Research Article Journal of Orthoptera Research 2021, 30(2): 131-143 A new species of pygmy mole cricket (Orthoptera: Tridactylidae) from the Lake Wales ridge of Florida and new records of Ellipes eisneri from the northern Brooksville ridge BRANDON Woo! 1 Cornell University, Comstock Hall, Department of Entomology, Ithaca, NY 14853, USA. Corresponding author: Brandon Woo (bmw95 @cornell.edu) Academic editor: Hugh Rowell | Received 18 March 2021 | Accepted 30 April 2021 | Published 27 September 2021 http://zoobank.org/91AGE44D-F43E-45A1-9E57-D2854959D32B Citation: Woo B (2021) A new species of pygmy mole cricket (Orthoptera: Tridactylidae) from the Lake Wales Ridge of Florida and new records of Ellipes eisneri from the northern Brooksville Ridge. Journal of Orthoptera Research 30(2): 131-143. https://doi.org/10.3897/jor.30.65603 Abstract Pygmy mole crickets (Orthoptera: Tridactylidae) are usually associated with the edges of ponds and streams, but in the sandy uplands of Florida, at least two lineages of these insects have evolved to live in xeric scrub and sandhill habitats. Very little work has been done with scrub tridactylids since they are tiny and often difficult to collect. In this paper, the pygmy mole cricket Ellipes deyrupi sp. nov. is described from the northern Lake Wales Ridge of Florida. It is sympatric at all known locations with another scrub tridactylid, Neotridactylus archboldi Deyrup & Eisner, but has not been found co-occurring with the closely related Ellipes eisneri Deyrup. The hab- its of this new species are convergent with both N. archboldi and E. eisneri. In addition, new records of Ellipes eisneri are presented, extending this spe- cies’ known distribution to the Northern Brooksville Ridge. Both species of scrub Ellipes are found in restricted geographic ranges and suffer from a lack of study and recognition. Conservation implications for these two species are discussed. Keywords Distribution, Florida endemics, sandhill, scrub, taxonomy, Tridactylinae Introduction The pygmy mole crickets, or Tridactylidae (Orthoptera: Caelifera: Tridactyloidea), are a family of small, burrowing or- thopterans with a worldwide distribution (Deyrup and Eisner 1996). There are currently seven species known in the Nearctic re- gion, with five recorded in Florida (Giinther 1975, 1977). Three of these, Neotridactylus apicialis (Say, 1825), Ellipes minuta (Scudder, 1862), and E. gurneyi Giinther, 1977, are fully winged, widespread in the southern USA, and occur in wet, sandy areas along water, as is typical for most members of the family. The other two species, N. archboldi Deyrup & Eisner, 1996, and E. eisneri Deyrup, 2005, are flightless and endemic to ridge systems of central Florida, where they occur in open patches of bare sand within scrub (N. archboldi) or sandhill (E. eisneri) habitats. They are able to occur in these xeric environments because of the presence of a subsurface layer of algae in the bare patches of sand they inhabit. This pro- vides them with a source of food that is transiently available when rains occur, at which time the insects come to the surface and con- struct feeding burrows (Deyrup and Eisner 1996). Their habitat and food restrictions, as well as their flightless nature, mean that populations of scrub tridactylids are often isolated from each oth- er, both within and between different ridge systems. The Florida scrub is a unique and threatened habitat restricted to a series of ridges and dunes formed from Miocene, Pliocene, or Pleistocene shorelines, during which time the rest of the state was underwater (Deyrup 1989). These ridges served as biotic refuges during glaciation events, and today serve as ecological islands, where isolated populations of many species have diverged into distinct species. Despite being located in a subtropical region with high precipitation, the scrub is a rather dry environment owing to the excessive draining of water through its fine-grained sands (Menges 1999). Because of these conditions, it supports a distinct community of organisms specially adapted to drought. Florida scrub is characterized by low evergreen oaks (Quercus spp.) and sand pine (Pinus clausa (Chapm. ex Engelm.)) Vasey ex Sarg.), often with the ericaceous shrub Ceratiola ericoides Michx., inter- spersed with patches of bare white sand where many endemic herbaceous plants and lichens occur (Menges 1999). Sandhill, or ‘high pine’ is another xeric habitat that is found on the Florida sand ridges but which also extends into other southeastern states (Myers 1985). This environment has a canopy of longleaf pine (Pinus palustris Mill.) and turkey oak (Quercus laevis Walter) and is more densely herbaceous than scrub, with abundant Aristida spp. grasses dominating (Menges 1999). Both scrub and sandhill are fire-maintained ecosystems, with sandhill burning more frequent- ly than scrub (Myers 1985). The oldest and longest of Florida’s sand ridges, and the one with the highest levels of endemism, is the Lake Wales Ridge (LWR), which extends in central Florida from just south of the Orlando area to just north of Lake Okeechobee (Deyrup 1990). The LWR contains a distinctive type of habitat, called yellow sand scrub, which is an intermediary between classic scrub and sandhill (Deyrup and Carrel 2012). This habitat occurs on higher elevat- ed sites with yellow sand and large populations of scrub hickory JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) 132 (Carya floridana Sarg.) and can vary from dense to more open scrub. There appears to be several arthropod species restricted to yellow sand scrub, including the Highlands tiger beetle (Cicindeli- dia highlandensis Choate, 1984), a candidate for federal protection (Deyrup and Carrel 2012). The LWR is one of the best studied ridg- es since it is the home of the Archbold Biological Station (ABS), which has conducted research on LWR endemic organisms since 1941. A total of 91 species of arthropods are known to be restricted to scrub habitat on the LWR (Deyrup and Carrel 2012), including six described species of Orthopterans. Most Florida scrub endemic arthropods are poorly known due to their small sizes, restricted geographic ranges, and habitat spe- cializations (Deyrup 1990), which are often such that only tar- geted collecting will reveal individuals. The scrub tridactylids are particularly difficult to locate, since they are only reliably found at or near the surface of the sand within a few hours after a rain. Their presence can sometimes be predicted in periods of drier weather, however, by the presence of small, excavated mounds of sand (tumuli) created when the insects burrow vertically down- ward in patches of open sand. Even when populations are located, individuals are a challenge to capture due to their impressive leap- ing abilities. Deyrup and Carrel (2012) completed an extensive sampling effort of LWR endemic arthropods across 23 different protected scrub preserves. In the course of that work, specimens of an unusual tridactylid were collected on the east side of the north-central LWR that clearly represented an undescribed species. The species was recognized as new by Deyrup (pers. com.) but has remained undescribed until now. Methods Collection.—Initial specimens of the new species were collected in yellow bowl traps (always following rain) during a 2009 survey of arthropods endemic to the Lake Wales Ridge. Additional material was collected by the author in August 2018 at the Tiger Creek Pre- serve and in August 2019 at two roadside sites east of Lake Marion. These two localities, as well as roadside sites on the Brooksville Ridge that might harbor Ellipes eisneri populations, were located us- ing Google Maps. The author made several trips to the Brooksville Ridge during June and July 2018 in search of E. eisneri, but dry con- ditions kept the insects deep underground and out of reach. On July 28 and 29, 2018, the weather was more favorable, and specimens of E. eisneri were collected at five sites. All tridactylid specimens were collected either by carefully digging up the raised burrows immedi- ately after rain or by sifting dry sand around the tumuli. Determination.—Specimens of E. eisneri were identified using Dey- rup (2005). No other known tridactylid species has the abbrevi- ated wings and pale color pattern of E. eisneri, except for the new species described below. The new species is a member of the genus Ellipes Scudder, 1902, due to the extreme reduction of the hind tarsus. Its pale white and brown coloration distinguishes it from all other species of the genus, except for E. albicollatus Giinther, 1977 and E. eisneri. E. albicollatus is described as having the caudal margin of the pronotum all white, with a white spot in the middle of the anal field of the tegmen. This species differs in other aspects of its coloration as well as in the shape of the female subgenital plate and is also known only from Brazil. Range, as well as the characters given below, will separate E. eisneri and the new species. Measurements and terminology.—Measurements were made to the nearest 0.5 mm by hand using a ruler and dissecting microscope B. WOO (Wild Heerbrugg, Heerbrugg, Switzerland). Terminology follows Deyrup (2005). Photographs and maps.—Live nymphs and adults of both species were photographed in situ using a Canon EOS Rebel T3 with an attached Canon 100 mm macro lens and an external flash (Sun- pak Auto 383 Super connected with a CowboyStudio 4 Channel Wireless Hot Shoe Flash Trigger & Receiver). Collected nymphs and adults were photographed in a whitebox studio setup us- ing the same equipment, with the addition of Canon extension tubes (EF25 and EF12 II) to the camera. Images of the habitat and tumuli were taken with an iPhone. Images of preserved specimens were made using a Canon EOS 6D with attached MPE 65 mm macro lens (20X zoom lens used for the terminalia and stridulatory apparatus) and Macro Twin Lite MT-24EX flash on a Cognisys Stackshot 3X system. Images were stacked using Zerene Stacker v.1.04 (Zerene Systems LLC, Richland, WA). All images were processed using Adobe Lightroom CC 2015.8 to crop and adjust white balance. Composite figures were created using Adobe Photoshop CC, v.14.2.1. Maps were created with SimpleMappr (Shorthouse 2010). GPS data were acquired using Google Maps. Depositories.— ABS Archbold Biological Station Arthropod Collection, Ve- nus, FL, USA; ANSP Academy of Natural Sciences, Philadelphia, PA, USA; CUIC Cornell University Insect Collection, Ithaca, NY, USA; FSCA Florida State Collection of Arthropods, Gainesville, FL, USA; UMMZ University of Michigan Museum of Zoology, Ann Arbor, MI, USA. Results Key to the species of Tridactylidae found in Florida (modi- fied from Gunther 1975 and Gunther 1977). 1 Larger, body length 3.8-10 mm; hind tarsus present.....Neotridactylus 2 - Smaller, length 3-5 mm; hind tarsus reduced to a tiny flap concealed betweensthe hind tibial: spurs .<.5..1279...04sqsrarttsse tote, a ) & OR ae: e ; . < : - ra cael, Fig. 2. Live nymph and adult images of Ellipes deyrupi sp. nov. A. Dorsal and lateral view of adult from Tiger Creek Preserve; B. Nymph from same locality; C. Adult from Sarasota Court; D. Adult burrowing in sand at Tiger Creek; E. Nymph burrowing at same locality. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) Fig. 3. Male Ellipes deyrupi sp. nov. A. Dorsal view of holotype; B. Lateral view of same; C. Underside of right tegmen showing stridula- tory file. Inset shows entire tegmen; D. Terminalia. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) Fig. 4. Females of upland Ellipes spp. A. Dorsal view of allotype E. deyrupi sp. nov.; B. Lateral view of same; C. Subgenital plate of same; D. Subgenital plate of female Ellipes eisneri from Floral City. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) 138 common wherever tridactylids were found (Fig. 1C, D, F). At the Tiger Creek Preserve, typical plants in the habitat included Aristida sp. grasses, Ware's hairsedge (Bulbostylis warei (Torr.) C.B.Clarke), paper nailwort (Paronychia chartacea Fernald), and sandhill sky- blue lupine (Lupinus cumulicola Small). At the roadside sites east of Lake Marion, typical plants included paper nailwort (Paronychia chartacea Fernald), sandhill sky-blue lupine (Lupinus cumulicola Small), scrub Stylisma (Stylisma abdita Myint), prickly pear cac- tus (Opuntia sp.), and Panicum sp. grasses along the road's edge. Lichens present in this habitat included Cladonia leporina Fr., C. evansii Abbayes, and C. subtenuis (Abbayes) Mattick. Photographs of nymphs and adults of E. deyrupi are posted by the author to BugGuide.net for public viewing, at https://bug- guide.net/node/view/1575492/bgimage Ellipes eisneri Deyrup, 2005 Figs 6-8 Ellipes eisneri Deyrup, 2005: 142. Type material.—Holotype: USA ¢ 3; Florida: Citrus County: near Inverness, Withlacoochee Forest, Citrus Area, Forest Road 13, 1.3 mi. south State Road 44; 03 Apr. 1995; sandhill habitat with bare sand, dug from vertical burrow; M. Deyrup leg.; FSCA [not examined]. Paratypes: USA e 4 3, 4 9; same information as ho- lotype; dry pinned specimens; 1 3’, 1 9, FSCA; 1 3, 1 2, ANSP e 2 3, 10 9; “Pine Oak Estates”, State Road 488, 3.7 mi. south of junction with U.S. 41; 04 Ap. 1996; sandhill area with bare sand; specimens in ethanol; M. Deyrup leg.; ABS [examined]. Material examined.—USA, Florida ¢ 2 9;Citrus County: Floral City, side of road on Bedford Dr.; 28.746, -82.299; 28 Jul. 2018; speci- mens in ethanol; B. Woo leg.; CUIC ¢ 1 9;Citrus County, Lecanto, side of road on 490; 28.832, -82.519; 28 Jul. 2018; photograph; B. Woo leg.; CUIC ¢ 1 2, 5 nymphs, Marion County, Dunnellon, side of road on SW Rainbow Lakes Blvd; 29.140, -82.493; 29 Jul. 2018; specimens in ethanol; B. Woo leg.; CUIC ¢ 4 9, 4 nymphs;; Mar- ion County, Dunnellon, round dune off Indian Hill Dr.; 29.169, -82.531; 29 Jul. 2018; specimens in ethanol; B. Woo leg.; CUIC ¢ 1 nymph; Levy County, Bronson, side of road on NE 75" St.; 29.443, -82.539; 30 Jul. 2018; specimen in ethanol; B. Woo leg.; CUIC. Remark.—These are the only two sites from which the species has been known prior to this work. Herein, I present the following specimen data and localities. For photographs of living and pre- served specimens of E. eisneri, see Figs 7 and 8. Distribution.—Found on the Brooksville Ridge in Levy, Marion, and Citrus Counties, FL (Fig. 5). Probably also in adjacent Her- nando County. Habitat.—Specimens were collected in bare sand patches within open sandhill habitats (Fig. 6A, C, E). These areas have scat- tered lichens and forbs and are usually surrounded by oaks and pines. Tumuli and raised feeding burrows were observed at all sites (Fig. 6B, D). Typical plants present included longleaf pine (Pinus palustris Mill.), chapman oak (Quercus chapmanii Sarg.), turkey oak (Quercus laevis Walter), oak ridge lupine (Lupinus diffusus Nutt.), and dogfennel (Eupatorium capillifolium (Lam.) Small) along the road’s edge. Lichens present in these habitats included Cladonia leporina Fr., C. evansii Abbayes, and C. subtenuis (Abbayes) Mattick. B. WOO Fig. 5. Distribution of Florida upland Ellipes spp. Green stars indi- cate collection localities of Ellipes deyrupi sp. nov.; red dots indicate previous (2005) known collection localities for Ellipes eisneri; blue dots indicate newly reported collection localities for E. eisneri. Discussion Ellipes deyrupi represents the third species of Tridactylidae known to be restricted to sandy Florida uplands. Its discovery vindicates Deyrup’s 1996 and 2005 statements that more species were likely to be found in such habitats and suggests that there are still more undiscovered species hiding in similar disjunct xeric habitats of the southeastern U.S. The new species can be collected using yellow bowl traps after a rain or by sifting in the sand where tumuli and feeding burrows are present. In the right conditions, it can be quite abundant, but only those knowing what to look for are likely to come across it. The genus Ellipes is now known to contain two rather distinct upland species with restricted ranges in Florida, whereas populations of scrub Neotridactylus are more ambiguous, perhaps consisting of only one wide-ranging, variable scrub species in a similar situation to the Florida endemic cock- roach Arenivaga floridensis Caudell, which is also restricted to scrub and sandhill (Lamb et al. 2018). The status of scrub Neotridactylus will be explored in a forthcoming paper. The situation of the two species of scrub Ellipes, in which there is one species (E. eisneri) on the Brooksville Ridge and another species (E. deyrupi) on the northern LWR, shares several similari- ties with that of Florida Geopsammodius Gordon & Pittino, 1992 scarab beetles. These blind, flightless beetles also live just below the surface of the sand in scrub and sandhill habitats, and the au- thor often found and collected them while searching for tridac- tylids (specimens are in the CUIC). Geopsammodius withlacoochee Skelley, 2006 is a Brooksville Ridge endemic like E. eisneri, and G. morrisi Skelley, 2006 is only known from the ridge habitat around Lake Marion (including the Catfish Creek Preserve), where E. deyrupi was found (Skelley 2006, Deyrup and Carrel 2012). The circumstances are somewhat different, however, as E. deyrupi also occurs at the Tiger Creek Preserve, which is inhabited by the more widespread southern LWR scarab G. relictillus (Deyrup & Wood- JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) Fig. 6. Habitat and traces of Ellipes eisneri. A. Habitat at Floral City; B. Tumuli at same locality; C. Habitat at SW Rainbow Lakes Blvd; D. Feeding burrow at same locality; E. Habitat at Indian Hill Dr. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) 140 B. WOO Fig. 7. Live nymph and adult images of Ellipes eisneri. A. Dorsal view of adult from Lecanto; B. Dorsal and lateral views of adult from Floral City; C. Lateral view of adult from Floral City, showing lack of tarsus; D. Nymph from Floral City; E. Nymph burrowing in sand at SW Rainbow Lakes Blvd. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) B. WOO 14] Fig. 8. Dorsal and lateral view of female Ellipes eisneri from Floral City. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) 142 ruff, 1991). More extensive surveys of both scrub tridactylids and Geopsammodius scarab beetles may reveal more about the biogeo- graphic patterns underlying their distributions in the state. The ecology and behavior of scrub tridactylids is largely un- known. The only species that has been studied in any detail at all is Neotridactylus archboldi, which feeds on a layer of subsurface algae (Deyrup and Eisner 1996). Adults and nymphs of Ellipes deyrupi have been collected from May to August and in Decem- ber; they likely occur year round but would be rather difficult to locate during periods of little to no rain. The males of E. deyrupi, like many other tridactylids, have a stridulatory apparatus com- prised of a scraper on the underside of the tegmen and a file on the fourth abdominal tergite. Deyrup and Eisner (1996) specu- lated that song may be an important isolating mechanism in tri- dactylids, but nothing is known of the nature or function of their calls, which are likely transmitted through the sand substrate. E. deyrupi co-occurs with N. archboldi at all locations where it has been found, often in the same patches of sand. In other areas of Florida that have been surveyed for scrub tridactylids, only flight- less Neotridactylus have been found, except on the Brooksville Ridge, where apparently only E. eisneri occurs. The fully winged Ellipes minuta sometimes occurs in scrub and sandhill (pers. obs), but this generalist species can fly and is likely present in these habitats only temporarily, when wet conditions prevail. Thus, the northern LWR appears to be the only known habitat where two scrub-restricted tridactylid species occur in sympatry. This brings up questions about interspecific competition for limited space and food resources. A vial containing live adults and nymphs of E. deyrupi and N. archboldi in sand substrate from the same local- ity was found to only contain N. archboldi after two months. This is, of course, anecdotal evidence, but it suggests that N. archboldi may be a better competitor in situations where space and food are limited. Geopsammodius scarab beetles, which likely feed on detritus in the sand (Skelley 2006), were often encountered in the same patches of sand, representing another potential competitor in these limited habitats. Ellipes deyrupi may be a species of conservation concern. If the species occurred further south on the LWR, Deyrup and Car- rel (2012) would most likely have detected it. The majority of its habitat on the northern LWR has been converted to housing or agriculture, and roadside sites where it still occurs are at risk of being developed. A thorough Google Maps search of this part of Polk County turned up no obvious unsurveyed areas that might contain E. deyrupi. In addition, the author could find no suitable E. deyrupi habitat while driving around in the intervening land between the three collection localities. At the Hickory Lake Scrub, only about 16.4 km south of the Tiger Creek Preserve, an hour- long search in December 2019 by the author turned up many N. archboldi but no E. deyrupi. At present, there are only two protected localities for the species, although fortunately these two preserves host large populations of E. deyrupi. Threats as described by Dey- rup and Carrel (2012) include its restriction to open bare sand patches within Florida scrub and specifically within the unique yellow sand scrub found only on the LWR. This habitat can be quickly overgrown by vegetation without prescribed burns, but too frequent fires can convert it to sandhill habitat. Thus, yellow sand scrub management requires a careful balance to maintain its distinctive qualities and organisms. The new records for E. eisneri reported here extend its range to the Northern Brooksville Ridge, and my suspicion is that it likely occurs across the entire ridge in suitable habitat. Its occurrence B. WOO on the northern part of the ridge is interesting because the other Brooksville Ridge endemic Orthopteran, the flightless grasshop- per Melanoplus withlacoocheensis Squitier et al., 1998, is restricted to the southern part of the ridge, with rigorous collecting efforts failing to locate it on the northern Brooksville ridge (Squitier et al. 1998). This was hypothesized to be due to the bisection of these two areas by the Withlacoochee River, which would serve as an effective barrier to flightless insects tied to specific habitats. Yet, E. eisneri occurs on both the Northern and Southern Brooksville Ridge, implying either that E. eisneri was historically present on the Ridge before the formation of the river or that it can navigate aquatic habitats. It might also have been possible for a population of tridactylids to be transported to the Northern Brooksville Ridge in bulk sand, perhaps due to human activity. Phylogenetic analy- sis of E. eisneri populations from both the northern and southern parts of the Brooksville Ridge might shed light on this situation. Ellipes eisneri was relatively abundant wherever found, judging by the number of tumuli and burrows present at all sites, so it is most likely not a rare species within its known range. While its distribution appears to be wider than previously known, the fact remains that it has not been found on any other of Florida’s sand ridges, and that most of the open sandhill habitat on the Brooks- ville ridge is under threat from human development. The Withla- coochee State Forest (Citrus Tract) is still the only known pro- tected site for E. eisneri, as the new localities reported here are all roadside sandhill habitats. The Bronson site, however, is adjacent to the Watermelon Pond Wildlife and Environmental Area, owned by the Florida Fish and Wildlife Conservation Commission. It is almost certain that there are populations of E. eisneri present in the extensive sandhills of this protected area. The nearby Goethe State Forest and Goethe Wildlife Management Area are also likely to hold populations of E. eisneri. It would be prudent to sample within these localities during rainy weather to determine whether E. eisneri occurs there. Acknowledgements I am extremely grateful to Mark Deyrup for offering me the chance to describe this species and for encouraging and support- ing my work on Florida tridactylids. I thank the staff of Archbold Biological Station for providing food and housing. Thanks are also due to Beatriz Pace-Aldana of the Nature Conservancy for granting me permission to collect specimens at the Tiger Creek Preserve and for providing helpful information about the area. Jason Dom- broskie reviewed the original manuscript and has supported my studies of tridactylids for several years. Kyhl Austin assisted with the Cognisys focus-stacking system and zoom lens. Ann Dunn ac- companied me on various collecting trips and shared my enthusi- asm for tiny, understudied animals. Angela Soto, Seth Raynor, and Scott Ward provided plant and lichen identifications. A portion of this work was supported by a summer internship at the Archbold Biological Station to B. Woo. I thank the two reviewers for their comments on the manuscript. The Orthopterists’ Society provided funding for publishing this paper. References Caudell AN (1918) Two new species of the blattid genus Arenivaga. Proceedings of the Entomological Society of Washington 20: 154-157. Choate PM (1984) A new species of Cicindela Linnaeus (Coleoptera: Cicindelidae) from Florida, and elevation of C. abdominalis scabrosa Schaupp to species level. Entomological News 95: 73-82. JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2) B. WOO Deyrup M (1989) Arthropods endemic to Florida scrub. Florida Scientist 52: 254-270. Deyrup M (1990) Arthropod footprints in the sands of time. Florida Ento- mologist 73: 529-538. https://doi.org/10.2307/3495269 Deyrup M (2005) A new species of flightless pygmy mole cricket from a Florida sand ridge (Orthoptera: Tridactylidae). Florida Entomologist 88: 141-145. https://doi.org/10.1653/0015-4040(2005)088[0141:AN SOFP]2.0.CO;2 Deyrup M, Carrel JE (2012) Conservation Status and Management of Lake Wales Ridge Arthropods Restricted to Scrub Habitat. Deyrup M, Eisner T (1996) Description and natural history of a new pygmy mole cricket from relict xeric uplands of Florida (Orthoptera: Tridactyli- dae). Memoirs of the Entomological Society of Washington 17: 59-67. Deyrup M, Woodruff RE (1991) A new flightless Psammodius from Florida’s in- land dunes (Coleoptera: Scarabaeidae). Coleopterists Bulletin 45: 75-80. Gordon RD, Pittino R (1992) Current status of the American genera and species of Psammodiini (Coleoptera: Scarabaeidae: Aphodiinae). Co- leopterists Bulletin 46: 260-273. Gunther KK (1975) Das Genus Neotridactylus Giinther, 1972. (Tridactyli- dae, Saltatoria, Insecta). Zoosystematics and Evolution 51: 305-365. https://doi.org/10.1002/mmnz.19750510208 Gunther KK (1977) Revision of the genus Ellipes Scudder, 1902 (Saltatoria, Tridactylidae). German Entomological Journal 24: 47-122. https:// doi.org/10.1002/mmnd.4800240104 Lamb T, Justice TC, Brewer MS, Moler PE, Hopkins H, Bond JE (2018) A biogeographical profile of the sand cockroach Arenivaga floridensis and its bearing on origin hypotheses for Florida scrub biota. Ecology and Evolution 8: 5254-5266. https://doi.org/10.1002/ece3.3885 143 Menges ES (1999) Ecology and conservation of Florida scrub. In: An- derson RC, Fralish JS, Baskin JM (Eds) (2007) Savannas, bar- rens, and rock outcrop plant communities of North America. Cambridge University Press, 7-22. https://doi.org/10.1017/ CBO9780511574627.002 Myers RL (1985) Fire and the dynamic relationship between Florida sand- hill and sand pine scrub vegetation. Bulletin of the Torrey Botanical Club 241-252. https://doi.org/10.2307/2996539 Say T (1825) Descriptions of new Hemipterous Insects collected in the Ex- pedition to the Rocky Mountains. Journal of the Academy of Natural Sciences, Philadelphia 4: 310. Scudder SH (1862) Materials for a monograph of the North American Or- thoptera, including a catalogue of the known New England species. Boston Journal of Natural History 7(3): 425. https://doi.org/10.5962/ bhl.part.11211 Scudder SH (1902) On the United States Orthoptera which have been Referred to the Genus Tridactylus. Psyche 9: 309. https://doi. org/10.1155/1902/35962 Shorthouse DP (2010) SimpleMappr, an online tool to produce publica- tion-quality point maps. [Retrieved from:] https://www.simplemappr. net [Accessed November 19, 2019] Skelley PE (2006) A revision of the genus Geopsammodius Gordon and Pit- tino, 1992 (Scarabaeidae: Aphodiinae: Psammodiini). Insecta Mundi 20: 101-112. Squitier JM, Deyrup M, Capinera JL (1998) A new species of Mela- noplus (Orthoptera: Acrididae) from an isolated upland in pen- insular Florida. Florida Entomologist 81: 451-460. https://doi. org/10.2307/3495936 JOURNAL OF ORTHOPTERA RESEARCH 2021, 30(2)