2006 SOUTHEASTERN NATURALIST 5(4):587–598 Distribution and Status of the Cockroach Arenivaga floridensis Caudell, a Florida Sand Ridge Endemic Trip Lamb1,*, Teresa C. Justice1, and Michael Justice1 Abstract - Arenivaga floridensis (Florida sand cockroach) is a fossorial insect restricted to scrub and sandhill communities on ancient ridges in peninsular Florida. Ecologically insular by nature, these ridge communities have experienced further fragmentation and significant reduction following conversion to citrus agriculture and subsequent urbanization. To assess the status of A. floridensis, we updated its known distribution by conducting an extensive field survey of scrub and sandhill habitats throughout the state. Roaches (n = 325) were collected at 47 localities, confirming the species’ continued presence in nine out of eleven counties from which it has been reported, and providing 28 new localities for these nine counties. Moreover, we discovered 15 new populations representing six additional counties for which A. floridensis has either never been recorded (Gilchrist, Indian River, Martin, Palm Beach, and St. Lucie counties) or had records published (Orange County). We collected Arenivaga on 11 named peninsular ridges, including one major (Atlantic Coastal) and three minor (Bell, Gordonville, Orlando) ridges on which this species had not been previously reported. The nine new localities on the Atlantic Coastal Ridge constitute a significant range extension for A. floridensis, making it the most broadly distributed species of Florida’s endemic ridge fauna. Introduction Florida scrub is a xeric shrubland partitioned across a series of relict beach ridges and bars (Menges 1999). Older ridges occupy the central part of the peninsula, whereas younger ridges are roughly coincident with present shorelines (Opdyke et al. 1984). Each major ridge system, arising sequentially and separately, has experienced rounds of isolation from the mainland and intra-ridge fragmentation associated with sea level fluctuation (MacNeil 1950, Webb 1990). Today these ridges retain scrub and related sandhill communities that were widespread throughout Florida from the early to mid-Pleistocene. Reduction and fragmentation of scrub habitat purportedly involved a prevailing shift from arid to mesic climatic conditions in the late Pleistocene (Watts and Hansen 1994, Webb 1990). Nonetheless, scrub has probably persisted on most inland ridges since the early Pleistocene (Webb 1990). Florida scrub occupies moderately to excessively drained quartzipsamment soils and supports a xeromorphic plant community maintained by low-frequency, high-intensity fires (Laessle 1958, Menges 1999). Major vegetation types vary, but typically involve evergreen shrub oaks 1Department of Biology, East Carolina University, Greenville, NC 27858. *Corresponding author – lamba@ecu.edu. 588 Southeastern Naturalist Vol. 5, No. 4 (Quercus chapmanii Sargent, Q. geminata Small, Q. inopina Ashe, Q. myrtifolia Willdenow), Ceratiola ericoides Michaux (Florida rosemary), and/or Pinus clausa (Chapman ex Engelm.) Vasey ex Sarg. (sand pine) (Myers 1990). Although species diversity is relatively low, a high level of endemism distinguishes the Florida scrub. The historic flux of suitable habitat, with its attendant opportunities for isolation, has probably played a central role in speciation of scrub biota. In all, over 90 species are endemic to scrub and scrub/sandhill ecotones on peninsular Florida’s ridges, including some 40 plants (Christman and Judd 1990), 3 vertebrates (Moler 1992, Rodgers et al. 1996), and 50+ arthropods (see Deyrup 1989, and new species descriptions cited below). Much of peninsular Florida’s xeric uplands have been converted to human use (citrus agriculture, urbanization), with concomitant loss of scrub and sandhill habitats. As a result, the once continuous stretches of Florida scrub within ridges have become highly fragmented, rendering a mosaic of small isolated patches. For example, the Lake Wales Ridge, the oldest of Florida’s ridges, has been reduced to a mere 15% of its original upland habitat (Menges 1999). Continued habitat loss and fragmentation have prompted detailed surveys of the ridge’s vertebrate (Ashton 2005, Branch et al. 2003, Clark et al. 1999, Greenberg et al. 1994, Hokit et al. 1999, McCoy et al. 2004, McDonald et al. 1999, Stith et al. 1996) and plant (McDonald and Hamrick 1996, Quintana-Ascencio and Menges 1996) endemics. Far less is known about the arthropod contingent, which probably represents more narrowly endemic forms, i.e., species confined to single ridges (Menges 1999). Aside from descriptions of new species (Deyrup 1996, 2004, 2005; Deyrup and Eisner 1996; Schuster 1994; Squitier et al. 1998), information on arthropods endemic to Florida ridges is limited to annotated lists (e.g., Deyrup 1989, 1990; Deyrup and Franz 1994). As such, even basic knowledge on local and regional distributions of endemic ridge arthropods remains largely incomplete. One of the more widely-distributed ridge arthropods, Arenivaga floridensis Caudell (Florida sand cockroach; Fig. 1), is also the sole eastern representative of a genus associated with arid regions of the American Southwest. In fact, prior to Caudell’s (1918) description, the few specimens of Arenivaga collected in Florida were considered to be conspecific with a southwestern species, A. apacha (Saussure). Members of the genus (family Polyphagidae) are psammophilic and characterized by wingless, fossorial females (Hebard 1920). Hubbell (1961) suggested the ancestral lineage of A. floridensis probably entered Florida during the late Pliocene-early Pleistocene, when prevailing xeric conditions facilitated dispersal of semiarid biota from the American Southwest. Arenivaga floridensis inhabits sandhills as well as scrub and is thought to require patches of open sand (Deyrup 1994a). Specimens have been collected in the burrows of Peromyscus polionotus (Osgood) (oldfield mouse) 2006 T. Lamb, T.C. Justice, and M. Justice 589 (Young 1949) and in Pogonomyrmex badius (Latreille) (harvester ant) mounds (Deyrup 1994a). Males occasionally engage in low, erratic flight at dusk, appearing sporadically under lights at night, whereas females and juveniles are probably completely fossorial (Deyrup 1994a). This species has not been reported outside of peninsular Florida, making it the state’s only endemic cockroach (Atkinson et al. 1990). Reported locality records are restricted to ridge systems in Alachua, Citrus, Clay, Highlands, Lake, Levy, Marion, Polk, Pinellas, Putnam, and Volusia counties (Atkinson et al. 1990, Deyrup 1994a). Here we present the results of a comprehensive field survey designed to examine the distribution and status of A. floridensis. We provide several new records and report significant range extensions for this species based on extensive sampling efforts across Florida’s ridge systems. Figure 1. Male (top) and female Arenivaga floridensis. 590 Southeastern Naturalist Vol. 5, No. 4 Methods Our update of the geographic distribution of Arenivaga floridensis involved (1) queries to entomological collections managers throughout the United States and (2) an extensive field survey of all six major ridge systems, as well as nine minor ridges, throughout peninsular Florida (Fig. 2). We also surveyed appropriate habitat in the Florida panhandle and south- Figure 2. The major (named) and minor (numbered) sand ridge systems of peninsular Florida. Minor ridges, listed north to south, are: (1) Bell, (2) Crescent City, (3) Cotton Plant, (4) Deland, (5) Orlando, (6) Winter Haven, (7) Lakeland, (8) Gordonville, (9) Henry, and (10) Bombing Range. 2006 T. Lamb, T.C. Justice, and M. Justice 591 eastern Georgia. Our survey efforts were conducted intermittently from December 2000 through October 2003, an interim in which we managed to secure samples for every month of the year. Roaches were collected by sifting sand samples through 0.5-m2 wood-framed, wire-mesh sieves. We used a two-tier sieve system, in which samples passed through 7.0-mm2 then 3.0-mm2 screens. Adult roaches were retained in the 7.0-mm2 screen, and all but the smallest nymphs were captured in the 3.0-mm2 screen. Each site involved a minimal 2-person-hr. search effort, during which samples from appropriate microhabitat, (i.e., loose sand adjacent to vegetative cover) were sifted. At night in appropriate habitat, we examined insects attracted to white lights for male roaches. Specimens were preserved in 95% ethanol and subsequently categorized as male, female, or nymph. Collection sites were georeferenced; for those few specimens with unrecorded geographic coordinates, we estimated latitude and longitude based on locality data. Localities were plotted using ArcView V 3.0 (ESRI). Specimens are currently under study to assess possible geographic variation and will be deposited in the arthropod collection at the Archbold Biological Station. Results and Discussion Range revision We sampled 63 sites in scrub or sandhill habitats throughout peninsular Florida, the Florida panhandle, and extreme southeastern Georgia (along Trail Ridge). Roaches (n = 325) were collected at 47 localities (Fig. 3, Appendix 1). The continued presence of roach populations at five historical localities was confirmed for Highlands, Hillsborough, Levy, Polk, and Putnam counties (based on locality data for specimens in the Florida State Collection of Arthropods; see Appendix 1). We also discovered 27 new localities representing several counties in which Arenivaga has been documented previously, including Citrus (n = 1 site), Clay (n = 1), Highlands (n = 8), Lake (n = 3), Levy (n = 1), Marion (n = 2), Polk (n = 10), and Volusia (n = 1). We confirmed the continued presence of Arenivaga in Orange County (n = 1 site)—a county unreported in the literature, but one for which we found unpublished records at the Academy of Natural Sciences, Philadelphia (6 specimens) and University of Michigan Museum of Zoology (1 specimen). Moreover, we discovered 14 new localities representing five additional counties in which Arenivaga has never been recorded: Gilchrist, Indian River, Martin, Palm Beach, and St. Lucie. In all, we collected roaches on 11 named ridges. These included one major (Atlantic Coastal) and three minor ridges (Bell, Gordonville, and Orlando) from which this species has not been previously reported. Records for the Atlantic Coastal Ridge constitute a significant range extension for A. floridensis, with nine localities being discovered along the ridge’s southern sector (from Roseland south to Juno Beach). Of equal geographic significance was our finding of an unreported museum record 592 Southeastern Naturalist Vol. 5, No. 4 (Academy of Natural Sciences, Philadelphia) for the northern portion of Atlantic Coastal Ridge. The locality for this single specimen collected by J. A.G. Rehn on April 7, 1940 is listed simply as Jacksonville, FL. Assuming reasonable geographic accuracy of the Rehn locale, it seems likely that A. floridensis inhabited the length of the Atlantic Coastal Ridge prior to habitat destruction associated with development. Figure 3. Collection localities for Arenivaga floridensis surveyed from December 2000 through October 2003. Stars represent localities where roaches have been collected previously; circles represent new localities; the open square approximates Rehn’s Jacksonville locality from 1940. Locality data are listed in Appendix 1. 2006 T. Lamb, T.C. Justice, and M. Justice 593 Efforts to locate roaches in appropriate habitats in the panhandle (four sites), upper peninsular Florida (six sites), or in southeastern Georgia (six sites along the northern portion of Trail Ridge) were unsuccessful. Despite our failure to recover roaches from these sites, we cannot conclude that Arenivaga is absent at these localities or immediate vicinities. However, it is worth noting that the Mt. Dora Ridge represents the northern range terminus for much of the ridge fauna, with ranges of the few more widespread ridge endemics extending north and west onto the Brooksville Ridge. In both cases, distributional limits lie south and/or east of the Suwannee River, a drainage flanked by extensive wetlands to its north and west. Further west, in Florida’s eastern panhandle, the Apalachicola drainage also represents a well-documented biogeographic barrier (Avise 1992, 2000). For these reasons and in light of our collecting efforts, we consider it likely that the northwestern range boundary of A. floridensis corresponds closely with northern termini of the Bell and Northern Brooksville ridges. Additional sampling will be required to make definitive statements regarding the northern distribution limits of A. floridensis, particularly for suitable scrub or sand hill habitats peripheral to the species’ range as we have now defined it. Microhabitat preferences Observations made during our extensive field efforts allow some preliminary comments on microhabitat preferences of Arenivaga floridensis within scrub and sandhill communities. Roaches were most often collected in loose sand under light leaf litter adjacent to Quercus geminata Small (sand live oak), but we also found individuals under deeper oak leaf litter or frond litter of Serenoa repens Hooker (saw palmetto). A few individuals were sifted from sand aprons in front of Gopherus polyphemus (Daudin) (Gopher Tortoise) burrows, and a single specimen was collected in a nest of the ant Leptothorax pergandei Emery. Localities with open sand, including trail and road edges as well as road banks, also yielded roaches, which were generally present in or near such disturbed sites. We never observed roaches in habitats where sand had become compacted or encrusted. However, we noted that roaches did occur in otherwise overgrown scrub by paths, roads, or similar disturbances if loose, open sand was present. Thus, our field observations corroborate Deyrup’s (1994a) view that A. floridensis “lives only in areas that are managed or allowed to burn periodically to provide patches of open sand.” Although we collected six male roaches near lights at night, the majority of male specimens (n =14) were collected sifting sand. Males, females, and juveniles were collected year-round. Species status The Florida Committee on Rare and Endangered Plants and Animals classified Arenivaga floridensis as rare, based on (1) rapid loss of Florida scrub and sandhills habitat throughout the state and (2) limited population/ locality data (Deyrup 1994a). As noted above, we have documented 42 new localities for this species, providing significant—those sites along 594 Southeastern Naturalist Vol. 5, No. 4 the Atlantic Coastal Ridge, in particular—range extensions. Fortunately, many of these new localities occur on protected lands overseen by private (e.g., The Nature Conservancy), state, or federal agencies (Appendix 1). Moreover, this distributional update makes A. floridensis the most widespread faunal ridge-endemic species, exceeding slightly the geographic range of Schistocerca ceratiola Hubbell and Walker (rosemary grasshopper) (Deyrup 1994b). Clearly, these findings hold promise regarding the status and future of A. floridensis, but they are not without qualification. Habitat specificity, the female’s flightless condition, apparent endemism to scrub and sandhills, and “striking differences in color in different parts of the range” (Deyrup 1994a) make this species an ideal candidate for a detailed study of geographic variation. The potential for long-term isolation among roach populations between ridges provides opportunities for genetic divergence. Morphological, genetic, and phylogeographic surveys of other ridge taxa (Branch et al. 2003, Breininger et al. 1996, Clark et al. 1999, McDonald and Hamrick 1996, McDonald et al. 1999) have repeatedly revealed significant inter-ridge variation that has and will continue to play an important roll in managing biotic diversity of this unique ecosystem. To this end, we are currently examining DNA sequence variation among the populations reported herein to determine whether (and to what degree) A. floridensis exhibits phylogeographic structure comparable to other ridge endemics. Acknowledgments We graciously thank the following sifters for their assistance in the field: Mark Deyrup, Michael Evans, Paul Moler, Jackson Mosely, and Paul Skelley. Mark O’Brien, Paul Skelley, and Jason Weintraub provided locality data for material in their following respective institutions: University of Michigan Museum of Zoology, Florida State Collection of Arthropods, and the Academy of Natural Sciences, Philadelphia. David Beamer helped generate the ridge maps. Research was supported by contract NG01-002 with the Florida Fish and Wildlife Conservation Commission to T. Lamb and T.C. Justice, and by the Theodore Roosevelt Fund and a summer internship at the Archbold Biological Station to T.C. Justice. Literature Cited Ashton, K.G. 2005. Life history of a fossorial lizard, Neoseps reynoldsi. Journal of Herpetology 39:389–395. Atkinson, T.H., P.G. Koehler, and R.S. Patterson. 1990. Annotated checklist of the cockroaches of Florida (Dictyoptera: Blattaria: Blatidae, Polyphagidae, Blattellidae, Bladeridae). Florida Entomologist 73:303–327. Avise, J.C. 1992. 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Transactions of the American Entomological Society 46:197–217. Hokit, D.G., B.M. Stith, and L.C. Branch. 1999. Effects of landscape structure in Florida scrub: A population perspective. Ecological Applications 9:124–134. Hubbell, T.H. 1961. Endemism and speciation in relation to Pleistocene changes in Florida and the southeastern coastal plain. Eleventh International Congress of Entomology, Wein, 1960 1:466–469. Laessle, A.M. 1958. The origin and successional relationship of sandhill vegetation and sand-pine scrub. Ecological Monographs 28:361–387. MacNeil, F.S. 1950. Pleistocene shore lines in Florida and Georgia. Geological Survey Professional Paper 221-F:95–107. McCoy, E.D., P.P. Hartman, and H.R. Mushinsky. 2004. Population biology of the rare Florida Scrub Lizard in fragmented habitat. Herpetologica 60:54–61. McDonald, D.B., and J.L. Hamrick. 1996. Genetic variation in some plants of Florida scrub. American Journal of Botany 83:21–27. 596 Southeastern Naturalist Vol. 5, No. 4 McDonald, D.B., W.K. Potts, J.W. Fitzpatrick, and G.E. Woolfenden. 1999. Contrasting genetic structure in sister species of North American scrub-jays. Proceedings of the Royal Society B, London 266:1117–1125. Menges, E.S. 1999. Ecology and conservation of Florida scrub. Pp. 7–22, In R.C.Anderson, J.S. Fralish, and J. Baskin (Eds.). The Savanna, Barren, and Rock Outcrop Communities of North America. Cambridge University Press, Cambridge, UK. 470 pp. Moler, P.E. 1992. Rare and Endangered Biota of Florida. III. Amphibians and Reptiles. University Press of Florida, Gainesville, FL. 272 pp. Myers, R.L. 1990. Scrub and high pine. Pp. 150–193, In R.L. Myers and J.J.Ewels (Eds.). Ecosystems of Florida. University of Central Florida Press, Orlando, FL. 765 pp. Opdyke, N.D., D.P. Spangler, D.L. Smith, D.S. Jones, and R.C. Lindquist. 1984. Origin of the epeirogenic uplift of Pliocene-Pleistocene beach ridges in Florida and development of the Florida karst. Geology 12:226–228. Quintana-Ascencio, P.F., and E.S. Menges. 1996. Inferring metapopulation dynamics from patch-level incidence of Florida scrub plants. Conservation Biology 10:1210–1027. Rodgers, J.A., H.W. Kale, and H.T. Smith. 1996. Rare and Endangered Biota of Florida. V. Birds. University Press of Florida, Gainesville, FL. 736 pp. Schuster, J.C. 1994. Odontotaenius floridanus new species (Coleoptera: Passalidae): A second US passalid beetle. Florida Entomologist 77:474-479. Squitier, J.M., M. Deyrup, and J.L. Capinera. 1998. A new species of Melanoplus (Orthoptera: Acrididae) from an isolated upland in peninsular Florida. Florida Entomologist 81:451–460. Stith, B.M., J.W. Fitzpatrick, G.E. Woolfenden, and B. Pranty. 1996. Classification and conservation of metapopulations: A case study of the Florida Scrub-jay. Pp. 187–215, In D. McCollough (Ed.). Metapopulations and Wildlife Conservation. Island Press, Washington, DC. Watts, W.A., and B.C.S. Hansen. 1994. Pre-Holocene and Holocene pollen records of vegetation history from the Florida peninsula and their climatic implications. Palaeogeography, Palaeoclimatology, and Palaeoecology 109:163–176. Webb, S.D. 1990. Historical biogeography. Pp. 70–100, In R.L. Myers and J.J. Ewels (Eds.). Ecosystems of Florida. University of Central Florida Press, Orlando, FL. 765 pp. Young, F.N. 1949. Insects from the burrows of Peromyscus polionotus. Florida Entomologist 32:77. 2006 T. Lamb, T.C. Justice, and M. Justice 597 Appendix 1. Collection locality data, including number of individuals collected (n), county, ridge, latitude and longitude (in decimal degrees), and collection date for Arenivaga floridensis. Locality n County Ridge Latitude Longitude Collection date Alexander SpringsB 2 Lake Mt. Dora 29.12346 81.57789 3/16/2002 Archbold Biological StationA,B 10 Highlands Lake Wales 27.18185 81.35217 11/22/2000; 5/16/2001 ArcherA 12 Levy N. Brooksville 29.50267 82.57250 2/16/2002 Bartow 2 Polk Gordonville 27.96510 81.77340 2/21/2003 Bell 4 Gilchrist Bell 29.79037 82.85362 10/15/2001 BrooksvilleB 14 Citrus S. Brooksville 28.79871 82.38119 5/13/2001; 9/3/2001 Carter CreekB 4 Highlands Lake Wales 27.54008 81.40860 8/14/2002 Cedar KeyB 11 Levy not named 29.18520 83.01901 5/19/2003 Clermont A 3 Lake Lake Wales 28.55583 81.74233 5/27/2003 Clermont B 2 Lake Lake Wales 28.54625 81.70100 5/28/2003 Deland 31 Volusia Deland 29.01872 81.23567 12/19/2002 Dundee 2 Polk Lake Wales 28.02648 81.63408 12/17/2001 East of Archbold 2 Highlands Lake Wales 27.23783 81.30600 5/29/2003 FrostproofB 3 Polk Lake Wales 27.69647 81.53964 9/3/2001 Ft. Pierce 9 St. Lucie Atlantic Coastal 27.47035 80.33381 3/13/2002 Gold HeadB 5 Clay S. Trail Ridge 29.82782 81.94990 3/10/2001 Highlands HammockB 7 Highlands Lake Wales 27.46728 81.52069 6/4/2002 Hobe Sound A 4 Martin Atlantic Coastal 27.10840 80.16827 10/17/2001 Hobe Sound B 21 Martin Atlantic Coastal 27.05972 80.14045 3/13/2002; 5/17/2002 J. Dickinson S.P.B 2 Martin Atlantic Coastal 27.01693 80.11015 3/13/2002 Josephine Creek 9 Highlands Lake Wales 27.36860 81.40173 4/19/2003 Juno Beach A 4 Palm Beach Atlantic Coastal 26.86850 80.05610 5/1/2003 Juno Beach B 7 Palm Beach Atlantic Coastal 26.87847 80.05540 5/1/2003 JupiterB 4 Palm Beach Atlantic Coastal 26.91608 80.07305 5/1/2003 Lake Marion 13 Polk Lake Wales 28.07444 81.54659 4/10/2002 598 Southeastern Naturalist Vol. 5, No. 4 Locality n County Ridge Latitude Longitude Collection date Lake Wales 5 Polk Lake Wales 27.91855 81.60485 12/17/2001 Ocala Natlional Forest AB 4 Marion Mt. Dora 29.36503 81.82204 5/14/2001 Ocala Natlional Forest BB 2 Marion Mt. Dora Unavailable Unavailable 5/29/2003 OrdwayA,B 2 Putnam S. Trail Ridge 29.72840 81.97990 9/2/2001 Orlando 3 Orange Orlando 28.54230 81.31950 5/27/2003 Placid Lakes 2 Highlands Lake Wales 27.24169 81.40142 7/3/2002 Roseland 2 Indian River Atlantic Coastal 27.82842 80.47814 3/17/2002 Saddle BlanketB 20 Polk Lake Wales 27.66950 81.57633 4/12/2003 SebringA 6 Highlands Lake Wales 27.44177 81.41863 5/16/01; 9/3/01 Silver Lake 7 Highlands Lake Wales 27.56431 81.52315 6/11/2002 South of Archbold 1 Highlands Lake Wales 27.10675 81.33240 3/8/2002 Sun ‘n Lakes 1 Highlands Lake Wales 27.24863 81.30063 8/2/2002 Sunray A 16 Polk Lake Wales 27.69037 81.56392 3/15/2002 Sunray B 3 Polk Lake Wales 27.70455 81.56120 3/15/2002 TampaA,B 8 Hillsborough not named 28.06980 82.39480 1/13/2002 Viking 3 St. Lucie Atlantic Coastal 27.54285 80.36295 12/22/00 WaltonB 7 St. Lucie Atlantic Coastal 27.29907 80.25837 8/31/01; 10/17/01 Warner Southern 5 Polk Lake Wales 27.70455 81.56120 12/17/2001 Weeki Wachee A 17 Hernando S. Brooksville 28.52517 82.56995 10/19/2002 Weeki Wachee B 2 Hernando S. Brooksville 28.51512 82.57318 10/19/2002 Wekiwa SpringsB 13 Orange Mt. Dora 28.71048 81.46653 12/16/01; 8/7/02 Winter Haven 9 Polk Winter Haven 28.04417 81.72717 5/30/2003 AConfirmed previous locality(Florida State Collection of Arthropods). BProtected site.