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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.
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 – firstname.lastname@example.org.
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
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
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.
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.
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.
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.
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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).