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ecology and conservation in extreme south
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<paragraph id="DB585652FFCDBC5AFF32F8FFFF081345" blockId="7.[142,812,1794,2024]" box="[142,222,1794,1817]" pageId="7" pageNumber="29">Florida.</paragraph>
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Our study is one of very few empirical studies of tortoises at the extreme southern limit of their geographic range (but see
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;
<bibRefCitation id="BF762BA3FFCDBC5AFD99F8C3FCF1130F" author="Whitfield, S. M. &amp; F. Ridgley &amp; D. Valle &amp; N. Atteberry" box="[549,807,1854,1876]" pageId="7" pageNumber="29" pagination="47 - 49" refId="ref10198" refString="Whitfield, S. M., F. Ridgley, D. Valle, and N. Atteberry. 2018. Seroprevalence of Mycoplasma agassizii and Mycoplasma testudineum in wild and waif Gopher Tortoises (Gopherus polyphemus) in Miami-Dade County, Florida, USA. Herpetological Review 49: 47 - 49." type="journal article" year="2018">Whitfield et al., 2018</bibRefCitation>
;
<bibRefCitation id="BF762BA3FFCDBC5AFF32F8A6FEA5132E" author="Figueroa, A. &amp; J. Lange &amp; S. M. Whitfield" box="[142,371,1883,1906]" pageId="7" pageNumber="29" pagination="27 - 34" refId="ref8241" refString="Figueroa, A., J. Lange, and S. M. Whitfield. 2021. Seed consumption by Gopher Tortoises (Gopherus polyphemus) in globally imperiled pine rockland ecosystems of South Florida, USA. Chelonian Conservation &amp; Biology 20: 27 - 34." type="journal article" year="2021">Figueroa et al., 2021</bibRefCitation>
).
<bibRefCitation id="BF762BA3FFCDBC5AFE3AF8A6FD16132E" author="Kushlan, J. A. &amp; F. J. Mazotti" box="[390,704,1883,1906]" pageId="7" pageNumber="29" pagination="231 - 239" refId="ref8772" refString="Kushlan, J. A., and F. J. Mazotti. 1984. Environmental effects on a coastal population of Gopher Tortoises. Journal of Herpetology 18: 231 - 239." type="journal article" year="1984">Kushlan and Mazotti (1984)</bibRefCitation>
studied a coastal population of tortoises at Cape Sable (Everglades National Park, Monroe County,
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), an isolated formation of coastal dunes on the extreme southwest of Floridas mainland. This is likely the most climatically similar site to the localities we describe, although the geology and ecology of the Cape Sable population is quite different with regard to vegetation and soils, and with responses to intense tropical storms and hurricanes. Still,
<bibRefCitation id="BF762BA3FFCDBC5AFB25FEF6FAAE157E" author="Waddle, J. H. &amp; F. J. Mazzotti &amp; K. G. Rice" box="[1177,1400,267,290]" pageId="7" pageNumber="29" pagination="277 - 284" refId="ref10155" refString="Waddle, J. H., F. J. Mazzotti, and K. G. Rice. 2006. Changes in abundance of Gopher Tortoise burrows at Cape Sable, Florida. Southeastern Naturalist 5: 277 - 284." type="journal article" year="2006">Waddle et al. (2006)</bibRefCitation>
documented dramatic declines in this population between the 1980s and 2000 and suggested that storm surge from repeated hurricanes had destabilized the sand dunes used by
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there. While both Zoo Miami and Deering were heavily impacted by hurricanes (including Hurricane Andrew and Hurricane Irma), the geology of these sites is apparently not as susceptible to storm impacts as is Cape Sable, as the limestone matrix interspersed with sand may prevent destabilization of sand formations.
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While we were able to identify a considerable number of burrows at each site, few of the burrows we observed were occupied. Only seven burrows at Zoo Miami were occupied and only nine at Deering were occupied. Past researchers have attempted to estimate the number of tortoises based on the number of burrows, but use of such correction factors often leads to inaccurate assessments of population size (
<bibRefCitation id="BF762BA3FFCDBC5AFCDFFD02FC3F1748" author="Burke, R." box="[867,1001,767,789]" pageId="7" pageNumber="29" pagination="92 - 93" refId="ref7932" refString="Burke, R. 1989. Burrow-to-tortoise conversion factors: comparison of three Gopher Tortoise survey techniques. Herpetological Review 20: 92 - 93." type="journal article" year="1989">Burke, 1989</bibRefCitation>
;
<bibRefCitation id="BF762BA3FFCDBC5AFC48FD02FB3F1748" author="Breininger, D. R. &amp; P. A. Schmalzer &amp; C. R. Hinkle" box="[1012,1257,767,789]" pageId="7" pageNumber="29" pagination="317 - 321" refId="ref7839" refString="Breininger, D. R., P. A. Schmalzer, and C. R. Hinkle. 1991. Estimating occupancy of Gopher Tortoise (Gopherus polyphemus) burrows in coastal scrub and slash pine flatwoods. Journal of Herpetology 25: 317 - 321." type="journal article" year="1991">Breininger et al., 1991</bibRefCitation>
;
<bibRefCitation id="BF762BA3FFCDBC5AFB48FD02FC43176F" author="McCoy, E. D. &amp; H. R. Mushinsky" pageId="7" pageNumber="29" pagination="402 - 407" refId="ref8907" refString="McCoy, E. D., and H. R. Mushinsky. 1992. Studying a species in decline: gopher tortoises and the dilemma of '' correction factors. '' Herpetologica 48: 402 - 407." type="journal article" year="1992">McCoy and Mushinsky, 1992</bibRefCitation>
). Still, these two sites appear to represent small and isolated, but reproductive populations. More formal efforts to estimate population size will be critical to understanding management scenarios for the tortoises.
</paragraph>
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The continued presence of
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in pine rocklands is particularly surprising given the isolation of these preserves within a dense urban matrix. These tortoises face direct impacts from habitat loss, risk of injury or mortality by automobile collision, and direct displacement by people. In fact, despite extensive ongoing conversion of natural areas to low-density urban habitats, the state wildlife agency has (to our knowledge) never required surveys or issued removal permits for
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within the entirety of Miami-Dade County.
</paragraph>
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The rocky soils of the pine rocklands have been considered low quality tortoise habitat by both past biologists (
<bibRefCitation id="BF762BA3FFCDBC5AFA19FB28FB9D1155" author="Auffenberg, W. &amp; R. Franz" pageId="7" pageNumber="29" pagination="95 - 126" refId="ref7612" refString="Auffenberg, W., and R. Franz. 1982. The status and distribution of the Gopher Tortoise (Gopherus polyphemus), p. 95 - 126. In: North American Tortoises: Conservation and Ecology. R. B. Bury (ed.). US Department of the Interior, US Fish and Wildlife Service Wildlife Research Report No. 12, Washington, D. C." type="book chapter" year="1982">Auffenberg and Franz, 1982</bibRefCitation>
) and by state wildlife agencies (FWC, 2012), yet we are unaware of any empirical evidence for this claim. As we show, the rocky substrate appears to limit burrow depth; yet it is unclear that deep burrows are necessary in a subtropical climate or that such shallow burrows directly lead to a fitness disadvantage. In contrast, the frost-free climate of south
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allows nearly year-round tortoise activity (
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) and may shorten time to maturity (
<bibRefCitation id="BF762BA3FFCDBC5AFB94FA23FAFA11A8" author="Mushinsky, H. R. &amp; D. S. Wilson &amp; E. D. McCoy" box="[1064,1324,1501,1524]" pageId="7" pageNumber="29" pagination="119 - 128" refId="ref9139" refString="Mushinsky, H. R., D. S. Wilson, and E. D. McCoy. 1994. Growth and sexual dimorphism of Gopherus polyphemus in central Florida. Herpetologica 50: 119 - 128." type="journal article" year="1994">Mushinsky et al., 1994</bibRefCitation>
) or enable higher reproductive rates (
<bibRefCitation id="BF762BA3FFCDBC5AFB85FA06FACC124E" author="Ashton, K. G. &amp; R. Burke &amp; J. Layne" box="[1081,1306,1531,1554]" pageId="7" pageNumber="29" pagination="355 - 363" refId="ref7522" refString="Ashton, K. G., R. Burke, and J. Layne. 2007. Geographic variation in body and clutch size of Gopher Tortoises. Copeia 2007: 355 - 363." type="journal article" year="2007">Ashton et al., 2007</bibRefCitation>
) than in the more northern portions of the range.
</paragraph>
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It may appear surprising that a state-threatened species that constructs rather conspicuous burrows could go nearly unreported in an urban area for two decades (
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;
<bibRefCitation id="BF762BA3FFCDBC5AFC1FF973FBAB12F9" author="Krysko, K. L. &amp; K. M. Enge &amp; E. M. Donlan &amp; E. A. Golden &amp; J. P. Burgess &amp; K. W. Larson" box="[931,1149,1678,1701]" pageId="7" pageNumber="29" pagination="132 - 142" refId="ref8713" refString="Krysko, K. L., K. M. Enge, E. M. Donlan, E. A. Golden, J. P. Burgess, and K. W. Larson. 2010. The non-marine herpetofauna of Key Biscayne, Florida. Herpetological Conservation and Biology 5: 132 - 142." type="journal article" year="2010">Krysko et al., 2010</bibRefCitation>
). However, such data deficiencies are common for fauna within the pine rocklands of south
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. The
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(
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), a south
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endemic found in pine rocklands that is now listed under the ESA, was only formally described as a species in 2004 (
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). The rockland endemic Rim Rock Crowned Snake (
<taxonomicName id="1CE72DD1FFCDBC5AFB06F8C3FAA61308" authorityName="Telford" authorityYear="1966" box="[1210,1392,1854,1876]" class="Reptilia" family="Colubridae" genus="Tantilla" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="29" phylum="Chordata" rank="species" species="oolitica">
<emphasis id="E9938A40FFCDBC5AFB06F8C3FAA61308" box="[1210,1392,1854,1876]" italics="true" pageId="7" pageNumber="29">Tantilla oolitica</emphasis>
</taxonomicName>
) has been documented by biologists fewer than 40 times (
<bibRefCitation id="BF762BA3FFCDBC5AFA0DF8A1FC4213D3" author="Hines, K. N." pageId="7" pageNumber="29" pagination="352 - 356" refId="ref8492" refString="Hines, K. N. 2011. Status and distribution of the Rim Rock Crowned Snake, Tantilla oolitica. Herpetological Review 42: 352 - 356." type="journal article" year="2011">Hines, 2011</bibRefCitation>
;
<bibRefCitation id="BF762BA3FFCDBC5AFC1BF887FB9313CC" author="USFWS &amp; U. S. Fish &amp; Wildlife Service" box="[935,1093,1913,1936]" pageId="7" pageNumber="29" pagination="37568 - 37579" refId="ref10073" refString="USFWS (U. S. Fish and Wildlife Service). 2015. Endangered and Threatened Wildlife and Plants; 90 - Day Findings on 31 Petitions. Rim Rock Crowned Snake in Florida. 80 FR 37568 - 37579." type="journal article" year="2015">USFWS, 2015</bibRefCitation>
). The pine rockland endemic Miami Tiger Beetle (
<taxonomicName id="1CE72DD1FFCDBC5AFC55F86BFB1E13F0" authorityName="Cartwright" authorityYear="1939" box="[1001,1224,1942,1964]" class="Insecta" family="Carabidae" genus="Cicindelidia" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="29" phylum="Arthropoda" rank="species" species="floridana">
<emphasis id="E9938A40FFCDBC5AFC55F86BFB1E13F0" box="[1001,1224,1942,1964]" italics="true" pageId="7" pageNumber="29">Cicindelidia floridana</emphasis>
</taxonomicName>
) was undetected from 1934 until its rediscovery in 2007 (
<bibRefCitation id="BF762BA3FFCDBC5AFB23F849FAA01397" author="Brzoska, D. &amp; C. B. Knisley &amp; J. Slotten" box="[1183,1398,1972,1995]" pageId="7" pageNumber="29" pagination="1 - 7" refId="ref7888" refString="Brzoska, D., C. B. Knisley, and J. Slotten. 2011. Rediscovery of Cicindela scabrosa floridana Cartwright (Coleoptera: Cicindelidae) and its elevation to species level. Insecta Mundi 2011: 1 - 7." type="journal article" year="2011">Brzoska et al., 2011</bibRefCitation>
) and is also now protected under the ESA (
<bibRefCitation id="BF762BA3FFCDBC5AFB08F82FFA9913B4" author="USFWS &amp; U. S. Fish &amp; Wildlife Service" box="[1204,1359,2001,2024]" pageId="7" pageNumber="29" pagination="68985 - 69007" refId="ref10115" refString="USFWS (U. S. Fish and Wildlife Service). 2016. Endangered and Threatened Wildlife and Plants; Endangered Species Status for the Miami Tiger Beetle (Cicindelidia floridana). 81 FR 68985 - 69007." type="journal article" year="2016">USFWS, 2016</bibRefCitation>
). Two butterfly species and more than ten pine rockland plant species have been added to the ESA since 2010 (
<bibRefCitation id="BF762BA3FFC2BC55FE48FF2CFD5A14BA" author="Salvato, M. H." box="[500,652,209,231]" pageId="8" pageNumber="30" pagination="53 - 57" refId="ref9592" refString="Salvato, M. H. 2003. Butterfly conservation and hostplant fluctuations: the relationship between Strymon acis bartrami and Anaea troglodyta floridalis on Croton linearis in Florida (Lepidoptera: Lycaenidae and Nymphalidae). Holarctic Lepidoptera 10: 53 - 57." type="journal article" year="2003">Salvato, 2003</bibRefCitation>
;
<bibRefCitation id="BF762BA3FFC2BC55FD25FF2CFF1A1559" author="Possley, J. &amp; S. Hodges &amp; E. Magnaghi &amp; J. Maschinski" pageId="8" pageNumber="30" pagination="81 - 87" refId="ref9486" refString="Possley, J., S. Hodges, E. Magnaghi, and J. Maschinski. 2016. Distribution of Croton linearis in Miami-Dade County preserves with potential for supporting the federally endangered butterflies Strymon acis bartrami and Anaea troglodyta floridalis. Natural Areas Journal 36: 81 - 87." type="journal article" year="2016">Possley et al., 2016</bibRefCitation>
;
<bibRefCitation id="BF762BA3FFC2BC55FF60FF12FE781559" author="Trotta, L. B. &amp; B. Baiser &amp; J. Possley &amp; D. Li &amp; J. Lange &amp; S. Martin &amp; E. B. Sessa" box="[220,430,238,261]" pageId="8" pageNumber="30" pagination="1735 - 1747" refId="ref9900" refString="Trotta, L. B., B. Baiser, J. Possley, D. Li, J. Lange, S. Martin, and E. B. Sessa. 2018. Community phylogeny of the globally critically imperiled pine rockland ecosystem. American Journal of Botany 105: 1735 - 1747." type="journal article" year="2018">Trotta et al., 2018</bibRefCitation>
). A large and distinctive pine rockland trapdoor spider,
<taxonomicName id="1CE72DD1FFC2BC55FE14FEF1FD56157E" authorityName="Godwin &amp; Bond" authorityYear="2021" box="[424,640,268,290]" class="Arachnida" family="Halonoproctidae" genus="Ummidia" kingdom="Animalia" order="Araneae" pageId="8" pageNumber="30" phylum="Arthropoda" rank="species" species="richmond">
<emphasis id="E9938A40FFC2BC55FE14FEF1FD56157E" box="[424,640,268,290]" italics="true" pageId="8" pageNumber="30">Ummidia richmond</emphasis>
</taxonomicName>
, was only formally described in 2021, and is apparently also a south
<collectingRegion id="192398B0FFC2BC55FFDBFEB5FF651502" box="[103,179,328,350]" country="United States of America" name="Florida" pageId="8" pageNumber="30">Florida</collectingRegion>
endemic that occurs primarily in pine rocklands (
<bibRefCitation id="BF762BA3FFC2BC55FFD2FE9BFE5B1521" author="Godwin, R. L. &amp; J. E. Bond" box="[110,397,358,381]" pageId="8" pageNumber="30" pagination="1 - 165" refId="ref8418" refString="Godwin, R. L., and J. E. Bond. 2021. Taxonomic revision of the New World members of the trapdoor spider genus Ummidia Thorell (Araneae, Mygalomorphae, Halonoproctidae). ZooKeys 1027: 1 - 165." type="journal article" year="2021">Godwin and Bond, 2021</bibRefCitation>
). A data gap concerning
<taxonomicName id="1CE72DD1FFC2BC55FD0DFE9BFF1E15C6" authority="Tortoises" authorityName="Tortoises" class="Testudines" family="Testudinidae" genus="Gopher" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="30" phylum="Chordata" rank="genus">Gopher Tortoises</taxonomicName>
within natural areas embedded in urban Miami is not exceptional; rather, this is typical of a more expansive data deficiency regarding terrestrial fauna in extreme south
<collectingRegion id="192398B0FFC2BC55FFDBFE23FF6215A8" box="[103,180,478,500]" country="United States of America" name="Florida" pageId="8" pageNumber="30">Florida</collectingRegion>
—and pine rocklands in particular. In this environment, discoveries and rediscoveries of threatened species are surprisingly commonplace.
</paragraph>
<paragraph id="DB585652FFC2BC55FFC3FDC5FE6C178F" blockId="8.[103,773,179,979]" pageId="8" pageNumber="30">
The tortoise populations we examine in this study were anticipated to be extirpated by the beginning of this century (
<bibRefCitation id="BF762BA3FFC2BC55FFD2FD89FE0616D7" author="Auffenberg, W. &amp; R. Franz" box="[110,464,628,651]" pageId="8" pageNumber="30" pagination="95 - 126" refId="ref7612" refString="Auffenberg, W., and R. Franz. 1982. The status and distribution of the Gopher Tortoise (Gopherus polyphemus), p. 95 - 126. In: North American Tortoises: Conservation and Ecology. R. B. Bury (ed.). US Department of the Interior, US Fish and Wildlife Service Wildlife Research Report No. 12, Washington, D. C." type="book chapter" year="1982">Auffenberg and Franz, 1982</bibRefCitation>
) and have been mostly overlooked by both tortoise biologists and management agencies (
<bibRefCitation id="BF762BA3FFC2BC55FF6EFD4DFE33169A" author="Schwartz, T. S. &amp; S. A. Karl" box="[210,485,688,710]" pageId="8" pageNumber="30" pagination="917 - 928" refId="ref9637" refString="Schwartz, T. S., and S. A. Karl. 2005. Population and conservation genetics of the Gopher Tortoise (Gopherus polyphemus). Conservation Genetics 6: 917 - 928." type="journal article" year="2005">Schwartz and Karl, 2005</bibRefCitation>
;
<bibRefCitation id="BF762BA3FFC2BC55FE4EFD4DFD6C169A" author="Smith, L. L. &amp; T. D. Tuberville &amp; R. A. Seigel" box="[498,698,688,710]" pageId="8" pageNumber="30" pagination="326 - 330" refId="ref9674" refString="Smith, L. L., T. D. Tuberville, and R. A. Seigel. 2006. Workshop on the ecology, status, and management of the Gopher Tortoise (Gopherus polyphemus), Joseph W. Jones Ecological Research Center, 16 - 17 January 2003: Final Results and Recommendations. Chelonian Conservation and Biology 5: 326 - 330." type="journal article" year="2006">Smith et al., 2006</bibRefCitation>
; FWC, 2012). This is surprising because
<taxonomicName id="1CE72DD1FFC2BC55FE77FD33FDAD16B8" authority="Tortoise" authorityName="Tortoise" box="[459,635,718,740]" class="Testudines" family="Testudinidae" genus="Gopher" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="30" phylum="Chordata" rank="genus">Gopher Tortoise</taxonomicName>
populations on the western extent of the geographic range have long been protected by the ESA in response to loss of habitat, and because the extent of habitat loss in the southern portion of the range has been far more rapid and extreme. The status of southern populations of
<taxonomicName id="1CE72DD1FFC2BC55FE2EFC99FD801726" authority="Tortoises" authorityName="Tortoises" box="[402,598,868,890]" class="Testudines" family="Testudinidae" genus="Gopher" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="30" phylum="Chordata" rank="genus">Gopher Tortoises</taxonomicName>
should be reexamined, as the severe and ongoing decline in this portion of the species range will likely require increased management attention to avoid extirpation.
</paragraph>
</subSubSection>
</treatment>
</document>