera with greater diversity being found in Central and/or South America. Although the species of the genus Beameria are primarily found in North America, north of Mexico, they represent the northernmost extension of the tribe Fidicinini. The species of Magicidada are restricted to North America, north of Mexico and also represent the northernmost extension of the tribe Taphurini. The Tibicininae has its greatest diversity in the New World with a large number of species found in southern South America (the Tettigadini) along with the North American Platypediini and the Tibicinini with representatives from North America, Europe, North Africa and Asia. The genera of the Platypediini and the North American Tibicinini are primarily distributed in North America, north of Mexico with the distribution of only a few species extending southward into Mexico. Both the genera Cicadetta and Tibicen have expansive distributions in the Palearctic and Oriental regions with Tibicen extending southward into the Neotropical region. The relatively low diversity of Cicadetta species and their distribution on the eastern portion of North America suggests immigration from the Palearctic region where there is greater diversity. The distribution of the three species groups within Tibicen illustrate a geographical separation between the eastern and western species groups. The large Tibicen species are sympatric with many of the eastern and western species. Further analyses should be performed to determine if these species groups represent distinct genera.

California (89 species, 46.6% of taxa) is the state with the greatest alpha diversity. Texas (57 species, 29.8% of taxa), Arizona (49 species, 25.7% of taxa), Colorado (36 species, 18.8% of taxa), and Utah (35 species, 18.3% of taxa) complete the top five states in biodiversity. Maine (4 species, 2.1% of taxa), New Hampshire (4 species, 2.1% of taxa), and Rhode Island (5 species, 2.6% of taxa) exhibit the lowest alpha diversity containing only species that extend over much of the eastern or northern part of North America. Although there is a size effect to biodiversity [182], there is a good relationship between the diversity of habitats within these states and the observed diversity. Mountain terrain within these states also may increase the diversity as has been suggested for other regions of the world [8,13,183]. The variety of plant communities provides the opportunity to support a greater variety of cicadas if the insects specialize on particular types of plants as hosts.

Texas, Arizona, Colorado and Utah also illustrate how phylogeny can influence regional diversity. These states have representatives of genera that that are unique to the Nearctic but also representatives of genera that have examples in the Neotropical and Palaearctic regions. California is an area noted for the high concentration of species from the genera Platypedia, Okanagana, and Clidophleps and is the only state in which the species of Tibicinoides is found. Only four California species are from genera that are found primarily in Mexico [26] and appear to have colonized southern California from the east or southeast.

California (35 species, 18.3% of taxa) is also the state with the greatest number of species found only in a single state. It is followed by Texas (15 species, 7.9% of taxa), Arizona (10 species, 5.2% of taxa), Utah (3 species, 1.6% of taxa) and Florida, Ohio, Oregon and Washington (1 species each, 0.5% of taxa each) for states with unique diversity. The same factors influencing alpha diversity are probably responsible for the single states in which particular species are found. Limited collecting and access to Canadian collections makes analysis of Canadian provinces difficult, so the summary comments are restricted to the lower 48 United States.

The known distribution of several species is limited due to the accessibility to collection sites. For example, 19 species from four genera have been recorded as collected at Cajon Pass in San Bernardino County, California and 14 species from four genera were collected at Buckman Springs, San Diego County, California. Both of these sites are found where major roadways cross mountain ranges and represent the only location in the geographic area where collecting is possible without hiking into areas without roadways. We can expect further range extensions with continued collecting.

Several regions need additional collection efforts. There are gaps in the distribution of species through central Nevada and within certain portions of the Rocky Mountains. The distance between the most northern examples of some species in Canada (e.g., Platypedia areolata (Figure 37)) and Okanagana luteobasalis (Figure 56) and the remaining distribution of the species suggests there are large areas of Canada that also need to be surveyed in more detail.

A review of the distributions of subspecies suggests that the status of the subspecies be re-evaluated. The current subspecies of cicadas were produced with the publication of the last edition of the Code [58] when varieties were converted to subspecies following Article 45.6.4. Although some subspecies appear to have geographic separation (e.g., the subspecies of Tibicen lyricen (Figure 11) and Tibicen pronotalis (Figure 21)), others are restricted to individual locations (e.g., Okanagana mariposa oregonensis (Figure 50)) or are surrounded by the nominotypical subspecies (e.g., Okanagana tristis rubrobasalis (Figure 51) or Diceroprocta swalei castanea (Figure 4)). We have shown [27,29,30] that several taxa originally described as varieties are actually species. The distributions of the remaining subspecies illustrated here will hopefully stimulate further study to determine the status.

3. Experimental Section

Location data were collected from the cicada specimens housed in more than 110 institutional and private collections (see list in acknowledgements). These data were combined with data from our own field studies between 1985 and 2004 and specimens sent to AFS for determination. Label data were placed in spreadsheets and missing information, i.e., counties or coordinates, was collected from maps and websites, particularly the USGS Geographic Names Information System [184] and Heavens-Above GmbH [185]. The distribution maps were produced with iMAP version 3.5 (Biovolution, Leuven, Belgium). A limited number of specimens from Mexico are plotted on some on the figures even though the scope of the paper is north of Mexico to give a more complete illustration of the distribution of the species since the maps used often included the northern Mexican states. Specimens collected during agricultural inspections (e.g., a specimen of Neocicada hieroglyphica collected at an agricultural inspection site in California) were not included in the distribution map because the source of the specimen could not be verified and these specimens were transported to the state in question with agricultural products.

Species are grouped on maps to illustrate one of several potential concepts without making the map unclear or confusing with respect to the distributions of the species illustrated. Subspecies are placed on the same map in order to illustrate the comparative distribution of each subspecies. Species that form a complex of related taxa are also mapped on a single map when possible. The remaining species are grouped together based on the ability to demonstrate the distributions of the collected species without interference with one another.

4. Conclusions

We have shown that cicadas show an affinity for specific plant communities and are limited in their distribution by the plants within these communities. Most species have extensive ranges but the known range of some species is limited (e.g., Okanagana georgi and O. rhadine) either due to habitat isolation (e.g., O. georgi) or to limited collecting effort (e.g., O. rhadine). Some species with limited distributions north of Mexico (e.g., Cornuplura nigroalbata or Diceroprocta lata) have more extensive ranges into Mexico. The first look provided here for the distributions of subspecies suggests that several subspecies may simply be local color variations. Further collecting efforts in areas that have tradiationally had limited access should fill in the distribution pattern for some western species. We hopefully have provided a useful tool for future researchers in locating the cicadas of North America, north of Mexico.

Acknowledgments

This work would not have been possible without the assistance of following individuals who provided access to collections or specimens used to construct the maps: S. Alm (University of Rhode Island), J. Amrine (University of West Virginia), D. Arnold (K.C. Emerson Entomology Museum, Oklahoma State University), C. Barr (Essig Museum of Entomology, University of California Berkeley), C. Bartlett (University of Delaware), 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), H. Brailovsky (Universidad Nacional Autónoma de México), J. Brambila, D. Ziesk, and S. Halbert (Florida State Collection of Arthropods), R. Brooks, G. Byers and K. Segelquist (Snow Entomology Museum, University of Kansas), R. Brown (Mississippi Entomological Museum, Mississippi State University), F. Carle and M. May (Rutgers University), C. Carlton and V. Bayless (Louisiana Arthropod Museum), E. Cashatt (Illinois State Museum), M. Caterino (Santa Barbara Museum of Natural History), P. Cato and P. Beller (San Diego Museum of Natural History), D. Chandler (University of New Hampshire), B. Clark and J. Bellon (Orma J. Smith Museum of Natural History, Albertson College of Idaho), S. Clark (West Virginia Department of Agriculture), W. Clark (Auburn University), P. Clausen (University of Minnesota Insect Collection), N. Cobb (Colorado Plateau Museum of Arthropod Diversity, Northern Arizona University), C. Cockendolpher (Museum of Texas Tech University), G. Courtney (Iowa State University), C. Covell (University of Louisville), Paula Cushing (Denver Natural History Museum), G. Dahlem (Cincinnati Museum of Natural History), S. Dash (University of Texas at El Paso), R. Davidson (Carnegie Museum of Natural History), L. Deitz and R. Blinn (North Carolina State University Insect Collection), J. Deckert (Museum für Naturkunde, Berlin), C. Donohue (Maine Forest Service), M. Douglas (Arizona State University), C. Durden (Texas Memorial Museum), M. Epstein, R. Froeschner and S. McKamey (Smithsonian Institution, National Museum of Natural History), S. Fullerton (Arthropod Collection, University of Central Florida), R. Gill (California State Collection of Arthropods), M. Goodrich (Spooner-Riegel-Goodrich Insect Collection, Eastern Illinois University), N. Greig (Houston Museum of Natural Sciences), R. Gundersen (St. Cloud State University), B. Gustafsson and G. Lindberg (Swedish Museum of Natural History), R. Gwiazdowski (University of Massachusetts), W. Hanson (Utah State University), M.S. and J.E. Heath, S. Heydon (Bohart Museum of Entomology, University of California Davis), K. Hobson (Sam Noble Oklahoma Museum of Natural History, University of Oklahoma), R. Hoffman (Virginia Museum of Natural History), C. Holliday (Lafayette College), J. Honda (J. Gordon Edwards Entomology Museum, San Jose State University), M. Horton and J. Morse (Clemson University), R. Hurley and M. Ivie (Montana Entomology Collection), E. Johnson (Staten Island Institute of Arts and Sciences), P. Johnson (Severin-McDaniel Insect Research Collection, South Dakota State University), J. Keiper (Cleveland Museum of Natural History), K.C. Kim (Frost Entomological Museum, Penn State University), J. Knight (Nevada Department of Agriculture Entomology Laboratory), B. Kondratieff (C.P. Gillette Museum of Biodiversity, Colorado State University), S. Krauth (University of Wisconsin), N. Kristensen (Natural History Museum of Denmark), J. Labonte (Oregon Department of Agriculture), P. Lago (University of Mississippi), J. Liebherr and R. Hoebeke (Cornell University Insect Collection), C. Marshall (Oregon State Arthropod Collection), T. MacRae, T. McCabe (New York State Museum), D. Marshall and K.B.R. Hill, K. McCravy (Western Illinois University), J. McPherson (Southern Illinois University), H. Mejlon (Uppsala University Zoological Museum), F. Merickel (W.F. Barr Entomological Museum, University of Idaho), C. Mitter (University of Maryland), R. Norton (State University of New York College of Environmental Science & Forestry, Syracuse), L. and C. O’Brien (Florida A & M University Center for Biodiversity), M. O’Brien (University of Michigan Museum of Zoology), J. O’Donnell (University of Connecticut), C. Olson (University of Arizona), P. Parillo (Field Museum of Natural History), G. Parsons (A.J. Cook Arthropod Research Collection, Michigan State University), N. Penney (California Academy of Sciences), J.-M. Perron (Université Laval), M. Potts (University of Kentucky), P. Perkins (Museum of Comparative Zoology, Harvard University), G. Perrigo, (Gillaspy Insect & Arthropod Museum, Texas A & M Univeristy-Kingsville), M. Potzler (Buffalo Museum of Science), A. Provonsha (Purdue University), R. Pupedis (Peabody Museum of Natural History, Yale University), B. Ratcliff (University of Nebraska), D. Richman (New Mexico State University Arthropod Museum), D. Rider and J. Fauske (North Dakota State University), E. Riley (Texas A & M University Entomology Collection), I. Robertson (Boise State University), M. Rosenzweig (Virginia Polytechnic Institute State University), C. Roy (Zaddock Thompson Natural History Museum, University of Vermont), E. Scherbrooke (Southwestern Research Station, American Museum of Natural History), R. Schuh and S. Oygur (American Museum of Natural History), O. Schwarz and C. Boake (University of Tennessee-Knoxville), A. Sharkov and C. Freeman (Museum of Biodiversity, Ohio State University), S. Shaw (University of Wyoming), R. Sites (Wilbur R. Enns Entomology Museum, University of Missouri), C. Smith (Georgia Museum of Natural History), J. Stimmel (Pennsylvania Department of Agriculture), Eric Toolson (Museum of Southwest Biology, University of New Mexico), Milton Ward (University of Alabama), M. Webb (Natural History Museum, London), J. Weintraub and D. Azuma (Philadelphia Academy of Natural Sciences), K. Welch (Connecticut Agricultural Experiment Station), D. Whitman (Illinois State University), W. Xie (Natural History Museum of Los Angeles County), D. Yanga (Entomology Research Museum, University of California Riverside), D. Yanga and K. Zeiders (Illinois Natural History Survey), W.C. Yearian (University of Arkansas Arthropod Museum), A. Young (Milwaukee Public Museum), R. Zack (M.T. James Entomological Collection, Washington State University), G. Zolnerowich (Kansas State Museum of Entomological and Prairie Arthropod Research), and J. Zyla. Special thanks are offered to J. Cooley (University of Connecticut) for producing and providing the Magicicada maps. Periodical cicada maps were generated using data obtained by John Cooley, Kathy Hill, David Marshall, Chris Simon, and their coauthors. The periodical cicada mapping project was supported by the Frank Ammermann Endowment of the University of Michigan Museum of Zoology Insect Division, the National Geographic Society, and various National Science Foundations grants to Chris Simon. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the authors and do not necessarily reflect the views of the funding agencies. AFS wishes to acknowledge the financial support of Sr. John Karen Frei of Barry University, a sabbatical leave at Barry University, Barry University Faculty Senate Mini-Grant, and a Collection Study Grant from the American Museum of Natural History. AFS was provided with financial support from Sr. John Karen Frei of Barry University, a sabbatical leave at Barry University, Barry University Faculty Senate Mini-Grants, and a Collection Study Grant from the American Museum of Natural History. Two anonymous reviewers made suggestions that improved the manuscript.

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