Fleshy-Fruit

Federal Register / Vol. 78, No. 149 / Friday, August 2, 2013 / Proposed Rules 47109

DEPARTMENT OF THE INTERIOR as form letters), our preferred format is respective ranges due to threats related a spreadsheet in Microsoft Excel. to: Fish and Wildlife Service (2) By hard copy: Submit by U.S. mail • For Short’s bladderpod, potential or hand-delivery to: Public Comments future construction and ongoing 50 CFR Part 17 Processing, Attn: FWS–R4–ES–2013– maintenance of transportation rights-of- [Docket No. FWS–R4–ES–2013–0087; 0087; Division of Policy and Directives way; prolonged inundation and soil 4500030113] Management; U.S. Fish and Wildlife erosion due to flooding and water level Service; 4401 N. Fairfax Drive, MS manipulation; overstory shading due to RIN 1018–AZ11 2042–PDM; Arlington, VA 22203. forest succession and shading and We request that you send comments Endangered and Threatened Wildlife competition from invasive, nonnative only by the methods described above. and Plants; Endangered Status for plant species; and small population We will post all information received on Physaria globosa (Short’s bladderpod), sizes. http://www.regulations.gov. This • Helianthus verticillatus (whorled For whorled sunflower, mechanical generally means that we will post any sunflower), and Leavenworthia crassa or chemical vegetation management for personal information you provide us (fleshy-fruit gladecress) industrial forestry, right-of-way (see the Information Requested section maintenance, or agriculture; shading AGENCY: Fish and Wildlife Service, below for more details). and competition resulting from Interior. FOR FURTHER INFORMATION CONTACT: vegetation succession; limited ACTION: Proposed rule. Mary E. Jennings, Field Supervisor, U.S. distribution and small population sizes. Fish and Wildlife Service, Tennessee • For fleshy-fruit gladecress, loss of SUMMARY: We, the U.S. Fish and Ecological Services Field Office, 446 habitat due to residential and industrial Wildlife Service, propose to list Neal Street, Cookeville, TN 38501; by development; conversion of agricultural Physaria globosa (Short’s bladderpod), telephone 931–528–6481; or by sites for use as pasture; mowing and Helianthus verticillatus (whorled facsimile 931–528–7075. Persons who herbicide treatment prior to seed sunflower), and Leavenworthia crassa use a telecommunications device for the production; and off-road vehicles and (fleshy-fruit gladecress) as endangered deaf (TDD) may call the Federal dumping. under the Endangered Species Act of Information Relay Service (FIRS) at We will seek peer review. We are 1973, as amended (Act). If we finalize 800–877–8339. seeking comments from knowledgeable this rule as proposed, it would extend SUPPLEMENTARY INFORMATION: individuals with scientific expertise to the Act’s protections to Physaria review our analysis of the best available globosa (Short’s bladderpod), Executive Summary science and application of that science Helianthus verticillatus (whorled Why we need to publish a rule. Under and to provide any additional sunflower), and Leavenworthia crassa the Act, if we intend to list a species are information to improve this proposed (fleshy-fruit gladecress) to conserve endangered or threatened throughout all rule. Because we will consider all these species. or a significant portion of its range, we comments and information we receive DATES: We will accept all comments are required to promptly publish a during the comment period, our final received or postmarked on or before proposal in the Federal Register to list determinations may differ from this October 1, 2013. Comments submitted the species as endangered or threatened proposal. electronically using the Federal and make a determination on our Information Requested eRulemaking Portal (see ADDRESSES proposal within 1 year. Listing a species section, below) must be received by as an endangered or threatened species We intend that any final action 11:59 p.m. Eastern Time on the closing can only be completed by issuing a rule. resulting from this proposed rule will be date. We must receive requests for This rule proposes to add three plants based on the best scientific and public hearings, in writing, at the to the Federal List of Endangered and commercial data available and be as address shown in the FOR FURTHER Threatened Plants. We are proposing to accurate and as effective as possible. INFORMATION CONTACT section by list Short’s bladderpod, whorled Therefore, we request comments or September 16, 2013. sunflower, and fleshy-fruit gladecress as information from other concerned ADDRESSES: You may submit comments endangered species under the Act. governmental agencies, Native by one of the following methods: Elsewhere in today’s Federal Register, American tribes, the scientific (1) Electronically: Go to the Federal we propose to designate critical habitat community, industry, or any other eRulemaking Portal: http:// for the Short’s bladderpod, freshy-fruit interested parties concerning this www.regulations.gov. In the Search gladecress, and the whorled sunflower. proposed rule. We particularly seek field, enter Docket No. FWS–R4–ES– The basis for our action. Under the comments concerning: 2013–0087, which is the docket number Act, we may determine that a species is (1) The species’ biology, range, and for this rulemaking. Then, in the Search an endangered or threatened species population trends, including: panel on the left side of the screen, based on any of five factors: (A) The (a) Habitat requirements for feeding, under the Document Type heading, present or threatened destruction, reproducing, and sheltering; click on the Proposed Rules link to modification, or curtailment of its (b) Genetics and taxonomy; locate this document. You may submit habitat or range; (B) overutilization for (c) Historical and current range, a comment by clicking on ‘‘Comment commercial, recreational, scientific, or including distribution patterns; Now!’’ If your comments will fit in the educational purposes; (C) disease or (d) Historical and current population provided comment box, please use this predation; (D) the inadequacy of levels, and current and projected trends; feature of http://www.regulations.gov, as existing regulatory mechanisms; or (E) and it is most compatible with our comment other natural or manmade factors (e) Past and ongoing conservation review procedures. If you attach your affecting its continued existence. measures for these species, their comments as a separate document, our We have determined that listing is habitats or both. preferred file format is Microsoft Word. warranted for these species, which are (2) The factors that are the basis for If you attach multiple comments (such currently at risk throughout all of their making a listing determination for a

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species under section 4(a) of the Act, hours, at the U.S. Fish and Wildlife definitions in the Act (58 FR 51144, which are: Service, Tennessee Ecological Services September 30, 1993). (a) The present or threatened Field Office (see FOR FURTHER In 1993, the Service eliminated destruction, modification, or INFORMATION CONTACT). candidate categories, and Short’s curtailment of its habitat or range; bladderpod and the two varieties of (b) Overutilization for commercial, Background fleshy-fruit gladecress were no longer recreational, scientific, or educational Previous Federal Actions candidates until they were again purposes; The Act requires the Service to elevated to candidate status on October (c) Disease or predation; identify species of wildlife and plants 25, 1999 (64 FR 57534). The 1999 (d) The inadequacy of existing review elevated the species regulatory mechanisms; or that are endangered or threatened, based on the best available scientific and Leavenworthia crassa (fleshy-fruit (e) Other natural or manmade factors gladecress) to candidate status, but did affecting its continued existence. commercial data. The Act directed the Secretary of the Smithsonian Institution not recognize intraspecific taxa (3) Biological, commercial trade, or (varieties) due to changes in other relevant data concerning any to prepare a report on endangered and threatened plant species, which was scientifically accepted taxonomy. threats (or lack thereof) to this species Whorled sunflower was first listed as a and regulations that may be addressing published as House Document No. 94–51. The Service published a notice candidate species in the 1999 review. those threats. All three of these species were then (4) Additional information concerning in the Federal Register on July 1, 1975 (40 FR 27824), in which we announced included in subsequent candidate the historical and current status, range, notices of review on October 30, 2001 distribution, and population size of that more than 3,000 native plant taxa named in the Smithsonian’s report and (66 FR 54808), June 13, 2002 (67 FR these species, including the locations of 40657), May 4, 2004 (69 FR 24876), May any additional populations of these other taxa added by the 1975 notice would be reviewed for possible 11, 2005 (70 FR 24870), September 12, species. 2006 (71 FR 53756), December 6, 2007 (5) Current or planned activities in the inclusion in the List of Endangered and (72 FR 69034), December 10, 2008 (73 areas occupied by these species and Threatened Plants. The 1975 notice was FR 75176), November 9, 2009 (74 FR possible impacts of these activities on superseded on December 15, 1980 (45 57804), November 10, 2010 (75 FR them. FR 82480), by a new comprehensive 69222), October 26, 2011 (76 FR 66370), Please note that submissions merely notice of review for native plants that and November 21, 2012 (77 FR 69994). stating support for or opposition to the took into account the earlier action under consideration without Smithsonian report and other Species Information providing supporting information, accumulated information. On November Short’s bladderpod although noted, will not be considered 28, 1983 (48 FR 53640), a supplemental in making a determination, as section plant notice of review noted the status Physaria globosa is a member of the 4(b)(1)(A) of the Act directs that of various taxa. Complete updates of the mustard family (Brassicaceae) known determinations as to whether any plant notice were published on from Posey County, Indiana; Clark, species is an endangered or threatened September 27, 1985 (50 FR 39526) and Franklin and Woodford Counties, species must be made ‘‘solely on the on February 21, 1990 (55 FR 6184). Kentucky; and Cheatham, Davidson, basis of the best scientific and In these reviews, Short’s bladderpod Dickson, Jackson, Montgomery, Smith, commercial data available.’’ (as Lesquerella globosa) was listed as a and Trousdale Counties, Tennessee. The You may submit your comments and Category 2 candidate, taxa for which following description is based on Flora materials concerning this proposed rule information in the possession of the of North America (http:// by one of the methods listed in the Service indicated that proposing to list www.efloras.org/florataxon.aspx?flora_ ADDRESSES section. We request that you the species as endangered or threatened id=1&taxon_id=250095135, accessed on send comments only by the methods was possibly appropriate, but for which December 7, 2012) and Gleason and described in the ADDRESSES section. sufficient data on biological Chronquist (1991, p. 187). If you submit information via http:// vulnerability and threat were not Short’s bladderpod is an upright www.regulations.gov, your entire available to support listing rules. biennial or perennial (lives for 2 years submission—including any personal Further biological research and field or longer) with several stems, some identifying information—will be posted study usually was necessary to ascertain branched at the base, reaching heights on the Web site. If your submission is the status of taxa in this category. up to 50 centimeters (cm) (20 inches made via a hardcopy that includes Fleshy-fruit gladecress was (in.)), and which are leafy to the base of personal identifying information, you recognized as consisting of two varietal the inflorescence (a group or cluster of may request at the top of your document taxa in these reviews, Leavenworthia flowers arranged on a stem that is that we withhold this information from crassa var. crassa and L. crassa var. composed of a main branch or a public review. However, we cannot elongata. In the 1980 review, var. crassa complicated arrangement of branches). guarantee that we will be able to do so. was listed as a Category 2 candidate, The basal leaves, borne on short petioles We will post all hardcopy submissions while var. elongata was listed as a (stalks) are 2.5 to 5 cm (1 to 2 in.) in on http://www.regulations.gov. Please Category 1 candidate, taxa for which the length and 0.5 to 1.5 cm (0.2 to 0.6 in.) include sufficient information with your Service had sufficient information to wide, obovate (egg-shaped and flat, with comments to allow us to verify any support listing as either endangered or the narrow end attached to the stalk) or scientific or commercial information threatened. In the 1983, 1985, and 1990 oblanceolate (with the widest portion of you include. reviews both varieties of Leavenworthia the leaf blade beyond the middle) in Comments and materials we receive, crassa were listed as Category 2 shape, with a smooth or slightly wavy as well as supporting documentation we candidates. Many Category 2 candidate margin, and gray-green in color due to used in preparing this proposed rule, species were found not to warrant a layer of dense hairs. Leaves are will be available for public inspection listing, either because they were not gradually reduced in size and petiole on http://www.regulations.gov, or by endangered or threatened or because length higher up the stem. Numerous appointment, during normal business they did not qualify as species under the flowers are borne on a raceme (elongate,

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spike-shaped inflorescence to which during the prior 25 years and where the current distribution and status of individual flowers are attached by appropriate habitat existed with no Short’s bladderpod. Difficulty in slender pedicels, or stalks, which in evidence that the occurrence had been relating the species’ distribution at the Short’s bladderpod are longer than the destroyed (Shea population numbers 27 time of Shea’s (1993, entire) status flowers). The yellow flowers are through 30). survey to its current distribution comes composed of four spoon-shaped petals, • Extirpated—one occurrence where as a result of State conservation agencies 0.4 to 0.7 cm (0.16 to 0.28 in.) long. The the habitat had been severely altered revising the mapping of some element fruit is globose in shape and lightly (Shea population number 31). occurrences in these databases. In two • beset with stellate (star-shaped) hairs, Historical—5 occurrences where the instances, pairs of occurrences that Shea but becoming smooth with time. species had not been observed during (1993) considered distinct have been Taxonomy. A member of the mustard the prior 25 years, but where combined into single element family (Brassicaceae), Short’s appropriate habitat remained (Shea occurrences (Table 1). Conversely, bladderpod was first described as population numbers 32 through 36). • TNHID (2012) treats as two distinct Vesicaria globosa by Desvaux in 1814 Locality information incomplete— element occurrences the two locations (Payson 1922, pp. 103–236). Because of 14 occurrences for which location that Shea (1993, p. 85, 108) mapped information was insufficient to confirm several distinctive characters, Watson together as population number 23. One the species’ presence or absence, despite (1888, pp. 249–255) proposed that the of these occurrences (TN EO number 22) searches having been attempted in some American species of the genus Vesicaria was extant as of 2012 (Table 1), while cases (Shea population numbers 37 be placed in the genus Lesquerella. This the other (TN EO number 2) is through 50). Many of these putative treatment was recognized as valid, until extirpated (Table 2). Based on current occurrences were based on herbarium Al-Shehbaz and O’Kane (2002, entire) mapping, State conservation agencies specimens dating from the late-19th to reunited most of the genus Lesquerella now recognize 24 element occurrences mid-20th centuries that contained little with the genus Physaria. This that correspond to populations that information about sites from which they determination was supported by Shea (1993, entire) found extant in were collected. Except for the molecular, morphological, cytological, 1992. Of these 24 occurrences, 18 were populations numbered 37, 42, and 50, biogeographic, and ecological lines of extant in 2012. Accounting for evidence (Al-Shehbaz and O’Kane 2002, Shea (1993) searched for suitable habitat or Short’s bladderpod plants in areas rediscovery of the two Tennessee p. 320). Flora of North America occurrences that Shea (1993, pp. 10–14) recognizes this change, using the associated with these occurrences but did not find the species. did not find during 1992, and recent scientific name Physaria globosa for changes in element occurrence Later surveys found Short’s bladderpod Short’s bladderpod (http:// mapping, a total of 20 occurrences that _ extant at two of these sites, Tennessee www.efloras.org/florataxon.aspx?flora were documented by Shea (1993, entire) _ element occurrence (EO) numbers 8 and id=1&taxon id=250095135, accessed on were still considered extant as of 2012 12, which correspond to Shea’s April 20, 2011). (Table 1). Distribution and Status. In a 1992 population numbers 34 and 29, status survey for Short’s bladderpod, respectively. The approximate range of abundance Shea (1993, pp. 6–15) observed the We used data provided to us by shown in Table 1 is primarily based on species at only 26 of 50 historical sites: conservation agencies in the States individual plants. As a result of 1 in Indiana, 14 in Kentucky, and 11 in where the species occurs (Indiana location, it was impossible to enumerate Tennessee. The remaining sites were Natural Heritage Data Center (INHDC) individual plants. This resulted in are classified as follows (Shea 1993, p. 10– 2012, Kentucky Natural Heritage two instances where TNHID surveyed 14): Program (KNHP) 2012, Tennessee these populations from a boat and • Status uncertain—4 occurrences (Tennessee Natural Heritage Inventory reported the approximate range in where the species had been observed Database (TNHID) 2012) to determine clusters.

TABLE 1—LIST OF KNOWN EXTANT SHORT’S BLADDERPOD OCCURRENCES BY STATE AND COUNTY, WITH ELEMENT OC- CURRENCE (EO) NUMBERS ASSIGNED BY STATE NATURAL HERITAGE PROGRAMS (INHDC (2012), KNHP (2012), TNHID (2012)), NUMBERS ASSIGNED TO POPULATIONS REPORTED IN SHEA (1993), AND FIRST AND LAST YEARS OF KNOWN OBSERVATIONS

EO Number Approximate State County (Shea Popu- First Last range of Land ownership lation Number) observed observed abundance

Indiana ...... Posey ...... 1 (1) 1941–05–06 2012 3–1000s ...... IDNR. Kentucky ...... Clark ...... 1 (3) 1957 2009–05–21 2 ...... Private. Franklin ...... 4 (11, 12) 1979 2011–04–19 100–500 ...... Private. 7 (10) 1981 2004–05–17 1–100 ...... Private. 11 (13) 1983 2003–06–01 1–52 ...... Private. 18 (4) 1992 2012–05–09 20–350 ...... City of Frankfort. 22 (9) 1990-Pres 2012–05–08 2–200 ...... private; Ken- tucky State Nature Pre- serves Com- mission. 23 (14) 1990 2011–04–26 60–500 ...... Private. Woodford ...... 28 2005–05–06 2010–06–02 few...... Private. Tennessee ...... Cheatham ...... 1 (18) 1956–03–02 2008–04–23 100s–1000s COE; private. 15 (17) 1955–04–24 2008–04–29 few–20 ...... COE.

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EO Number Approximate State County (Shea Popu- First Last range of Land ownership lation Number) observed observed abundance

17 (16) 1953–04–26 2012–06–15 20–∼1500 .... Town of Ashland City; private. 29 1998–05–12 2008–04–29 ∼50 ...... COE; private. 30 1998–05–12 2008–04–29 10–25...... COE; private. Davidson; Cheatham ...... 10 (21,22) 1935 2012–06–15 10s–1000s .. Private. Davidson ...... 4 (19) 1971–05–16 2012–06–15 100s–1000s private; COE easement. 8 (34) 1886–04–22 2008–05–02 ∼50 ...... private; COE easement. Dickson ...... 32 2008–04–29 2008–04–29 ∼7 clusters ... COE. Jackson ...... 26 1998–05–08 2008–05–06 3 clusters..... COE. 27 1998–05–08 2008–05–06 ∼50 ...... COE. Montgomery ...... 12 (29) 1946–04–27 2008–05–09 ∼50 ...... private; COE easement. 22 (23a) 1969–04–28 2008–05–02 20–50 ...... private; COE easement. 28 1998–04–23 2008–04–29 ∼300 ...... private; COE easement. Smith ...... 24 1998–05–05 2008–05–06 ∼10 ...... COE. Trousdale ...... 3 (25) 1969–05–08 2008–05–06 40–500 ...... COE; private. 21 (26) 1992–04–30 2008–05–12 100–250 ...... COE; private. IDNR is the Indiana Department of Natural Resources. COE is the U.S. Army Corps of Engineers. Pres is present.

Despite the rediscovery of the two the loss of 10 occurrences during the after 1992, have since been extirpated Tennessee occurrences and the last 20 years (Table 1). Seven of the (Table 2). This constitutes a loss of 27 discovery of 10 additional occurrences occurrences that Shea (1993, pp. 44–71) percent of all occurrences that were since the 1992 status survey, only 26 observed in 1992, and three others extant during 1992 or later. extant occurrences of Short’s (Kentucky EO number 27 and Tennessee bladderpod are known to remain due to EO numbers 23 and 25) that were seen

TABLE 2—LIST OF EXTIRPATED SHORT’S BLADDERPOD OCCURRENCES BY STATE AND COUNTY, WITH ELEMENT OCCUR- RENCE (EO) NUMBERS ASSIGNED BY STATE NATURAL HERITAGE PROGRAMS (INHDC (2012), KNHP (2012), TNHID (2012)), NUMBERS ASSIGNED TO POPULATIONS REPORTED IN SHEA (1993), AND FIRST AND LAST YEARS OF KNOWN OBSERVATIONS

EO Number State County (Shea Popu- First observed Last observed Abundance Land ownership lation Number)

Kentucky ...... Bourbon ...... * 19 (2) 1963–04–27 2005–06–09 10–120 private. Fayette ...... 12 (38) 1931 1931–05–24 n/a private. 16 (37) 1892 1900–05–09 n/a private. Franklin ...... * 2 (6) 1979–05 1992–05–04 11 private. * 3 (8) 1979 1994–05–12 4 private. 5 (39) 1880 1880–06 n/a private. 8 (27) 1981 1981–05–03 ∼40 private. 14 (40) 1856 1856–05 n/a private. * 20 (5) 1992 1992–05–19 21 private. * 21 (7) 1992 1992–05–12 7 private. Jessamine ...... 6 (42) 1942 1942–05–16 n/a private. 13 (32) 1939 1939–04–27 n/a private. 17 (28) 1991–Pre 1991–Pre n/a private. + 27 1990 1993–05–10 1–7 private. Madison ...... 10 (43) 1903 1903–05–16 n/a private. Mercer ...... 24 (44) 1916 1916–05–13 1–7 private. Nelson ...... 25 1935–pre 1935–pre n/a private. Powell ...... 15 (45) 1923 1923–05–26 n/a private. Scott ...... * 9 (15) 1930 1992–05–19 2 private. Tennessee ...... Cheatham ...... 14 (33) 1969–04–29 1969–04–29 n/a private. Davidson ...... * 9 (20) 1974–04–16 1998–04–16 20–29 private; COE easement. + 23 1997–05–09 1997–05–09 ∼200 private.

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EO Number State County (Shea Popu- First observed Last observed Abundance Land ownership lation Number)

Jackson ...... + 25 1998–07–24 1998–07–24 5 COE Maury ...... 7 (31) 1955–04–23 1955–04–23 n/a private. Montgomery ...... 2 (23b) 1968–05–07 1992–04–28 1 private. 13 (30) 1975–05–25 1975–05–25 n/a private. 18 (35) 1967–06–01 1967–06–01 n/a private. 31 1979–04–09 1979–04–09 ...... private. Smith ...... 20 (24) 1992–05–01 1998–04–17 30 private; COE easement. * Occurrences observed by Shea (1993), but which are now considered extirpated. + Occurrences not documented in Shea (1993) that have been observed since 1992, but which are now considered extirpated. COE is the U.S. Army Corps of Engineers. Pres is present.

No records exist in State-maintained various threats, we only use the 55 herbaceous vegetation between a road databases for seven populations that records that have been verified and are and forested bank of a cypress slough Shea (1993, pp. 12–13) treated as currently tracked in State-maintained (M. Homoya, Natural Heritage Program historical or lacking sufficient locality databases. Botanist, Indiana Department of Natural information to verify (population There are 19 extant Short’s Resources (IDNR), December 2012), is number 41 from Kentucky, and numbers bladderpod occurrences that are located unique among populations of the 36 and 46 through 50 from Tennessee). on city, State, or federal lands. The species. The occurrence in Indiana is Therefore, Table 1 and Table 2 do not Indiana occurrence is on lands owned within the Shawnee Hills Section of the include entries for these Shea by the State of Indiana and managed by Interior Low Plateaus Physiographic population numbers. Shea (1993, p. 15) the Indiana Department of Natural Province (Quarterman and Powell 1978, also determined that four historical Resources (IDNR). A portion of one pp. 30–31), on a site underlain by reports for the species were erroneous: occurrence in Kentucky is located in a undifferentiated outwash from the One each from Monroe County, Indiana, State nature preserve owned and Wisconsinan glaciation (Indiana and Vinton County, Ohio; and one each managed by the Kentucky State Nature Geologic Survey 2002) as opposed to the from unknown counties in Kansas and Preserves Commission (KSNPC), and calcareous geology on which the species Vermont. another occurs in a park owned by the occurs in Kentucky and Tennessee. The There are now 8 known extant City of Frankfort, where access is soil at the Indiana site is Weinbach silt occurrences in Kentucky, 17 in limited, but no specific management is loam, which forms in acid alluvium on Tennessee, and 1 in Posey County, provided for the species or its habitat. river terraces, and is nearly level with Indiana (Table 1). Extant occurrences in In Tennessee, there are 15 occurrences 0 to 2 percent slopes (USDA 1979, p. Kentucky are distributed among Clark that are entirely or partially located on 89). This site is on a terrace adjacent to (1), Franklin (6), and Woodford (1) lands owned or leased by the U.S. Army an oxbow swamp formed in an Counties, and in Tennessee among Corps of Engineers (Corps) adjacent to abandoned meander of the Wabash Cheatham (5), Davidson (2), Dickson (1), the Cumberland River. Some of these River (Quarterman and Powell 1978, p. Jackson (2), Montgomery (3), Smith (1), Corps lands are wildlife management 244). and Trousdale (2) Counties. One areas (WMA) cooperatively managed by Kentucky occurrences are located on Tennessee occurrence straddles the the Tennessee Wildlife Resources bluffs and hillsides adjacent to the county line between Cheatham and Agency (TWRA). The plants at EO Kentucky River or its tributaries within Davidson Counties. There are 19 numbers 29 and 32 are located in the Bluegrass Section of the Interior occurrences in Kentucky and 10 in TWRA’s Cheatham WMA, and those at Low Plateaus Province (Fenneman 1938, Tennessee that have either been EO numbers 24 through 27 are located pp. 411–448; Quarterman and Powell extirpated or for which inadequate in TWRA’s Cordell Hull WMA. Part of 1978, pp. 30–31). Extant occurrences in information exists to relocate them. one occurrence in Tennessee is located Kentucky predominantly are found on Adding the seven populations that Shea on lands owned by Ashland City. the Ordovician age Lexington Limestone (1993, p. 12–13) treated as either Habitat. Short’s bladderpod typically and Tanglewood Limestone Members historical or lacking complete locality grows on steep, rocky, wooded slopes (Kentucky Geological Survey, http:// information, and which are not and talus (sloping mass of rock www.arcgis.com/home/item.html?id= represented in State-maintained fragments below a bluff or ledge) areas. d32dc6edbf9245cdbac3fd7e255d3974, databases used to create Tables 1 and 2, It also occurs along tops, bases, and accessed on January 25, 2013), and the these numbers rise to 20 for Kentucky ledges of bluffs. The species usually is Fairmount-Rock outcrop Complex is the and 16 for Tennessee. Thus, there is a found in these habitats near rivers or prevalent soil type at most of the sites total of 62 occurrences that have been streams and on south- to west-facing where the species is found (U.S. reported for Short’s bladderpod. slopes. Most populations are closely Department of Agriculture (USDA), Soil However, when reporting percentages of associated with calcareous outcrops Survey Geographic Database, available all known occurrences that are now or (Shea 1993, p. 16). The Short’s online at http://soildatamart.nrcs.usda. historically were in the case of bladderpod site in Indiana, where the gov, accessed on January 30, 2013). Soils extirpated occurrences, affected by species is found in a narrow strip of of the Fairmount series formed from

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weathered limestone interbedded with Complex (USDA, Soil Survey fruit set in the Indiana population and, thin layers of calcareous shale and are Geographic Database, available online at based on lack of seed production from shallow, well-drained, and slowly http://soildatamart.nrcs.usda.gov, plants in a greenhouse from which permeable. As implied in the name of accessed on January 30, 2013), which pollinators were excluded, she this complex, limestone outcrops are formed from weathered cherty concluded that the species likely is self- common on the steeply sloped sites limestone on sloping to very steep sites incompatible. Self-incompatibility has where this soil occurs, especially along and are somewhat excessively well- been reported in other species of river bluffs (USDA 1985, p. 64). drained. Depth to bedrock within Physaria (Tepedino et al. 2012, p. 142; Tennessee occurrences are located Sulphura soils is less than 16 cm (40 in), Edens-Meier et al. 2011, p. 292; primarily on steep hills or bluffs but deeper in Bodine soils, and chert Claerbout et al. 2007, p. 134; Bateman adjacent to the Cumberland River content is high near the surface of these 1955, p. 64), and the molecular within the Highland Rim and Central soils (USDA 1981, pp. 46–47). mechanisms underlying self-recognition (also known as Nashville) Basin Ordovician age limestones of the between pollen and stigma and Sections of the Interior Low Plateaus Leipers and Cathey Formations, Bigby- subsequent pollen rejection have been Province (Fenneman 1938, pp. 411–448; Cannon Limestone, and Hermitage well studied in the Brassicaceae Quarterman and Powell 1978, pp. 30– Formation are the predominant geologic (Takayama and Isogai 2005, pp. 468– 31). Three occurrences in Cheatham formations underlying occurrences in 474). Dr. Baskin (pers. comm., December County are adjacent to the Harpeth Smith, Trousdale, and Jackson Counties 2012) also observed that seeds produced River near its confluence with the (Wilson et al. 1972, Wilson 1975, by Short’s bladderpod apparently are Cumberland River. Extant occurrences Wilson et al. 1980, Kerrigan and Wilson capable of forming a seed bank, as seeds in Tennessee are found across a wider 2002). In these counties, Short’s that were planted in a greenhouse were range of geology and soils than those in bladderpod occurs across a wider range observed to germinate and produce Indiana or Kentucky. The Mississippian of soil series, all of which are formed seedlings over several years, rather than age Fort Payne Formation, which from weathered limestone or all germinating in the year they were includes limestone and calcareous interbedded siltstone and limestone on planted. siltstone, and Warsaw Limestone are the steeply sloped or hilly sites. The soils The pollinators for Short’s bladderpod predominant geologic formations are shallow, are rocky, or contain areas have not been studied, but Rollins and underlying occurrences in Cheatham, of bedrock outcrop (USDA 2001, pp. 19– Shaw (1973, p. 6) reported that bees and Dickson, and Montgomery Counties 20, 28, 59, 64; USDA 2004a, pp. 22–23, flies were repeatedly observed visiting (Moore et al. 1967, Wilson 1972, Marsh 36–37, 83, 87; USDA 2004b, pp. 21, 75, flowers of other congeners. The majority et al. 1973, Finlayson et al. 1980). In 82). of floral foragers observed visiting Cheatham and Dickson Counties, the Within the physical settings described Physaria filiformis (Missouri main soil mapped in locations where above, the most vigorous (Shea 1992, p. bladderpod) were true bees representing Short’s bladderpod occurs is simply 24) and stable (TDEC 2009, p. 1) Short’s five families, with greater than 50 ‘‘Rock outcrop, very steep’’ (USDA, Soil bladderpod occurrences are found in percent from the family Halictidae. The Survey Geographic Database, available forested sites where the canopy has families Apidae and Andrenidae also online at http://soildatamart.nrcs.usda. remained relatively open over time. were well represented among bee gov, accessed on January 30, 2013). In Common woody species associated with pollinators of this species, the most Montgomery County, Baxter soils and Short’s bladderpod are Acer negundo dependable and frequent of which were Rock outcrop and Bodine cherty silt (box elder), Acer rubrum (red maple), ground-nesters. Several flies of the loam are the soil types on which Short’s Aesculus glabra (Ohio buckeye), Celtis family Syrphidae also carried Missouri bladderpod occurs (USDA, Soil Survey laevigata (hackberry), Cercis canadensis bladderpod pollen (Edens-Meier et al. Geographic Database, available online at (redbud), Fraxinus Americana (white 2011, pp. 293). Tepedino et al. (2012, http://soildatamart.nrcs.usda.gov, ash), Juniperus virginiana (eastern red pp. 143–145) found that native ground- accessed on January 30, 2013). Baxter cedar), Lonicera japonica (Japanese nesting bees from the families soils formed from weathered cherty honey suckle), Parthenocissus Andrenidae and Halictidae were the limestone, and where they are mapped quinquefolia (Virginia creeper), most reliable pollinators visiting flowers as Baxter soils and Rock outcrop they Symphoricarpos orbiculatus (coral of three Physaria species, but they are steeply sloped and Rock outcrop can berry) and Ulmus americana (American reported fewer numbers of pollen- make up as much as 20 percent of the elm). Common herbaceous associates carrying flies from the families map unit (USDA 1975, pp. 12–14). include Alliaria petiolata (garlic Tachinidae and Conopidae. They Bodine soils are well-drained, cherty mustard), Camassia scilloides (wild estimated that maximum flight distance soils that formed from weathered cherty hyacinth), Chaerophyllum procumbens ranged from 100 to 1400 meters (m) (330 limestone; are steeply sloped; and (spreading chervil), Delphinium tricorne to 4593 feet (ft)) for the Andrenids and include areas near the escarpment (dwarf larkspur), Galium aparine 40 to 100 m (130 to 330 ft) for the adjacent to the Cumberland River (cleavers), Lamium sp. (dead nettle), Halictid bees they collected. floodplain where cherty limestone Phacelia bipinnatifida (forest phacelia), Whorled Sunflower bedrock is exposed (USDA 1975, pp. Polygonatum biflorum (Solomon’s seal), 16–17). Sedum pulchellum (stonecrop), Silene Helianthus verticillatus is a member Silurian age limestone and shale of virginica (fire-pink), and Verbascum of the sunflower family known from the Waynes Group and the Brassfield thapsus (common mullein) (Shea 1993, Cherokee County, Alabama; Floyd Limestone and Ordovician age p. 19). County, Georgia; and McNairy and limestone of the Leipers and Catheys Biology. Published literature on the Madison Counties, Tennessee. It is a Formations are the predominant biology of Short’s bladderpod is lacking. perennial arising from horizontal, geologic formations underlying the The species flowers during April and tuberous-thickened roots with slender occurrences located in Davidson County May (Gleason and Chronquist 1991, p. rhizomes. The stems are slender, erect, (Wilson 1979). The dominant soils on 187, Shea 1993, p. 20). Dr. Carol Baskin and up to 2 meters (m) (6 feet (ft)) tall. which Short’s bladderpod occurs in this (Professor, University of Kentucky, pers. The leaves are opposite on the lower county are the Bodine-Sulphura comm., December 2012) observed low stem, verticillate (whorled) in groups of

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3 to 4 at the mid-stem, and alternate or Chester County, Tennessee, in 1892. studies showed through comparative opposite in the inflorescence at the end. Small distinguished it from the related genetic studies with its putative parents, Individual leaves are firm in texture and H. giganteus by its smooth and hairless H. grosseserratus and H. angustifolius, have a prominent mid-vein, but lack stems; narrow, entire leaf blades; and that whorled sunflower is a good prominent lateral veins found in many narrowly linear-lanceolate involucre (a taxonomic species of non-hybrid origin members of the genus. The leaves are collection or rosette of bracts subtending (Ellis et al. 2006, pp. 2351–2352). linear-lanceolate in shape, narrowing at a flower cluster, umbel, or the like) Distribution and Status. There are the tip to a point, and 7.5 to 18.5 cm (3.0 bracts (a leaflike or scalelike plant part, four whorled sunflower populations to 7.2 in.) long and 0.7 to 2.0 cm (0.3 usually small, sometimes showy or known to be extant, each consisting of to 0.8 in.) wide. The flowers are brightly colored, and located just below multiple tracked subpopulations (Table arranged in a branched inflorescence a flower, a flower stalk, or an 3) (Alabama Natural Heritage Program typically consisting of 3 to 7 heads. The inflorescence). No additional collections (ANHP) 2012, Georgia Department of heads are about 1 cm high (0.4 in.), are of this species had been made when about 1.5 cm (0.6 in.) wide, and have Beatley (1963, p. 153) speculated that Natural Resources (GDNR), TNHID deep yellow ray flowers and lighter the specimens (which lacked basal parts 2012). In Floyd County, Georgia, there is yellow disk flowers. The seeds are 0.4 and mature seeds) from this single one population comprised of four to 0.5 cm (0.16 to 0.2 in.) long. collection site perhaps represented a subpopulations. There is one population Several members of the aster family single aberrant individual formed from in Cherokee County, Alabama, are similar in appearance to whorled hybridization of an opposite- and comprised of two subpopulations. sunflower, with minor morphological alternate-leaved Helianthus species. Populations in Georgia and Alabama are differences being apparent. Helianthus With no new material to examine, less than 2 km (1.2 mi) apart. In grosseserratus is similar to whorled Heiser et al. (1969, p. 209) and Tennessee, there is one population sunflower but its leaves typically are Cronquist (1980, p. 36) accepted comprised of six subpopulations in arranged in an alternating pattern, Beatley’s suggestion that whorled McNairy County and one population which differs from the whorled sunflower was a hybrid. comprised of four subpopulations in arrangement of H. verticillatus. The rediscovery of the species in Madison County. Table 3 lists these Helianthus angustifolius can be 1994, in Georgia, provided ample populations and subpopulations, and confused with H. verticillatus but it has material for reexamination of this relates them to EO numbers used by narrower leaves and reddish disk species’ taxonomic status. Plants State conservation agencies to track flowers, as opposed to the yellow disk throughout these new populations were their status. The population in Floyd flowers of H. verticillatus (Schotz 2001, found to conform to the morphology of County, Georgia, is located on lands p. 1). Helianthus giganteus often the type collection of whorled owned by The Campbell Group, a exhibits whorled leaves, but H. sunflower. Morphological studies and timber investment management verticillatus leaves have only the root-tip chromosome counts by organization. This site is referred to as midvein prominent while H. giganteus Matthews et al. (2002, pp. 17–23) the Coosa Valley Prairie and is protected has lateral veins evident on the leaves validated this taxon’s status as a by a conservation easement held by The (Matthews et al. 2002, p. 22). distinct, diploid species. The taxonomic Nature Conservancy, which jointly Taxonomy. Whorled sunflower was validity of this species was also manages the property with The described by J.K. Small (1898, p. 479), confirmed through genetic studies by Campbell Group. All other sites also are based on a collection by S.M. Bain from Ellis et al. (2006, pp. 2345–2355). Their on private lands but are not protected.

TABLE 3—LIST OF WHORLED SUNFLOWER POPULATIONS AND SUBPOPULATIONS BY STATE AND COUNTY, WITH COR- RESPONDING SITE NAMES AND ELEMENT OCCURRENCE (EO) NUMBERS FROM STATE CONSERVATION AGENCY DATA- BASES IN ALABAMA, GEORGIA, AND TENNESSEE

Heritage Population Subpopulation Site name EO (County, State) number(s) Number

Cherokee, AL ...... 1 Kanady Creek Prairie ...... AL_1 2 Locust Branch Prairie ...... AL_2 Floyd, GA ...... 1 Jefferson Road Wet Prairie ...... GA_1 2 Kanady Creek Wet Prairie ...... GA_4 3 Upper Mud Creek Wet Prairies ...... GA_5 4 Sunnybell Prairie...... GA_7 Madison, TN ...... 1–6 Turk Creek ...... TN_2 McNairy, TN ...... 1–4 Prairie Branch ...... TN_3

Status surveys have been conducted percent. Surveys during 2000 and 2002 2002, pp. 3–8; Schotz 2001, pp. 8–10); for this species throughout its range in Tennessee were unsuccessful at however, due to the species’ clonal (Nordman 1998, pp. 1–17; 1999, pp. 1– locating any additional sites (Lincicome growth habit, stem counts overestimate 5; Schotz 2001, pp. 1–14; Allison 2002, 2003, pp. 1–2). Surveys in 2006 resulted the true number of genetically distinct pp. 1–2; Lincicome 2003, pp. 1–2). in discovery of the population in individuals (genets). Ellis et al. (2006, p. Despite these extensive surveys, the McNairy County, Tennessee (Tennessee 2349) found that the genetic population number of known populations remains Division of Natural Areas 2008, p. 2). size is much smaller than the number of low. Schotz (2001, pp. 1, 10) located 1 Initial efforts to estimate population stems in a population and that a more new population out of 44 attempts, sizes of whorled sunflower relied on accurate population census could be representing a success rate of only 2 counting individual stems (Allison made at most whorled sunflower sites

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by counting obvious clusters of stems Botanist, Georgia Department of Natural The Alabama and Georgia populations rather than individual stems. However, Resources, pers. comm. February 2012). are located on flat to gently rolling Mandel (2010, p. 2056) reported that Based on the work of Ellis (2006) and uplands and along stream terraces in the individual clusters were much less Mandel (2010), summarized above, at headwaters of Mud Creek, a tributary to distinct in a portion of the Alabama site one time Alabama supported the largest the Coosa River. In Tennessee, the she sampled. population with an estimated 100 Madison County population occurs Ellis et al. (2006, p. 2351) counted 70 individuals at the Kanady Creek Prairie along Turk Creek, a tributary to the distinct clusters at the site in Madison, site, where whorled sunflower was first South Fork Forked Deer River, and in Tennessee, which closely equated to 70 found to occur in the State. However, adjacent uplands. The McNairy County separate individuals through genetic Schotz (2011, p. 11) found only 79 population occurs along Prairie Branch, analyses; however, not all clusters were stems, distributed among 8 clusters, at a headwater tributary to Muddy Creek sampled at this site (Mandel, pers. this site in 2011. Mandel (2010) in the Tuscumbia River drainage. comm., 2012). At the McNairy County, sampled only portions of the Georgia We used the Natural Resources Tennessee, population, 36 clusters of and Tennessee populations, thus Conservation Service’s Web Soil Survey plants were found growing along creek underestimating their sizes. Whorled to determine the soil types on which banks at the unplowed edges of sunflower likely is now most abundant whorled sunflower populations occur cultivated crop fields and extending in Georgia due to population growth in across its range (USDA, Web Soil into a railroad right-of-way (Tennessee response to habitat management by The Survey, available online at http:// Division of Natural Areas 2008, p. 3). Nature Conservancy and The Campbell websoilsurvey.nrcs.usda.gov/app/ Mandel (2010, p. 2056) sampled 19 Group at the Coosa Valley Prairie. HomePage.htm, accessed on January 30, clusters at the McNairy County Schotz estimated approximately 175 to 2013). The most prevalent soils where population and determined these 200 stems were present at the second the species occurs in Georgia are represented 24 genets; however, only Alabama site in September 2008 (Schotz Conasauga, Lyerly, Townley, and two of the four subpopulations mapped pers. comm. 2009), but there were only Wolftever silt loams and Dowellton silty at this population were sampled 42 stems found at this site in 2011 clay loam. The silt loam soils all formed (Mandel, pers. comm., 2012). (Schotz 2011, p. 14). No estimate of from weathered limestone or shale, and individual plants is available for this occupy various land forms from broad Mandel (2010, p. 2058) sampled the site. upland ridges to low stream terraces. Alabama subpopulation number 1 Habitat. Whorled sunflower is found These soils share the characteristics of (Table 3) using two methods. In one in moist, prairie-like remnants, which in being moderately well-drained to well- portion of the site, leaf tissue was a more natural condition exist as drained, being slightly to extremely collected from 15 distinct clusters, openings in woodlands and adjacent to acid, and having low to moderate which represented 24 genets. However, creeks. Today, the only whorled fertility and organic matter content and because distinct clusters were not sunflower site where these habitat clayey subsoils (USDA 1978a, pp. 24– obvious in another portion of this conditions are present over a relatively 54). The Dowellton silty clay loam subpopulation, Mandel (2010, p. 2058) large area is located in the Coosa Valley formed in alluvium (soil material sampled leaves from the first 100 stalks Prairie of northwest Georgia, where the deposited by running water) on low encountered in a 1-meter-wide transect species occurs in prairie openings and stream terraces and upland depressions run through the largest patch of whorled woodlands interspersed among lands is poorly drained, is moderate in sunflower in that area. These 100 stalks managed for pulpwood and timber fertility and organic content, is neutral were within an approximately 11-m (40- production. At one of the Alabama to strongly acid, and has a clayey ft) long portion of this transect, and subpopulations, whorled sunflower subsoil (USDA 1978a, pp. 28–29). represented 46 distinct genets. Mandel occurs in a narrow, open strip of Alabama subpopulations inhabit the (2010, p. 58) estimated that 400 stalks vegetation between a roadside and Gaylesville silty clay loam, a deep, were present in this area and that the adjacent forest. The second Alabama poorly drained, slowly permeable soil total number of genets was between 100 subpopulation occurs along a small formed from limestone on floodplains and 200. However, more recently only intermittent stream and adjacent and depressed areas in limestone 79 stems, distributed among 8 clusters, floodplain, in a site where an immature valleys (USDA 1978b, p. 20). These soils were found at this site (Alabama Natural hardwood forest was harvested in 1998. are strongly to extremely acid, with low Heritage Program 2011, p. 11). Whorled sunflower and associated natural fertility and medium organic Mandel (2010, p. 2056) sampled 15 prairie species responded favorably to content (USDA 1978b, p. 20). Conasauga clusters growing in a ‘‘wet prairie’’ at the timber removal, but the site was silt loams, discussed above, lay upslope the Georgia site, presumably soon converted into a loblolly pine of the Gaylesville soils at the Alabama representing EO number 1 from the plantation and the planted seedlings whorled sunflower sites. Georgia Natural Heritage Program have grown into a young, dense stand In Madison County, Tennessee, the database (Table 3). It was determined into which little light penetrates. As of population is primarily found on Falaya that these clusters represented 18 genets 2012, there were few whorled sunflower silt loam, which are poorly drained soils (Mandel 2010, p. 2058), but apparently plants or prairie associates present at that formed in alluvium derived from the other three subpopulations present this site. Known populations of this loess (loamy soil material believed to be at this population were not sampled. species in Tennessee are relegated deposited by wind) and are strongly to The true number of genets at this site is mostly to narrow bands of habitat very strongly acid (USDA 1978, p. 44). likely much greater, as others have between cultivated fields and creeks The McNairy County, Tennessee, reported vigorous growth of whorled and adjacent to roads and railroad population occurs on Iuka and Enville sunflower in response to prescribed rights-of-way. The largest concentration fine sandy loam soils, both of which fires that are used to manage the Coosa of plants in Tennessee is found at the occupy floodplains and are occasionally Valley Prairie conservation easement Madison County population, in a 1-ha flooded during winter and early spring area (M. Hodges, Georgia Director of (2.5-ac) patch of remnant, wet prairie (USDA 1997, pp. 73–76). Stewardship, The Nature Conservancy, habitat wedged between US Highway 45 The list of associated species in these pers. comm. May 2012; T. Patrick, and a railroad right-of-way. habitats indicates a community with

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strong prairie affinities. Dominant creating dense colonies. The stems can than all other groups and germination grasses of the tall grass prairie are reach over 4 m (13 ft) in height rates were lower than both Alabama present including Schizachyrium (Matthews et al. 2002, pp. 17–20). intra-population crosses and Alabama scoparium (little bluestem), Ellis and McCauley (2008, p. 1837) plants crossed with pollen from Sorghastrum nutans (Indian grass), investigated whether there were Tennessee plants (Ellis and McCauley Andropogon gerardii (big bluestem), and differences among populations of 2008, pp. 1839–1840). Panicum virgatum (switch grass). Other whorled sunflower with respect to Based on their results, Ellis and common herbaceous associates include achene viability and germination rates McCauley (2008, p. 1841) concluded Bidens bipinnata (Spanish needles), and whether those differences might that populations of whorled sunflower Carex cherokeensis (Cherokee sedge), have a genetic basis. They conducted are not interchangeable with respect to Hypericum sphaerocarpum (roundseed this experiment for two generations of phenotypic fitness-related characters St. Johnswort), Helianthus angustifolius plants, the second generation produced (i.e., achene viability and germination (swamp sunflower), Helenium from intra-population crosses of first rates) and suggested that the potential autumnale (common sneezeweed), generation plants. They also explored exists for genetic rescue of the Lobelia cardinalis (cardinal flower), whether isolation of populations from Tennessee population by transplanting Pycnanthemum virginianum (Virginia one another could have fitness either seeds or seedlings produced from mountain mint), Physostegia virginiana consequences, by conducting inter- crosses between Tennessee and (obedient plant), Saccharum giganteum population crosses and evaluating Alabama plants into the Tennessee (sugarcane plumegrass), Silphium whether they found: (1) Evidence of population. genetic rescue expressed as higher terebinthinaceum (prairie rosinweed), Fleshy-fruit Gladecress Sporobolus heterolepis (prairie fitness of hybrid individuals as dropseed), and Symphyotrichum novae- compared to any or all of the parental Leavenworthia crassa is a glabrous angliae (New England aster) (Tennessee populations; and (2) evidence of (morphological feature is smooth, Division of Natural Areas 2008, p. 5; outbreeding depression. Their study glossy, having no trichomes (bristles or Matthews et al. 2002, p. 23; Schotz included material from the Alabama, hair-like structures)) winter annual 2001, p. 3). Some of these areas are also Georgia, and Madison County, known from Lawrence and Morgan habitat for a number of other rare Tennessee, populations. However, they Counties, Alabama. It usually grows species including Marshallia mohrii were unsuccessful in cultivating plants from 10 to 30 cm (4 to 12 in) tall. The (Mohr’s Barbara’s buttons), which is from the Georgia population, where the leaves are mostly basal, forming a federally listed as threatened. flower heads contained few viable rosette, and entire to very deeply, Biology. There is little published achenes, which produced low pinnately (multiple leaflets attached in information available concerning the germination rates (Ellis and McCauley rows along a central stem) lobed or biology of the whorled sunflower, and 2008, pp. 1837–1838). divided, to 8 cm (3.1 in) long. Flowers the cause for its current rarity is not The number of crosses that produced are on elongating stems, and the petals known. Ellis et al. (2006, pp. 2349– no viable achenes was higher in the are approximately 0.8 to 1.5 cm (0.3 to 2350) investigated genetic diversity in intra-population Tennessee crosses than 0.6 in.) long, obovate to spatulate, and the Georgia, Alabama, and Madison in any other pair of crossings. Those emarginate (notched at the tip). Flower County, Tennessee, populations of achenes that were produced by first color is either yellow with orange or whorled sunflower and found high generation Tennessee intra-population white with yellow, usually with both levels of genetic diversity at the crosses exhibited lower germination color forms intermixed in a single population and species levels despite its rates than Alabama achenes, and second population. The fruit is globe-shaped or apparent rarity. They speculated that generation Tennessee achenes from slightly more elongate and about 1.2 cm this is indicative of a species that was intra-population crosses exhibited both (0.5 in) long with a slender beak at the more widespread in the past and lower viability and germination rates tip, which is 0.25 to 0.60 cm (0.1 to 0.24 perhaps became rare relatively recently than the Alabama achenes. However, in) in length. Seeds are dark brown, (Ellis et al. 2006, pp. 2351–2352). survival rates of germinated achenes did nearly round in shape and winged. Whorled sunflower populations not differ among these populations in Taxonomy. Fleshy-fruit gladecress exhibited moderate levels of either generation (Ellis and McCauley was described by Rollins in 1963, from differentiation based on markers that are 2008, p. 1840). Ellis and McCauley material collected in 1959, from Morgan presumed to be selectively neutral, and (2008, p. 1840) suggested three possible County, Alabama. Rollins (1963, pp. 61– since these populations are mechanisms that could explain these 68) delineated the species into two geographically distinct and ecological results, none of which are mutually varieties (var. crassa and var. elongata) conditions vary somewhat among them exclusive: (1) Limited mate availability based on differences in fruit length. Ellis et al. (2006, p. 2353) concluded in the Tennessee population due to However, herbarium and field studies that they likely are as differentiated, if limited diversity of self-incompatibility have shown var. elongata to have not more so, at adaptive loci (the alleles; (2) more extensive inbreeding variation in fruit length within the range specific location of a gene or DNA within the Tennessee population; or (3) of fruit lengths for var. crassa (McDaniel sequence on a chromosome). differential adaptation between the two and Lyons 1987, p. 2–3). Thus, the Whorled sunflower is a self- populations. species is treated as one taxon incompatible, clonal perennial and When Tennessee plants were crossed throughout this document. This taxon flowers from August into October with pollen from Alabama plants, the was brought to the attention of the (Matthews et al. 2002, pp. 17–20; Ellis second generation mean achene scientific community in 1957, by and McCauley 2008, p. 1837). The viability and germination rates were venerable botanist Reed C. Rollins, who species is easily cultivated and seed equal to or greater than those of distinguished the taxon from similar germination is high in the laboratory. Alabama intra-population crosses or species based on reproductive Upon transplanting, this species has Alabama plants crossed with pollen morphology. been shown to reproduce rapidly from from Tennessee plants. Mean achene Fleshy-fruit gladecress’s globular to rhizomes (a horizontal underground viability of Tennessee intra-population oblong fruit with a smooth exterior stem that produces roots and shoots), second generation crosses was lower distinguishes it from another gladecress

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species, Leavenworthia alabamica p. 6). Surveys by Lyons (in litt. 1981 to Lawrence Counties, Alabama (Table 4). (Alabama gladecress), which has a much R. Sutter), McDaniel and Lyons (1987, p. One of these occurs on U.S. Forest more elongated linear fruit with 5–6), and Hilton (1997, p. 12) were Service (USFS) lands, where it is corrugated surfaces. Alabama gladecress unsuccessful at locating a number of formally protected. The majority of also does not usually have the yellow historical sites for fleshy-fruit other sites are actively grazed, a practice and orange flower forms found mixed in gladecress. McDaniel and Lyons (1987) that has, for the most part, maintained populations of fleshy-fruit gladecress failed to locate eight sites previously favorable growing conditions for the (McDaniel and Lyons 1987, p. 10). reported by Rollins (1963, p. 63), and species. However, adjusting grazing Distribution and Status. Fleshy-fruit Lloyd (1965) and Hilton (1997, p.12) patterns to take place during the gladecress is endemic to a 21-km (13- were unsuccessful at locating seven species’ dormant cycle would greatly mi) radius area in north central Alabama sites listed in McDaniel and Lyons reduce potential mortality of in Lawrence and Morgan Counties (1987, p. 5–6). (Rollins 1963, p. 63). A 1961 record Currently there are six known extant reproducing plants while maintaining from Lauderdale County has never been occurrences of fleshy-fruit gladecress ideal habitat conditions. confirmed (McDaniel and Lyons 1987, documented, three each in Morgan and

TABLE 4—LOCATION, SITE NAMES AND DESCRIPTIONS, AND ELEMENT OCCURRENCE (EO) RANKS FOR KNOWN EXTANT FLESHY-FRUIT GLADECRESS OCCURRENCES

EO County Population designation Rank Historic site description Land ownership

Lawrence ...... Bluebird Glades ...... D ..... Described by ALNHP in 1995 as approx. Private & State ROW. 0.2-ha (0.5-ac) site with 1200 plants; by 2009 was reduced to 600 plants. Stover Branch Glades ...... C ..... Two subpopulations, most in pasture, 3.16 Private. ha (7.8 ac); 2,200 to 2,500 plants; maintained by livestock management, found in 1961. Indian Tomb Hollow Glade ...... A ..... 0.46-ha (1.1-ac) site with 1,200 to 1,300 Federal—USFS. plants; discovered 1977. Morgan ...... Cedar Plains South ...... C ..... 0.04-ha (0.1-ac) site with 75 to 100 plants; Private. discovered 1968. Cedar Plains North ...... B ..... 1.7-ha (4.2-ac) site with 5,000 to 6,000 Private. plants; discovered 1968. Massey Glade ...... C ..... 2.75-ha (6.8-ac) site with 2,300 to 2,500 Private. plants; discovered 1961. ALNHP is the Alabama Natural Heritage Program. ROW is right-of-way.

The Alabama Natural Heritage Habitat. This species is a component moisture, and the presence or absence of Program determines EO ranks ranging of glade flora and occurs in association a woody vegetation component. Few from A to D for sites and populations of with limestone outcroppings. The terms undisturbed examples of this rare species, with A indicating the ‘‘glade’’ and ‘‘cedar glades’’ are used community type remain (Hilton 1997, status of the EO is considered to be interchangeably to refer to shallow- pp. 5, 8; McDaniel and Lyons 1987, p. excellent, B good, C marginal, and D soiled, open areas that are dominated by 11; Baskin and Baskin 1985, p. 1; poor. The EO rank is based on a herbaceous plants and characterized by Rollins 1963, p. 5–6). combination of standardized criteria exposed sheets of limestone or gravel. Populations of fleshy-fruit gladecress including quality, condition, viability, Eastern red cedar (Juniperus virginiana) are now located in glade-like remnants and defensibility. Hilton (1997, pp. 13– trees are frequent in the deeper soils exhibiting various degrees of 26) developed the specific criteria for along the edges of the glades (Hilton disturbance, including pastures, determining EO ranks for fleshy-fruit 1997, p. 1; Baskin et al. 1986, p. 138; roadside rights-of-way, and cultivated or gladecress and its habitat. Based on Baskin and Baskin 1985, p. 1). Glades plowed fields (Hilton 1997, p. 5). As these criteria, only one of the six can vary in size from as small as a few with most of the cedar glade endemics, occurrences is A-ranked. It consists of square meters to larger than 1 square fleshy-fruit gladecress exhibits weedy an estimated 1200+ plants in a relatively kilometer (km2) (0.37 square miles tendencies, and it is not uncommon to undisturbed glade (Schotz 2009, p. 10). (mi2)) and are characterized as having find the species growing in altered Of the remaining occurrences, one has an open, sunny aspect (lacking canopy) habitats. However, none of the cedar approximately 5,000 to 6,000 plants, but (Quarterman 1950, p. 1; Rollins 1963, p. glade endemics appear to have spread is B-ranked because the site where it is 5). Historically, glades in northern very far from their original glade located is heavily grazed. Three Alabama occurred as glade complexes habitats; thus the geographic range of occurrences are C-ranked (2 occurrences where sparsely vegetated patches of fleshy fruit gladecress is probably very have approximately 2400 plants in a exposed, or nearly exposed, limestone similar to what it was in pre-settlement degraded glade community; the other occurred in a matrix of woody times (Baskin et al. 1986, p. 140). occurrence has 75 to 100 plants but is vegetation to form a mosaic of habitats All species within the small genus located in high-quality habitat), and one grading into one another (Hilton 1997, Leavenworthia are adapted to the is D-ranked (600 plants in a residential pp. 1, 5, 64). Herbaceous diversity was unique physical characteristics of glade area with no potential for habitat irregular over these complexes, affected habitats, perhaps the most important of restoration) (Schotz 2009). by changes in soil gradient and these being a combination of shallow

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depth and high calcium content of soils CFR part 424, set forth the procedures observed on a steep slope above the and their tendency to have temporarily for adding species to the Federal Lists road cut adjacent to Kentucky EO 7 in high moisture content at or very near of Endangered and Threatened Wildlife 2004 (KNHP 2012), no plants were the surface (Rollins 1963, pp. 4–6). and Plants. Under section 4(a)(1) of the found at the base of the bluff, where 21 Typically, only a few inches of soil Act, we may list a species based on any plants had been observed in 1992 (Shea overlie the bedrock, or, in spots, the soil of the following five factors: (A) The 1993, p. 60) before the road cut had may be almost lacking and the surface present or threatened destruction, altered the habitat. Poorly timed barren. The glade habitats that support modification, or curtailment of its mowing or indiscriminate herbicide all Leavenworthia species are extremely habitat or range; (B) overutilization for application along the road cut at the wet during the late winter and early commercial, recreational, scientific, or base of this bluff could cause mortality spring, and become extremely dry in educational purposes; (C) disease or of seedlings produced there from seeds summer (Rollins 1963, p. 5). predation; (D) the inadequacy of that are dispersed from the plants on the In northern Alabama, cedar glades existing regulatory mechanisms; and (E) slope above. According to data from the primarily are distributed within the other natural or manmade factors KNHP (2012), a road cut was present in Moulton Valley subdivision of the affecting its continued existence. Listing 2004, and no Short’s bladderpod could Interior Low Plateau Physiographic actions may be warranted based on any be found at Kentucky EO 2, where in Province, and a few glades are scattered of the above threat factors, singly or in 1992 Shea (1993, p. 52) observed 11 up the Eastern Valley subdivision of the combination. Each of these factors is Short’s bladderpod plants and observed Tennessee Valley (Hilton 1997, p. 1). discussed below. no apparent threats to the population. Most of these glades are concentrated in Much of the habitat downslope of a the Moulton Valley, a level area Short’s Bladderpod road, where Tennessee EO 20 once underlain by Mississippian age A. The Present or Threatened occurred but is no longer extant, was limestone stretching across Morgan, Destruction, Modification, or found to be covered with rip rap in Lawrence, Franklin, and Colbert Curtailment of Its Habitat or Range 2008, and the remaining habitat above Counties in northwestern Alabama. and below the road was overgrown Glades occur in association with Shea (1993, pp. 22–23 and 42–92) and (TDEC 2009, p. 10). Road construction outcrops of Bangor Limestone and Tennessee Department of Environment destroyed suitable habitat around typically are level with exposed sheets and Conservation (2009, p. 1–3) Tennessee EO 23, and Short’s of limestone or limestone gravel discussed several threats that have bladderpod is no longer present at the interspersed with fingers of cedar- destroyed or modified Short’s site (TNHID 2012). Based on these data, hardwood vegetation. The Bangor bladderpod habitat and could cause five Short’s bladderpod occurrences (9 Limestone underlying the Moulton further habitat loss or modification in percent) have been lost to habitat Valley tapers to an end in eastern the future. These include transportation destruction or modification associated Morgan County, where it meets the right-of-way construction and with road construction or maintenance. sandstone of Brindley Mountain. maintenance; impoundments and Shea (1993, p. 22) identified roadside Limestone is often near the soil surface, reservoir water level manipulation; maintenance as a threat to 12 and can be seen in rocky cultivated overstory shading due to forest occurrences, including two discussed fields and as small outcroppings at the succession; competition and shading above: Indiana EO 1; Kentucky EOs 1 base of low-lying forested hills (Hilton from invasive, nonnative plant species; through 4, 7, 19, and 23; and Tennessee 1997). trash dumping; commercial and EOs 2, 4, 10, and 22. In addition, Biology. Fleshy-fruit gladecress is an residential construction; and livestock Kentucky EO 27 is located along a annual, spring-flowering member of the grazing. Predictions of increased mowed roadside (KNHP 2012), and mustard family (Brassicaceae). As an frequency, duration, and intensity of TDEC (2009, p. 2) reported that annual, the seeds germinate in the fall, droughts across the species’ range, and Tennessee EOs 3 and 15 could be overwinter as rosettes, and commence a increased flooding in the Midwest affected by roadside maintenance. month-long flowering period beginning region, could portend adverse effects for Indiana EO 1 is an extant roadside in mid-March. The first seeds mature in Short’s bladderpod and its habitat. We occurrence, where the species’ late April, and during most years the discuss each of these threats in greater persistence depends on periodic plants dry and drop all of their seeds by detail below. clearing of competing vegetation and the end of May. It is unlikely that all Transportation Right-of-Way associated soil disturbance to prevent seeds produced in spring germinate the Construction and Maintenance succession of the vegetation at the site next fall, but the length of dormancy in to a forested condition that would be the soil is not known (McDaniel and During the status survey for this unsuitable for Short’s bladderpod Lyons 1987, p. 10); thus we do not know species, Shea (1993, p. 22) observed that (Homoya, pers. comm., December 2012). whether the species is capable of Short’s bladderpod habitat at three sites Nonetheless, poorly timed mowing or forming a seed bank. Native bees in the (Kentucky EO 7; Tennessee EOs 7, 14) indiscriminate herbicide application families’ Andrenidae and Halictidae had been destroyed or degraded by road could negatively affect this occurrence (sweat bees), including the species construction or maintenance activities. by disrupting reproductive cycles or Halictus ligatus (sweat bee), were Neither of these Tennessee occurrences causing direct mortality of Short’s observed carrying pollen from is extant today (TNHID 2012). Shea bladderpod plants. In total, roadside Leavenworthia crassa (fleshy-fruit (1993, p. 60) observed 48 plants at maintenance has been identified as a gladecress) and L. alabamica (Alabama Kentucky EO 7 in 1992, but noted that threat to 15 occurrences. gladecress) in northern Alabama (Lloyd the population had been much more Short’s bladderpod is considered 1965). extensive prior to improvements of U.S. extirpated from four of the eight sites in 421. Shea (1993, p. 22) also indicated Kentucky where roadside maintenance Summary of Factors Affecting the that roadside maintenance posed a has been identified as a threat to the Species continuing threat to the species at this species. Neither Kentucky EO 2, lost to Section 4 of the Act (16 U.S.C. 1533), location. Although approximately 100 road construction as discussed above, and its implementing regulations at 50 Short’s bladderpod plants were nor EO 3 is extant. No plants were

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found at Kentucky EO 3 during searches absent during a search in 1998, and no Nonetheless, the potential exists for in 2004 and 2008; however, only a few plants were found during a monitoring road widening projects or vegetation plants had been observed here in 1994 visit in 2008. As noted above, Short’s management efforts along road rights-of- and earlier (KNHP 2012), and the cause bladderpod was no longer present when way to destroy or modify habitat, cause for the species’ current absence is not TDEC (2009, p. 10) observed in 2008 mortality of individual plants, or known. Despite the presence of 17 that the roadside habitat at Tennessee diminish reproductive output at a large Short’s bladderpod plants at Kentucky EO 20 had been covered with rip rap proportion of sites where the species EO 19 during 2005, none were found and the remaining habitat above and occurs. during visits in 2004 and 2011 (KNHP below the road was overgrown. There are seven extant Short’s 2012). While roadside maintenance Short’s bladderpod remains extant at bladderpod occurrences, and three sites could have contributed to loss of this five of the seven sites in Tennessee from which the species is thought to be population, observations by Kentucky where roadside maintenance has been extirpated, located in or adjacent to the Natural Heritage Program (2012) identified as a threat to the species. Old Tennessee Central Railroad right-of- indicate that shading or competition More than 500 Short’s bladderpod way (TDEC 2009, p. 3, TNHID 2012), from invasive species is likely a primary plants were found at Tennessee EO 3 in portions of which are not actively used cause. Short’s bladderpod was last seen 2008 (TDEC 2009, p. 6), where Shea or maintained or have been sold to other at Kentucky EO 27 in 1993, when seven (1993, p. 89) found 40 plants in 1992. rail companies. There were hundreds to plants were found along a mowed This occurrence is located along a thousands of Short’s bladderpod plants roadside dominated by fescue and other south-facing wooded slope, north of the each at three of these occurrences weeds (KNHP 2012). This occurrence Cumberland River, but very little of its (Tennessee EOs 1, 10, and 17) when was determined to be extirpated during habitat would be vulnerable to TDEC (2009, p. 4) monitored the species a 2011 site visit by KNHP (2012) staff. maintenance associated with the road in 2008. The Nashville Area Short’s bladderpod remains extant at right-of-way to the immediate west. Metropolitan Planning Organization four of the eight sites in Kentucky where Tennessee EOs 4 and 10 are located (NAMPO) (2010, p. 98) 2035 Regional roadside maintenance has been along a roadside approximately 0.5 km Transportation Plan reported that the identified as a threat to the species. (0.3 mi) apart, and both occurrences are Old Tennessee Central Railroad, which Kentucky EO 1 is considered extant, but estimated to number in the hundreds to follows the Cumberland River and only three Short’s bladderpod plants— thousands of plants (TDEC 2009, p. 6– passes through Ashland City, was found two in 1992, and one in 2009—have 8). While roadside maintenance could to be the most practical alignment for a been observed at this site since the adversely affect plants located along the proposed commuter rail to improve species was first observed there in 1975. base of the roadside bluffs on which intercity commute options between the Kentucky EO 4 was treated as two they occur, the majorities of these cities of Nashville and Clarksville, separate populations by Shea (1993, pp. occurrences are located on ledges and Tennessee. While no plans have been 62–65), which are now tracked as a bluff tops where roadside maintenance produced for developing this proposed single occurrence (KNHP 2012). While would be unlikely to affect them. commuter rail system, the 2035 some plants at the base of the cliff Tennessee EO 15 is a small occurrence Regional Transportation Plan states that where Kentucky EO 4 is located are located adjacent to a bridge, on a steep this transportation option should be vulnerable to roadside mowing or limestone bluff overlooking the Harpeth developed by 2017 (NAMPO 2010, p. herbicide application, many of the River. While no impacts from roadside 98). Habitat modification or destruction plants are on the cliff face and maintenance have been observed, no resulting from such development could associated ledges, and no impacts from more than 20 plants have ever been potentially affect 27 percent of the roadside maintenance have been counted at this occurrence. Biologists known extant occurrences of the documented. Short’s bladderpod from TDEC (2009, p. 11) found species, including some occurrences abundance at this occurrence has ranged approximately 35 plants at Tennessee where the species is most abundant. from a low of approximately 56 EO 22, where Shea (1993, p. 85) found Flooding and Water Level Fluctuation individuals in 1998, to a high of at least 43 reproductive plants in 1992. No 400 individuals in 2004 (KNHP 2012). impacts from roadside maintenance Shea (1993, pp. 22–23) and TDEC As discussed above, there were were noted during this site visit. (2009, p. 2) noted that impoundments approximately 100 plants observed Four Short’s bladderpod occurrences and artificial water level manipulation above the road cut at Kentucky EO 7, (7 percent) apparently have been lost to threatened several Short’s bladderpod but roadside maintenance could prevent road construction or roadside occurrences. This threat might be better plants from becoming established at the maintenance. While 10 of the known characterized as flooding and water base of the road cut. Kentucky EO 23 extant occurrences (38 percent) are level fluctuation, regardless of cause, as has ranged in abundance from a low of located along roadsides, where some occurrences in free-flowing river 60 plants in 2008, to a high of at least maintenance activities such as mowing reaches are vulnerable to this threat. For 430 plants in 2001. In 2011, there were or herbicide application could affect example, the Indiana occurrence is more than 500 seedlings present at this them, there have been few documented located near an oxbow lake that was site, but no flowering plants were examples of such effects. In many created in a relict channel of the observed. While this occurrence is roadside locations, Short’s bladderpod Wabash River, and it is periodically located near a roadside, there have been occurs on steep slopes or bluffs, where inundated by floodwaters from the river. no documented impacts from roadside roadside maintenance would be In 2011, this occurrence was subjected maintenance. unlikely to affect the species unless the to a prolonged flood that killed most of Short’s bladderpod is considered road was widened, requiring alteration the Short’s bladderpod plants at this extirpated from two of the seven sites in or removal of the slope or bluff. location (Homoya, pers. comm., Tennessee where roadside maintenance Moreover, well-timed and carefully November 2012). There were thousands has been identified as a threat to the executed right-of-way maintenance of seedlings present at this site in 2010, species. At Tennessee EO 2, TDEC intended to control vegetation and this flood event likely eliminated (2009, p. 5) found the habitat to be too encroachment could be beneficial by the recruitment of most, if not all, of overgrown and Short’s bladderpod reducing shading and competition. those seedlings into the population. At

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least 100 plants were present at this site extirpation from even limited habitat is no longer found at two of these sites: in 2012 (Homoya, pers. comm., alteration or inundation. Kentucky EO 19, where canopy shading November 2012); however, it is not has been implicated in the species’ Overstory Shading known whether these were survivors of absence, and Tennessee EO 20, where the flood or new plants that had The most vigorous (Shea 1992, p. 24) most of the habitat for the species has sprouted from the seed bank. and stable (TDEC 2009, p. 1) Short’s been covered by rip-rap. While Short’s There are seven Tennessee bladderpod occurrences are found in bladderpod is presumed to be extant at occurrences that TDEC (2009, p. 2) locations where the canopy has Kentucky EO 1, there was only one reported could be affected by water remained relatively open over time. plant found at this site in 2009 (KNHP level manipulation. One of these, Overstory shading appears to have been 2012). The species was first collected at Tennessee EO 3, is located on a wooded a factor contributing to the this site in 1957, and despite several site slope above the upper reaches of waters disappearance of Short’s bladderpod at visits between then and 2009, only two impounded by Old Hickory Lake. There three sites in Kentucky (EO numbers 9, plants were seen there in 1992 (KNHP were more than 500 plants at this 19, and 20) and one in Tennessee (EO 2012). TDEC (2009, p. 3) lists trash location in 2008, and the position of 2) where Shea (1992, p. 4) observed dumping as a general threat to Short’s Short’s bladderpod within the forested heavy shading as a threat to the species bladderpod, but provides no specific area above the zone of routine water in 1992. Overstory shading has been information to support this conclusion. level fluctuation is unlikely to be identified as a threat to Indiana EO 1 (INHDC 2012), Kentucky EO 22 (KNHP Livestock Grazing affected by manipulation of water levels 2012), and Tennessee EOs 10, 21, and Livestock grazing historically in the lake. Shea (1993, p. 90) did not 24 (TNHID 2012), or 19 percent of presented a threat to Short’s mention water level manipulation in her known extant occurrences. Based on bladderpod, but we are not aware of any assessment of threats to this occurrence. these data, canopy shading has been threats currently posed by this land use. Tennessee EO 20, also in the upper implicated as a factor contributing to the In addition to potentially causing direct reaches of Old Hickory Lake, is disappearance of Short’s bladderpod harm to or loss of individual plants, presumed extirpated but was likely lost from four sites and has been identified livestock grazing on the steeply sloped to placement of rip rap along the as a limiting factor at nearly one-fifth of sites where Short’s bladderpod typically roadside where it occurs, as discussed remaining extant occurrences. occurs could increase soil erosion, above (please see Transportation Right- potentially uprooting individual plants Competition With Nonnative Plant of-Way Construction and Maintenance). and causing loss of the soil seed bank. Tennessee EO 12 is located on bluffs Species Shea (1993, p. 22) identified three overlooking the Cumberland River but Competition with or shading from Kentucky sites (EOs 9, 20, and 21) at not within an area managed as a invasive, nonnative herbaceous and which livestock (goats or cows) grazing reservoir or lake. Shea (1993, pp. 22–23) shrub species are cited in notes posed a threat to Short’s bladderpod. was unable to find this occurrence in concerning threats in database records None of these sites support the species 1992, and concluded that flooding at the for three of Kentucky’s (EO numbers 4, today, likely due to multiple factors that base of the bluff was the cause. In 2008, 11, and 18) (KNHP 2012) and five of degraded the habitat at those locations. TDEC (2009, p. 8) found approximately Tennessee’s (EO numbers 8, 10, 22, 24, In Tennessee, Shea (1993, p. 22) 50 plants at Tennessee EO 12, but they and 26) (TNHID 2012) extant Short’s reported that EO numbers 15 and 21 considered Short’s bladderpod habitat bladderpod occurrences. Homoya (pers. were threatened by grazing. However, to be vulnerable to flooding at this site comm., December 2012) also lists more recent data from TDEC (TNHID due to water level fluctuation and the invasive species among the threats 2012) indicate that Short’s bladderpod position of the plants at a low elevation affecting the single Indiana occurrence. has remained relatively stable at these on the bluff. Tennessee EOs 24 through The species most often mentioned by sites and grazing is not listed among 27 are found in soil at the river bank or these agencies include Lonicera threats observed at these locations. on bedrock ledges within about 1.5 m (5 japonica (Japanese honeysuckle), L. Commercial and Residential ft) of the waters of Cordell Hull maackii (bush honeysuckle), Alliaria Construction Reservoir (TNHID 2012), but, with the petiolata (garlic mustard), and Bromus exception of EO 27, no more than 10 tectorum (downy brome grass); While TDEC (2009, p. 3) lists plants have ever been counted at any of however, several other invasive, commercial and residential construction these sites. These three occurrences are nonnative species occur in sites where among potential threats to Short’s vulnerable to the effects of water level Short’s bladderpod exists, including bladderpod, there is little fluctuation, as evidenced by observed Ligustrum spp. (privet), Rosa multiflora documentation of these impacts. erosion within the fluctuation zone (multiflora rose), and Glechoma Tennessee EO 31, which is based on a (TNHID 2012). Tennessee EO 27 appears hederacea (ground ivy). Competition single herbarium collection from 1979, to be at little risk of habitat alteration with or shading from these species was apparently lost due to construction due to water level fluctuation, as it is adversely affects Short’s bladderpod. activities at its location within the city located on bluff ledges above the zone While this threat has been specifically of Clarksville (TNHID 2012). The only of routine water level fluctuation. noted at approximately one-third of other reference we have found for this While the threat of flooding or water Short’s bladderpod occurrences, it likely particular threat was an observation by level fluctuation is present at only five is more widespread among occurrences TDEC (TNHID 2012) that an area in the extant occurrences (19 percent), one of of the species and has not been reported vicinity of Tennessee EO 21 had been these is the only Indiana population of in database records. subdivided for residential construction the species, where the species has on the bluffs overlooking Old Hickory numbered in excess of 1,000 plants in Trash Dumping Lake. Construction-related threats to the past (Homoya, pers. comm., Shea (1993, p. 22) identified three Short’s bladderpod could include direct November 2012). The four occurrences Short’s bladderpod sites at which trash destruction of habitat and the plants in Tennessee threatened by water level dumping posed a threat (Kentucky EOs found there or the indirect effects of fluctuation are small and vulnerable to 1 and 19, Tennessee EO 20). The species habitat alteration from sediment runoff

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and encroachment of invasive, pp. 111–122) provide some insight for Short’s bladderpod occurs by slowing nonnative plant species from areas evaluating the potential threat of climate rates of forest canopy development, disturbed during construction. change to the species. Most of the range increasing tree mortality, and increasing of Short’s bladderpod lies within the light availability for the species, which Climate Change geographic area included by Karl et al. could stimulate recruitment from Our analyses under the Act include (2009, pp. 111–122) in their summary of dormant seed banks and increase vigor consideration of ongoing and projected regional climate impacts affecting the of plants located in areas that are changes in climate. The terms ‘‘climate’’ Southeast region; however, the Indiana presently well-shaded. The predicted and ‘‘climate change’’ are defined by the occurrence of the species lies in the increase in flood frequency in the Intergovernmental Panel on Climate Midwest region, just west of its Midwest could place the Indiana Change (IPCC). ‘‘Climate’’ refers to the boundary with the Southeast region. population of the species at risk, as mean and variability of different types Since 1970, the average annual evidenced by the loss of large numbers of weather conditions over time, with 30 temperature across the Southeast has of seedlings during a prolonged flood at years being a typical period for such increased by about 2 °F, with the this site in 2011. While climate has measurements, although shorter or greatest increases occurring during changed in recent decades in regions longer periods also may be used (IPCC winter months. The geographic extent of where Short’s bladderpod occurs and 2007a, p. 78). The term ‘‘climate areas in the Southeast region affected by the rate of change likely will continue change’’ thus refers to a change in the moderate to severe spring and summer to increase into the future, we do not mean or variability of one or more drought has increased over the past have data to determine how the habitats measures of climate (e.g., temperature or three decades by 12 and 14 percent, where Short’s bladderpod occurs will be precipitation) that persists for an respectively (Karl et al. 2009, p. 111). affected by these changes and how the extended period, typically decades or These trends are expected to increase. species will respond to these changes. longer, whether the change is due to Rates of warming are predicted to more natural variability, human activity, or than double in comparison to what the Conservation Efforts To Reduce Habitat both (IPCC 2007, p. 78). Various types Southeast has experienced since 1975, Destruction, Modification, or of changes in climate can have direct or with the greatest increases projected for Curtailment of Its Range indirect effects on species. These effects summer months. Depending on the There have been limited conservation may be positive, neutral, or negative and emissions scenario used for modeling efforts directed towards reducing threats they may change over time, depending change, average temperatures are affecting Short’s bladderpod and its on the species and other relevant expected to increase by 4.5 °F to 9 °F habitat. The Indiana Department of considerations, such as the effects of by the 2080s (Karl et al. 2009, p. 111). Natural Resources acquired the single interactions of climate with other While there is considerable variability Indiana occurrence. IDNR controls variables (e.g., habitat fragmentation) in rainfall predictions throughout the competing vegetation by mowing along (IPCC 2007, pp. 8–14, 18–19). In our region, increases in evaporation of the roadside where Short’s bladderpod analyses, we use our expert judgment to moisture from soils and loss of water by occurs and attempts to stimulate weigh relevant information, including plants in response to warmer germination and seedling recruitment uncertainty, in our consideration of temperatures are expected to contribute with light soil disturbance. The species various aspects of climate change. to increased frequency, intensity, and has responded positively, at least in the The Intergovernmental Panel on duration of drought events (Karl et al. short term, to this management Climate Change (IPCC) concluded that 2009, p. 112). (Homoya, pers. comm., December 2012). evidence of warming of the climate Projected increases in winter and In Kentucky, a Landowner Incentive system is unequivocal (IPCC 2007a, p. spring rainfall for the Midwest region, Program grant was used to manage 30). Numerous long-term climate as well as predictions of more intense vegetation structure or control invasive changes have been observed including rainfall events throughout the year, are species at two occurrences in 2005. The changes in arctic temperatures and ice, expected to lead to more frequent effort to control bush honeysuckle at widespread changes in precipitation flooding. Despite these projected trends, Kentucky EO 19 provided only a short- amounts, ocean salinity, wind patterns the likelihood of drought is expected to term benefit, if any, for Short’s and aspects of extreme weather increase in the Midwest due to bladderpod, as bush honeysuckle is including droughts, heavy precipitation, warming-induced increases in again well established at this site. heat waves and the intensity of tropical evapotranspiration rates and longer During 2011, no Short’s bladderpod cyclones (IPCC 2007b, p. 7). While intervals between precipitation events plants could be found at this site, and continued change is certain, the (Karl et al. 2009, pp. 120–121). the occurrence is presumed extirpated. magnitude and rate of change is Depending on timing and intensity of The removal of cedar trees at Kentucky unknown in many cases. Species that drought events, Short’s bladderpod EO 23 appears to have positively are dependent on specialized habitat could be adversely affected by increased affected habitat conditions for Short’s types, are limited in distribution, or mortality rates, reduced reproductive bladderpod, as there were more than have become restricted to the extreme output due to loss or reduced vigor of 500 plants, mostly seedlings, observed periphery of their range will be most mature plants, and reduced rates of seed at the site in 2011. The Kentucky State susceptible to the impacts of climate germination and seedling recruitment. Nature Preserve Commission acquired change. The species’ presumed ability to form a lands to establish the Rockcress Hills Estimates of the effects of climate seed bank should provide some State Nature Preserve, where Kentucky change using available climate models resilience to drought-induced EO 22 is located and where the federally lack the geographic precision needed to population declines; however, multiple listed endangered Braun’s rockcress predict the magnitude of effects at a droughts in successive years could (listed as Arabis perstellata, but now scale small enough to discretely apply diminish this resilience and lead to the recognized as Boechera perstellata) also to the range of Short’s bladderpod. loss of occurrences. Conversely, occurs. As discussed above, this However, data on recent trends and increased drought frequency and occurrence is threatened by shading due predicted changes for the Southeast and severity could alter structure of to forest canopy development. These Midwest United States (Karl et al. 2009, vegetation communities in which conservation efforts have benefited three

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extant Short’s bladderpod occurrences, to Factor D under the Act, we interpret intent of this statute is not to ameliorate but significant habitat threats remain this language to require the Service to the threats identified for the species, but across the species’ range. consider relevant Federal, State, and it does provide information on the tribal laws, plans, regulations, and other species. Summary of Factor A such mechanisms that may minimize The Tennessee Rare Plant Protection The threats to Short’s bladderpod any of the threats we describe in threat and Conservation Act of 1985 (T.C.A. from habitat destruction and analyses under the other four factors, or 11–26–201) authorizes the Tennessee modification are occurring throughout otherwise enhance conservation of the Department of Environment and the entire range of the species. These species. We give strongest weight to Conservation (TDEC) to, among other threats include transportation right-of- statutes and their implementing things: conduct investigations on way construction and maintenance; regulations and to management species of rare plants throughout the flooding and water level fluctuation; direction that stems from those laws and state of Tennessee; maintain a listing of overstory shading; and competition with regulations. An example would be State species of plants determined to be nonnative plant species. The population governmental actions enforced under a endangered, threatened, or of special level impacts from these activities are State statute or constitution, or Federal concern within the state; and regulate expected to continue into the future. action under statute. the sale or export of endangered species Trash dumping, livestock grazing, and Having evaluated the significance of via a licensing system. This act forbids commercial and residential construction the threat as mitigated by any such persons from knowingly uprooting, have been recognized as threats to conservation efforts, we analyze under digging, taking, removing, damaging, habitat for this species, but there is little Factor D the extent to which existing destroying, possessing, or otherwise evidence that these are significant regulatory mechanisms are inadequate disturbing for any purpose, any threats to extant occurrences. to address the specific threats to the endangered species from private or species. Regulatory mechanisms, if they public lands without the written B. Overutilization for Commercial, exist, may reduce or eliminate the permission of the landowner, lessee, or Recreational, Scientific, or Educational impacts from one or more identified other person entitled to possession and Purposes threats. In this section, we review prescribes penalties for violations. The There has been limited collection of existing State and Federal regulatory TDEC may use the list of threatened and Short’s bladderpod seed for mechanisms to determine whether they special concern species when conservation purposes. The Missouri effectively reduce or remove threats to commenting on proposed public works Botanical Garden holds seed accessions Short’s bladderpod. projects in Tennessee, and the from the Indiana occurrence, four Short’s bladderpod is listed as department shall encourage voluntary Kentucky occurrences (EOs 4, 18, 19, endangered in Indiana, Kentucky, and efforts to prevent the plants on this list and 28), and two Tennessee occurrences Tennessee. In Indiana this listing does from becoming endangered species. (EOs 4 and 17). Kentucky EO 19 is no not provide legal protection for the This authority shall not, however, be longer extant, for reasons discussed species; although, listed species are used to interfere with, delay, or impede above, but Short’s bladderpod is still given special consideration when any public works project. found at all of the other occurrences planning government-funded projects. Thus, despite the fact that Short’s from which these accessions were Additionally, the Indiana site is located bladderpod is listed as endangered by collected. Dr. Carol Baskin (pers. on land owned by the IDNR where the states of Indiana, Kentucky, and comm., December 2012) collected seeds collection or damage to plants is Tennessee, these designations confer no from Indiana for research on seed prohibited. guarantee of protection to the species or ecology. We are not aware of The Kentucky Rare Plants Recognition its habitat, whether on privately owned commercial trade in Short’s bladderpod Act, Kentucky Revised Statutes (KRS), or state-owned lands, unless such at this time. Indiscriminate collecting chapter 146, section 600–619, directs protections are voluntarily extended to for scientific or other purposes could be the KSNPC to identify plants native to the species. a threat to the species due to the low Kentucky that are in danger of extirpation within Kentucky and report E. Other Natural or Manmade Factors numbers of individuals at most Affecting Its Continued Existence occurrences, but we have no data to every 4 years to the Governor and indicate that indiscriminate collecting General Assembly on the conditions and The ability of populations to adapt to of Short’s bladderpod has occurred. On needs of these endangered or threatened environmental change is dependent the contrary, collections for ex situ plants. This list of endangered or upon genetic variation, a property of conservation holdings could be an threatened plants in Kentucky is found populations that derives from its important component of future recovery in the Kentucky Administrative members possessing different forms efforts for the species. Regulations, title 400, chapter 3:040. (i.e., alleles) of the same gene (Primack The statute (KRS 146:600–619) 1998, p. 283). Small populations C. Disease or Predation recognizes the need to develop and occurring in isolation on the landscape We are not aware of any commercial maintain information regarding can lose genetic variation due to the or scientific data indicating that disease distribution, population, habitat needs, potentially strong influence of genetic or predation threatens the continued limiting factors, other biological data, drift, i.e., the random change in allele existence of Short’s bladderpod. and requirements for the survival of frequency from generation to generation plants native to Kentucky. This statute (Barrett and Kohn 1991, p. 8). Smaller D. The Inadequacy of Existing does not include any regulatory populations experience greater changes Regulatory Mechanisms prohibitions of activities or direct in allele frequency due to drift than do Section 4(b)(1)(A) of the Act requires protections for any species included in larger populations (Allendorf and the Service to take into account ‘‘those the list. It is expressly stated in KRS Luikart 2007, pp. 121–122). Loss of efforts, if any, being made by any State 146.615 that this list of endangered or genetic variation due to genetic drift or foreign nation, or any political threatened plants shall not obstruct or heightens susceptibility of small subdivision of a State or foreign nation, hinder any development or use of populations to adverse genetic effects, to protect such species. . . .’’ In relation public or private land. Furthermore, the including inbreeding depression and

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loss of evolutionary flexibility (Primack depression (Allendorf and Luikart 2007, occurrences, maximum recorded 1998, p. 283). Deleterious effects of loss p. 306). abundance ranged from 3 clusters of of genetic variation through drift have Evidence in plants of inbreeding plants at EO 26 to approximately 50 been termed drift load, which is depression due to small population size plants each at EOs 8, 12, 22, 27, and 29 expressed as a decline in mean is provided by Heschel and Paige (1995, (TNHID 2012). These small populations population performance of offspring in p. 128), who found that plants from are at risk of adverse effects from small populations (Willi et al. 2005, p. populations of Ipomopsis aggregata reduced genetic variation and associated 2260). (scarlet gilia) with 100 or fewer drift load, increased risk of inbreeding The likelihood that Short’s flowering individuals produced smaller depression, and reduced reproductive bladderpod is self-incompatible seeds with lower rates of germination output due to low availability of presents another threat related to small success compared to those from genetically compatible mates. Many of population sizes. Genetic populations with more than 100 these occurrences where population incompatibility prevents self- flowering individuals. Heschel and sizes are small are isolated from other fertilization or reduces successful Paige (1995, p. 131) also found that seed occurrences, decreasing the likelihood breeding among closely related sizes increased and germination success that they could be naturally individuals, which can decrease mean improved in response to transfer of reestablished via seed dispersal, in the fitness in small populations because of pollen into each of the small event that local extinction occurred. increased probability of an encounter of populations, which they interpreted as evidence that the reduced fitness Cumulative Effects From Factors A two incompatible haplotypes (specific through E combination of alleles at adjacent observed in small populations was locations (loci) on the chromosome that attributable, in part, to inbreeding Where two or more threats affect are inherited as a unit) (Willi et al. 2005, depression. Short’s bladderpod occurrences, the p. 2256), which would prevent seed Willi et al. (2005, pp. 2263) found effects of those threats could interact or production in self-incompatible plants. evidence of the three processes be compounded, producing a described above (reduced cross- In small populations, less common S- cumulative adverse effect that rises compatibility presumably due to lack of haplotypes (self-incompatibility above the incremental effect of either compatible mates carrying different S- haplotypes) might be easily lost due to threat alone. The most obvious cases in haplotypes, reduced fitness due to genetic drift, reducing the number of which cumulative adverse effects would inbreeding, and drift load due to loss of compatible mates within the population be significant are those in which small genetic variation) simultaneously (Byers and Meagher 1992, p. 356). populations (Factor E) are affected by affecting small populations of a plant, threats that result in destruction or In self-incompatible plants of the Ranunculus reptans (creeping Brassicaceae family, when pollen and modification of habitat (Factor A). Two buttercup). Populations with low allelic occurrences in Kentucky and six in stigma share S-haplotypes at the S-locus diversity, taken as an indication of long- (self-incompatibility locus, i.e., the Tennessee where small population size term small population size, had higher was identified as a threat also face position on a chromosome occupied by inbreeding levels. Inbreeding depression the self-incompatibility gene complex), threats to their habitats, as discussed in these populations was expressed as under Factor A above. The vulnerability pollen tube development is disrupted poor clonal performance and reduced on the stigma of the female reproductive of these occurrences to habitat seed production in offspring (F1 plants) modification or destruction is system (Takayama and Isogai 2005, p. produced by crosses between plants 469). The stigma is the receptive heightened by effects of small with high kinship coefficients. Drift population size discussed above, structure of the female reproductive load also was expressed as a reduction system in plants, which also includes reduced resilience to recover from acute in mean seed production of F1 plants in demographic effects of habitat the pistil and ovary, on which pollen long-term small populations (Willi et al. grains germinate and begin development disturbances, and low potential for 2005, p. 2260). recolonization due to isolation from of the pollen tube. Pollen tube formation In evaluating threats to Short’s other occurrences. is necessary for fertilization of the ovary bladderpod that could arise due to small and subsequent seed production to population size, we first evaluated the Whorled Sunflower occur. limited data available concerning A. The Present or Threatened Despite the presence of such a abundance at each of the occurrences mechanism functioning to reduce or across the species’ range. This Destruction, Modification, or eliminate reproductive output among represents a conservative classification Curtailment of Its Habitat or Range individuals sharing S-haplotypes, in of small population size, as available Whorled sunflower appears to be a small populations mating is likely to data typically do not discriminate narrow habitat specialist, occurring in occur among individuals that possess among life history stages, so the number natural wet meadows or prairies and different S-haplotypes but are of reproducing individuals is typically calcareous barrens. Such habitats likely genetically similar at other loci due to less than what is shown in the were more extensive in the eastern loss of alleles from the population abundance data in Table 1 (see United States before European through genetic drift (Byers and Distribution and Status for the Short’s settlement, subsequent fire suppression, Meagher 1992, p. 358). Mating between bladderpod, above). Less than 100 and conversion of habitat to cropland or such closely related individuals is individual plants have ever been residential areas (Allison 1995, p. 7). referred to as inbreeding. Inbreeding observed at one time at 12 (46 percent) Today these prairie areas are not very rates are higher in small populations of the extant occurrences in Kentucky extensive, and they often are degraded because most or all individuals in the (EOs 1, 11, and 28) and Tennessee (EOs or have been destroyed for a number of population are related, and inbred 8, 12, 15, 22, 24, 26, 27, 29, and 30). The reasons. Most remaining prairie individuals generally have reduced greatest number of plants ever observed vegetation in the geographic area where fitness as compared to non-inbred at the small Kentucky occurrences whorled sunflower occurs exists as individuals from the same population, a ranged from 2 at EO 1 to 52 at EO 11 remnants along roadside and utility phenomenon referred to as inbreeding (KNHP 2012). At the small Tennessee rights-of-way, where prairie-like

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conditions are artificially maintained subpopulations and could contribute to significant mortality (A. Bishop, (Allison 1995, p. 4). Where whorled habitat degradation caused by shading Botanist, TDEC, pers. comm., February sunflower habitat remains, it faces and competition (please see ‘‘Shading 2008; D. Lincicome, Natural Heritage threats due to indiscriminate use of and Competition’’ below) by improving Program Manager, TDEC, pers. comm., mechanical or chemical vegetation conditions for growth of planted pines. September 2006). Similarly, plants management for industrial forestry, During timber harvests, either to thin extending into the railroad right-of-way right-of-way maintenance, or (i.e., reduce density of pine trees in at the McNairy County, Tennessee, agricultural purposes that could order to improve growth conditions for population are vulnerable to adverse adversely affect it. Because the species remaining trees) or to clearcut the stand, effects from indiscriminate herbicide requires well-lit habitats for its growth whorled sunflower plants at this application for railroad right-of-way and reproduction, shading and subpopulation could be subjected to maintenance. A small cluster of plants competition due to vegetation indirect adverse effects from soil in one of the Georgia’s subpopulations succession in the absence of natural or disturbance or to direct mortality due to is located on the bank of a road adjacent human-caused disturbance also threaten movement of harvesting equipment. to the Coosa Valley Prairie easement whorled sunflower habitat. Right-of-Way Maintenance area and is not protected. These data indicate that effects of indiscriminate Industrial Forestry Practices Incompatible maintenance activities use of herbicides or mowing for Industrial forestry practices have in transportation rights-of-way have vegetation management in altered much suitable whorled adversely affected the whorled transportation rights-of-way could sunflower habitat in Georgia and sunflower in Alabama and Tennessee, adversely affect the whorled sunflower Alabama, and currently threaten one and could affect one subpopulation in populations in Alabama and Tennessee, known subpopulation in Alabama. Georgia. At one of the Alabama While surveying potential habitat for subpopulations, the whorled sunflower as well as a small subpopulation in additional populations, J. Allison occurs in a narrow strip of vegetation Georgia. (Botanist, Georgia Department of between a roadside and adjacent pine Agricultural Practices and Land Natural Resources, pers. comm., March forest, where it is vulnerable to Conversion 1999) observed that much of this mortality or reduced vigor and species’ prairie habitat in Georgia had reproductive output due to The whorled sunflower has not been been converted to pine plantations. indiscriminate use of herbicides or rediscovered at the type locality in Nearly all of the Georgia subpopulations mowing for right-of-way maintenance. Tennessee despite intensive surveys of and one of the Alabama subpopulations Poorly timed mowing of this right-of- that area (Nordman 1998, p. 1–2). of whorled sunflower are located on way prevented flowering and seed However, this record is from an 1892 lands that currently are owned by The production in some plants at this site in collection and locality information is Campbell Group, a timberland 2008; however, the Alabama vague, so it is not possible to determine investment advisory firm. The Georgia Department of Conservation and Natural why this population has been lost. In subpopulations on The Campbell Resources, Alabama Department of Tennessee, much of this species’ Group’s lands are protected from habitat Transportation, and Cherokee County suitable habitat presumably has been destruction or degradation by their Highway Department cooperated in converted for agricultural use, as inclusion in the conservation easement placing signs at the site to mark the substantial proportions of the counties area at the Coosa Valley Prairie, which presence of whorled sunflower and to in the State where the species have been was donated to The Nature Conservancy attempt to prevent this in the future (W. found have been in row crop production by the Temple-Inland Corporation, the Barger, Botanist, Alabama Department since 1850 (Table 5) (Waisanen and former owner of these lands. of Conservation and Natural Resources, Bliss 2002; GIS data available at With the exception of the pers. comm., February 2009); periodic http://landcover.usgs.gov/cropland, conservation easement area at the Coosa replacement might be needed due to accessed January 9, 2013). Because this Valley Prairie, The Campbell Group vandalism or removal of the signs species was not seen following the typically subsoil plows planting sites to (Barger, pers. comm., March 2012). initial 1892 collection until it was improve drainage and conditions for Regular coordination with parties rediscovered in 1994, and was not seen tree root development, and uses responsible for roadside maintenance at again in Tennessee until 1998, it is mechanical or chemical methods to this location will be necessary to avoid impossible to know the historical control competing vegetation when future adverse effects to the whorled distribution and abundance of its preparing sites for planting pine sunflower from indiscriminate mowing habitat. However, the data in Table 5 seedlings (J. King, Area Manager, The or herbicide application. indicate that land conversion to Campbell Group, LLC, pers. comm., Plants extending onto a roadside agricultural uses has a long and August 2012) on its lands in Floyd within a powerline right-of-way at the sustained history in the Tennessee County, Georgia, and Cherokee County, Madison County, Tennessee, population counties where the whorled sunflower Alabama. These practices could cause were subjected to herbicide spraying in has been found and likely has direct mortality of whorled sunflower association with roadside and powerline contributed to loss of habitat and plants at one of the Alabama maintenance in 2004, causing whorled sunflower populations.

TABLE 5—PROPORTIONS OF COUNTY LAND BASE CONSIDERED IMPROVED FARMLAND FOR TENNESSEE COUNTIES WHERE THE WHORLED SUNFLOWER HAS BEEN FOUND. REPORTED HERE FOR EACH COUNTY ARE THE HIGHEST AND LOW- EST PROPORTIONS ON RECORD FOR EACH COUNTY AND THE YEARS IN WHICH THEY OCCURRED AND VALUES FOR THE YEARS 1850 AND 1997, THE FIRST AND LAST YEARS INCLUDED IN WAISANEN AND BLISS (2002).

County High (year) Low (year) 1850 1997

Chester ...... 37 (1940) 18 (1850) 18 23 Madison ...... 54 (1949) 23 (1870) 28 29

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County High (year) Low (year) 1850 1997

McNairy ...... 33 (1920) 14 (1850) 14 20

Agricultural practices, including field plans are in place to ensure that this site Southeast has increased by about 2 °F, preparation, herbicide use, and receives periodic disturbance to with the greatest increases occurring harvesting of crops, are threats to both maintain open conditions needed for during winter months. The geographic of the known Tennessee populations, the growth and sexual reproduction of extent of areas in the Southeast region due to the species’ presence in habitats whorled sunflower. affected by moderate to severe spring adjacent to actively farmed crop fields The Alabama subpopulation on The and summer drought has increased over in both locations. In July 2009, TDEC Campbell Group’s lands is located in a the past three decades by 12 and 14 biologists observed that one clump site where the prior owner, Temple- percent, respectively (Karl et al. 2009, p. consisting of two whorled sunflower Inland Corporation, harvested an 111). These trends are expected to stems had been destroyed by row crop immature hardwood forest in 1998. increase. Rates of warming are predicted cultivation in a previously fallow field Initially this timber harvest was thought to more than double in comparison to at the McNairy County, Tennessee, to have adversely affected the whorled what the Southeast has experienced population. Unpaved access roads sunflower population, but these plants since 1975, with the greatest increases around the perimeter of this field had and associated prairie species projected for summer months. also been widened, encroaching on responded favorably within a few years Depending on the emissions scenario whorled sunflower plants (7 clumps, following the harvest. However, the site used for modeling change, average 140 stems) in an adjacent railroad right- was subsequently converted into a temperatures are expected to increase by of-way (Bishop, pers. comm., March loblolly pine plantation, and the trees 4.5 °F to 9 °F by the 2080s (Karl et al. 2010). With the exception of the have attained sufficient size and density 2009, p. 111). While there is approximately 1-ha (2.5-ac) patch of old to threaten whorled sunflower plants considerable variability in rainfall field habitat discussed above (see due to increased shading and predictions throughout the region, Habitat for the whorled sunflower, competition (Schotz 2011, p. 4). As of increases in evaporation of moisture above), the Madison County, Tennessee, 2012, there were few whorled sunflower from soils and loss of water by plants in whorled sunflower population is plants present at this site, and those response to warmer temperatures are distributed in narrow strips of present were in a suppressed, vegetative expected to contribute to increased vegetation along borders of row crop condition due to strong shading and frequency, intensity, and duration of fields and is vulnerable to mechanized competition from planted pines and drought events (Karl et al. 2009, p. 112). disturbance of these habitats or to vegetation growing in the understory. The predicted increase in drought effects from herbicide application. Encroachment by invasive, nonnative frequency, intensity, and duration could Based on this information we conclude plants following the timber harvest and adversely affect the moist prairie that habitat at both whorled sunflower establishment of the loblolly pine stand habitats inhabited by whorled populations in Tennessee face also is a threat at this site (Schotz 2011, sunflower, by reducing soil moisture significant threats associated with p. 12). The second Alabama and increasing sunflower mortality rates agricultural practices used in row crop subpopulation is relegated to a narrow or reducing flowering and seed production. strip of vegetation between a roadside production rates. A positive effect of and adjacent pine forest with a densely increased drought could result from Shading and Competition vegetated understory. The spatial extent increased mortality of woody vegetation Absent natural or human-caused of this subpopulation is limited by the and reduced rates of vegetation disturbance, habitats where whorled whorled sunflower’s inability to grow in succession, which diminishes habitat sunflower occurs are threatened by the shaded habitat of the adjacent forest. abundance and quality for whorled succession of vegetation to a shrub- Based on this information we sunflower. While climate has changed dominated or forested condition. The conclude that habitat degradation due to in recent decades in the region where largest concentration of plants at the shading and competition resulting from whorled sunflower occurs and the rate Madison County, Tennessee, population vegetation succession currently is a of change likely will continue to is located in a successional old field significant threat to two whorled increase into the future, we do not have approximately 1 ha (2.5 ac) in size, sunflower populations. Both of the data to determine how the habitats where vegetation succession threatens Alabama subpopulations and the largest where the whorled sunflower occurs to degrade the largest patch of contiguous patch of suitable occupied will be affected by these changes and contiguous habitat where the majority of habitat for the species in Tennessee are how the species will respond to these this population occurs. Woody species at risk from this threat. changes. present at this site include Acer negundo (box elder), Liquidambar Climate Change Conservation Efforts To Reduce Habitat styraciflua (sweetgum), and Salix nigra We discuss the topic of climate Destruction, Modification, or (black willow) (Tennessee Division of change in greater detail above in the Curtailment of Its Range Natural Areas 2006, p. 5), all of which Factor A threats analysis for Short’s Temple-Inland Corporation donated a can rapidly invade moist old field bladderpod, which is also applicable to conservation easement for the Coosa habitats if left unmanaged. No whorled sunflower. Since 1970, the Valley Prairie property in Georgia to conservation agreements or management average annual temperature across the The Nature Conservancy, thereby

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protecting most of the Georgia industrial forestry practices, Policy Act (GEPA; O.C.G.A. 12–16–1), population of this species. This site maintenance of transportation and enacted in 1991, requires that impacts to drains into the headwaters of Mud utility rights-of-way, agricultural protected species be addressed for all Creek. In 2002, The Georgia Department practices, and shading and competition. projects on State-owned lands, and for of Natural Resources and The Nature While a conservation easement and all projects undertaken by a Conservancy worked with staff of suitable habitat management alleviate municipality or county if funded half or Temple-Inland to develop a 10-year threats from industrial forestry that more by State funds, or by a State grant management plan for conservation of otherwise would adversely affect the of more than $250,000. The provisions rare species within this easement area. Georgia population, one of the Alabama of GEPA do not apply to actions of Site-specific management plans for whorled sunflower subpopulations nongovernmental entities. On private several open wet prairies, known to currently is threatened by industrial lands, the landowner has ultimate provide habitat for this species within forestry practices. The population-level authority over what protection efforts, if the easement, were developed. Temple- impacts from these activities are any, occur with regard to protected Inland implemented a prescribed burn expected to continue into the future. plants (Patrick et al. 1995, p. 1 of section and selective timber harvest on 243 ha titled ‘‘Legal Overview’’). B. Overutilization for Commercial, (600 ac) of the easement in 2001, to The Tennessee Rare Plant Protection Recreational, Scientific, or Educational improve habitat conditions for whorled and Conservation Act of 1985 (T.C.A. Purposes sunflower and other species. Temple- 11–26–201) authorizes the Tennessee Inland conducted additional burns The whorled sunflower currently is of Department of Environment and within the easement area between 2002 limited availability in the horticultural Conservation (TDEC) to, among other and 2006. Mechanical thinning and trade, although no negative impacts are things: conduct investigations on control of invasive, exotic plants was known to have occurred due to species of rare plants throughout the also a component of their management collection of wild material for state of Tennessee; maintain a listing of of this site. commercial sale. Nonetheless, the species of plants determined to be This easement area, now owned by conspicuous, attractive flowers of this endangered, threatened, or of special The Campbell Group, is cooperatively species combined with easy access of concern within the state; and regulate managed with The Nature Conservancy some sites leaves the species vulnerable the sale or export of endangered species based on a jointly developed to collection or poaching. Poaching from via a licensing system. This act forbids conservation management plan, which the small populations of whorled persons from knowingly uprooting, was revised in 2012, for the period sunflower that are known to exist could digging, taking, removing, damaging, extending through 2016. The contribute to altered demographic or destroying, possessing, or otherwise management goals for the site are based genetic structure of populations, disturbing for any purpose, any on the conservation easement and potentially diminishing their viability; endangered species from private or include long-term perpetuation and however, we have no information to public lands without the written restoration of the mosaic of prairies, suggest this currently is an active threat permission of the landowner, lessee, or woodlands, wetlands, creeks, and forest or has adversely affected populations in other person entitled to possession and while allowing for sustainable timber the past. prescribes penalties for violations. The harvesting. Protecting and enhancing TDEC may use the list of threatened and C. Disease or Predation native plant communities, especially special concern species when those supporting rare species, is the We are not aware of any commercial commenting on proposed public works primary management objective, and or scientific data indicating that disease projects in Tennessee, and the periodic timber harvesting is a or predation threatens the continued department shall encourage voluntary secondary objective. Portions of the tract existence of whorled sunflower. efforts to prevent the plants on this list either have been or will be planted into D. The Inadequacy of Existing from becoming endangered species. Pinus palustris (longleaf pine) as part of Regulatory Mechanisms This authority shall not, however, be the Longleaf Alliance partnership. used to interfere with, delay, or impede Prescribed fire is the primary Whorled sunflower is State-listed as any public works project. management tool used to perpetuate and endangered in Georgia and Tennessee, Thus, despite the fact that whorled restore the native plant communities but has no official State status in sunflower is listed as endangered by the and also serves silvicultural objectives. Alabama. The law that provides official states of Georgia and Tennessee, these Despite the existence of a protection to designated species of designations confer no guarantee of conservation plan and the cooperative plants in Georgia is known as the protection to the species or its habitat, partnership between The Nature Wildflower Preservation Act of 1973 whether on privately owned or state- Conservancy and The Campbell Group (O.C.G.A. 12–6–170). Under this law, no owned lands, unless such protections to implement the plan, management protected plant may be collected are voluntarily extended to the species with prescribed fire is not a binding without written landowner permission. by owners or managers of lands where condition of the conservation easement. No protected plant may be transported the species is present. Thus, the potential remains that this within Georgia without a transport tag E. Other Natural or Manmade Factors management could be discontinued in with a permit number affixed. Permits Affecting Its Continued Existence the event that the property was sold to are also used to regulate a wide array of a less cooperative landowner. conservation activities, including plant The whorled sunflower is vulnerable rescues, sale of protected species, and to localized extinction because of its Summary of Factor A propagation efforts for augmentation of extremely restricted distribution and The threats to whorled sunflower natural populations and establishment small population sizes at most known from habitat destruction and of new ones. No protected plants may be locations, which reduces the resilience modification are occurring throughout collected from State-owned lands of these populations to recover from the entire range of the species. These without the express permission of the acute demographic effects of threats to threats include mechanical or chemical Georgia Department of Natural its habitat discussed above under Factor vegetation management associated with Resources. The Georgia Environmental A. Whorled sunflower is dependent

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upon existence of prairie-like openings Fleshy-Fruit Gladecress Transportation Right-of-Way or remnant roadside prairie habitats for Maintenance Factor A. The Present or Threatened its survival. Alteration or elimination of Destruction, Modification, or Five of the six fleshy-fruit gladecress disturbance processes that maintain Curtailment of Its Habitat or Range occurrences extend onto roadsides or these openings could result in the are near roads, where mowing and extinction of populations of this species. This species is endemic to cedar glade herbicide application prior to seed set Further, the highly fragmented areas in north-central Alabama that have pose threats to the species. Three distribution of populations within been significantly altered from their historical sites near roads have not been Tennessee, combined with their original condition. More than a 50 relocated and a portion of one of the disjunct location with respect to those percent loss in glade habitat has extant populations was destroyed by in Georgia and Alabama, presumably occurred since European settlement road widening and grading in the 1980s precludes gene flow among them and (Hilton 1997), with resulting glade (McDaniel and Lyons 1987, p. 7–9). leaves little chance of natural habitats reduced to remnants Additional road widening at this site in recolonization of these populations in fragmented by agriculture and recent years has further reduced the size the event of localized extinctions. development. Hilton (1997) conducted a of this population (Schotz 2009, p. 14). Small population size could be thorough survey of cedar glade The largest population of this species affecting reproductive fitness of the communities in northern Alabama using has a dirt road traversing through a whorled sunflower. The findings of Ellis historical records, soil maps, portion of the site, which has made the and McCauley (2008, entire) suggest that topographic maps, geology, and aerial site vulnerable to off-road vehicles and the Madison County, Tennessee, photography; 22 high priority glades dumping (Hilton 1997, p. 31). Other population is reproductively less fit were identified. However, field surveys sites have also been negatively affected than the Alabama population. Ellis and found only five of these to be in good by trash dumping and off-road vehicles, McCauley (2008, p. 1840) offered two condition and restorable, and only two including the site on U.S. Forest Service land. The U.S. Forest Service has posted possible explanations for reduced of these were considered high-quality the area as closed and recently gated the reproductive fitness of the Tennessee sites (Hilton, pers. comm., 1999). population, including limited mate area to block all-terrain vehicle access to availability due to limited diversity of Agricultural Practices the site (T. Counts, U.S. Forest Service, self-incompatibility alleles, or more in litt. 2008), which appears to have extensive inbreeding. Both could be At four of the fleshy-fruit gladecress been effective at reducing damage to the contributing to reduced seed production populations, plants occur in pasture glade (A. Cochran, U.S. Forest Service, and viability rates. areas, on roadside rights-of-way, and/or in litt. 2005, Schotz in litt. 2007). The in planted fields surrounded by U.S. Forest Service continues to monitor Ellis and McCauley (2008, pp. 1837– agriculture or residential developments the glade site for impacts from 1838) could not assess the fitness of the (Hilton 1997, pp. 13–27). Periodic recreational vehicles and from other Georgia population because seed heads disturbance, such as plowing in row illegal vehicle activity (A. Cochran, collected for the study contained very crop farming, arrests succession and pers. comm., 2011). few viable achenes, which produced maintains populations in this type of poor germination rates. However, the Shading and Competition habitat; however, plowing or herbicide lack of viable achenes in seed heads application in the spring prior to seed Winter annuals, such as fleshy-fruit collected for this study suggests that set and dispersal could be detrimental gladecress, are excluded from many poor reproductive fitness could be a habitats because they are poor threat in this population, as well. to populations. Populations are enhanced by disturbance created from competitors (Baskin and Baskin 1985, p. Cumulative Effects From Factors A light grazing, but heavy grazing of 387). As with all annuals, this species’ through E pastures creates unfavorable conditions long-term survival at a locality is (i.e., soil compaction, nutrient dependent upon its ability to reproduce and reseed there every year. Thus, Where two or more threats affect enrichment) for fleshy-fruit gladecress. whorled sunflower populations, the populations decline and become at risk Plants have been severely trampled effects of those threats could interact or of local extinction if conditions remain where grazing is allowed during the be compounded, producing a unsuitable for reproduction for height of the plant’s flowering or cumulative adverse effect that rises successive years. The most vigorous above the incremental effect of either fruiting period. Grazing during the populations of the fleshy-fruit threat alone. Cumulative adverse effects reproductive period also reduces vigor gladecress are located in areas which are likely significant for whorled of the populations (Schotz, 2009, p. 2). receive full, or near full, sunlight at the sunflower because all of the populations Improving pastures with fertilizer canopy level and have limited are small and their reproductive fitness treatments or planting of forage grasses herbaceous competition (Hilton 1997, p. is likely diminished (Factor E), and the could eventually result in loss of 5). Rollins (1963, p. 17) documented the Alabama and Tennessee populations are populations due to competition. Lyons loss of fleshy-fruit gladecress affected by threats that result in (in litt. 1981 to R. Sutter) considered individuals caused by invading grasses destruction or modification of habitat that her failure to relocate many of the in an unweeded portion of an (Factor A). The vulnerability of these historical fleshy-fruit gladecress sites experimental plot, while fleshy-fruit occurrences to habitat modification or from the 1960s was due to the change gladecress individuals in the hand- destruction is heightened by the effects in agricultural practices from growing weeded part of the plot thrived. Hilton of small population size discussed corn to using those sites for cattle (1997, p. 12) was unable to relocate five above, reduced resilience to recover pastures. McDaniel and Lyons (1987, p. populations in abandoned fields and from acute demographic effects of these 11) considered the trend toward pastures, which McDaniel and Lyons disturbances, and low potential for converting agricultural sites from row (1987, p. 7–9) had noted as appearing recolonization due to isolation from crop cultivation to pasture as a primary suppressed due to competition from other occurrences. threat to the species. invading weedy species.

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Shading and competition are potential during the winter months. The gladecress inhabits Indian Tomb Hollow threats at the two largest populations of geographic extent of areas in the Glade, the one limestone glade present fleshy-fruit gladecress (Hilton 1997, p. Southeast region affected by moderate to on WBNF, with a surface area of 68). One site, reported to be widely severe spring and summer drought has approximately 2.7 ha (1.1 ac). WBNF open in 1968, is now partially shaded increased over the past three decades by conducted treatment of the nonnative due to closing of the canopy (Hilton 12 and 14 percent, respectively (Karl et invasive species Ligustrum sinense 1997, p.18). Nonnative plants, including al. 2009, p. 111). These trends are (Chinese privet) on the Indian Tomb Ligustrum vulgare (common privet) and expected to increase. Rates of warming Hollow Glade in the fall of 2009 and Lonicera maackii (bush honeysuckle), are predicted to more than double in summer of 2011. The U.S. Forest are a significant threat in many glades comparison to what the Southeast has Service has posted the area of the due to the ever present disturbances that experienced since 1975, with the gladecress population as closed to allow for their colonization (Hilton greatest increases projected for summer access and monitors impacts to the 1997, p. 68). Nonnative plant species months. Depending on the emissions glade from off-road vehicles. Seeds from pose a threat to one population of the scenario used for modeling change, the Indian Tomb Hollow Glade were fleshy-fruit gladecress, where they have average temperatures are expected to collected in May 2010, and sent to the established near an unimproved road increase by 4.5 °F to 9 °F by the 2080s traversing the site (Hilton 1997, p.18). (Karl et al. 2009. p. 111). While there is USDA National Center for Genetic Under natural conditions, cedar considerable variability in rainfall Resources Preservation for long-term glades are edaphically (related to or predictions throughout the region, storage. caused by particular soil conditions) increases in evaporation of moisture The Service funded a survey of cedar maintained through processes of from soils and loss of water by plants in glade habitats in the Moulton Valley drought and erosion interacting with response to warmer temperatures are physiographic region of northwestern other processes that disrupt expected to contribute to increased Alabama, the major area for this habitat encroachment of competing vegetation. frequency, intensity, and duration of type, in the late 1990s. A survey and Soils that develop on glades are easily drought events (Karl et al. 2009, p. 112). status update for all fleshy-fruit eroded, moving downslope or into A warmer climate with more frequent gladecress populations was part of that fractures in the substrate. The shallow droughts, but also extreme precipitation project. The Service recently funded soil, exposed rock, and frequently hot, events, may adversely affect fleshy-fruit surveys to update information on all dry summers create xeric conditions gladecress by altering the glade habitat populations of this species. All sites that regulate competition and shading the species requires. Ephemeral seeps were visited in 2006 and 2007, and from encroaching vegetation (Hilton and streams on glades provide surveys continued into 2009 (Schotz 1997, p. 5; McDaniel and Lyons 1987, p. microhabitats important to the 2009). This information will be used to 6; Baskin et al. 1986, p. 138; Rollins distribution of the species (Hilton 1997, develop conservation measures needed 1963, p. 5). Historically, periodic fires p. 5). Climate change may also improve to protect and enhance populations. also likely played a role in maintaining habitat conditions for invasive plant these communities (Shotz 2009, p. 1). species and other plants (USFWS 2010, Summary of Factor A Extant occurrences of fleshy-fruit p. 5). A positive effect of increased gladecress are primarily located in areas drought could result from increased The threats to fleshy-fruit gladecress modified for human use. These habitat mortality of woody vegetation and from habitat destruction and modifications have either eliminated or reduced rates of vegetation succession. modification are occurring throughout reduced the frequency of natural While climate has changed in recent the entire range of the species. These disturbance processes, such as fire, that decades in the region where fleshy-fruit threats include agricultural conversion would otherwise regulate encroachment gladecress occurs and the rate of change or incompatible practices, maintenance of competing vegetation. likely will continue to increase for the of transportation rights-of-way, foreseeable future, we are unable to residential and industrial development, Residential and Industrial Development determine how the habitats where and shading and competition. Hilton (pers. comm., 1999) considered fleshy-fruit gladecress occurs will be Conservation efforts of the U.S. Forest residential and industrial development affected by these changes and how the Service have removed threats associated that had taken place in the decade prior species will respond to these changes. with off-road vehicle use and to her study to be the primary threat to Conservation Efforts to Reduce Habitat encroachment of invasive species at one cedar glade communities and the Destruction, Modification, or site; however, maintenance of primary reason for the loss of cedar Curtailment of Its Range transportation right-of-ways and use of glade habitat. One of the six fleshy-fruit off-road vehicles could adversely affect The occurrence and its habitat on gladecress populations is located in the the remaining five extant populations. William B. Bankhead National Forest front yard of a private residence. The population-level impacts from these (WBNF) is protected due to its location However, at this time, we know of no activities are expected to continue into in a Native American cultural site and projects that would lead to the the future. destruction of habitat where this species the fact that cedar glade communities is currently located. are considered ‘‘rare communities’’ on Factor B. Overutilization for the WBNF and protected from Commercial, Recreational, Scientific, or Climate Change detrimental effects from agency actions Educational Purposes We discuss the topic of climate (A. Cochran, U.S. Forest Service, in litt. change in greater detail above in the 2005). A thorough survey of limestone There is no information to suggest Factor A threats analysis for Short’s and sandstone glades on the WBNF was that fleshy-fruit gladecress is collected bladderpod, which is also applicable to completed by Schotz in 2006. Nine for commercial, recreational, or the fleshy-fruit gladecress. Since, 1970, glades presently are known to occur on educational purposes, and we have no the average annual temperature across WBNF, with sandstone glades reason to believe that this factor will the Southeast has increased by about 2 constituting the largest percentage of become a threat to the species in the °F, with the greatest increases occurring glade surface area. The fleshy-fruit future.

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Factor C. Disease or Predation effects of threats discussed in the above bladderpod plants at five occurrences in sections. the past, and roadside maintenance or One occurrence was lost due to Three of the six populations of fleshy- road widening could adversely affect infection by mustard rust in the early fruit gladecress are small in size as a nearly 40 percent of the extant 1980s (Lyons and Antonovics 1991, p. result of effects of habitat loss discussed occurrences of the species due to their 274; McDaniel and Lyons 1987, p. 11). in the above sections. The loss of position in roadside habitats. Future We have no data to indicate whether populations and reductions in development of a commuter rail project this disease poses a significant long- population sizes have resulted in spatial to improve intercity commute options term threat to the species generally. isolation between these remnant between the cities of Nashville and There is no information regarding populations. These isolated populations Clarksville, Tennessee, could affect 27 predation of the species by wildlife. are vulnerable to extinction by percent of known extant occurrences, Grazing is ongoing across the range of reductions in genetic variation among including some locations where the the gladecress and occurs on portions of the populations (Klank et al. 2012, pp. species is found in greatest abundance. all extant population sites; however, 1–2; Shotz, pers. comm., 2013). Based Flooding and water level fluctuations there is no information to document that on this information we conclude that threaten 19 percent of extant Short’s cattle eat gladecress. No studies have the small number of populations and bladderpod occurrences, most notably been conducted to investigate the effect the small size of populations within the the single Indiana occurrence, where the of grazing or herbivory specifically on species’ limited range are significant species has been present in large fleshy-fruit gladecress. threats to fleshy-fruit gladecress. numbers but recently experienced a reduction in numbers due to prolonged Factor D. The Inadequacy of Existing Cumulative Effects From Factors A flooding. Overstory shading due to Regulatory Mechanisms Through E natural forest succession, combined The greatest threats to the gladecress Where two or more threats affect with shading and competition due to include loss of habitat and the plants fleshy-fruit gladecress occurrences, the invasive, nonnative shrubs and themselves due to actions that remove effects of those threats could interact or herbaceous species presents the most the substrate under the populations or be compounded, producing a widespread, imminent threat to Short’s that cover them up. These types of cumulative adverse effect that rises bladderpod, and has been implicated in actions have been associated with above the incremental effect of either the loss of several historic occurrences. conversion of native glades or pastures threat alone. Cumulative adverse effects These threats are expected to continue with glades and outcrops to other land could be significant for fleshy-fruit into the foreseeable future. uses and potentially herbicide gladecress because three of the six The Factor E analysis demonstrated applications for the purpose of extant populations are small (Factor E) that Short’s bladderpod is vulnerable to controlling invasive plants. State and and all but one of the extant occurrences adverse effects of small population size, Federal regulations that might help are affected by threats that result in the including potential for reduced genetic conserve rare species on State highway destruction or modification of habitat. variation, low numbers of compatible mates, increased likelihood of rights-of-way, including avoidance or The vulnerability of these occurrences inbreeding depression, and reduced minimization of habitat destruction, as to habitat modification or destruction is resilience to recover from acute well as regulations that would protect heightened by effects of small demographic effects of other threats to plants from herbicide applications, population size discussed above, the species and is habitat. Fewer than protect only already listed species, and reduced resilience to recover from acute 100 plants have ever been observed at therefore do not apply to gladecress. demographic effects of these one time at 12 (46 percent) of the 26 Likewise, no existing regulations protect disturbances, and low potential for extant occurrences, and many of these the species on privately owned land, recolonization due to isolation from occurrences are isolated from other where most of the remnant gladecress other occurrences. occurrences. Existing threats may be populations are found. Proposed Determinations exacerbated by the effects of ongoing Factor E. Other Natural or Manmade We have carefully assessed the best and future climate change, especially Factors Affecting Its Continued scientific and commercial data available projected increases in temperature and Existence regarding the past, present, and future increased frequency and severity of threats to Short’s bladderpod, whorled droughts in the Southeast and projected The fleshy-fruit gladecress is sunflower, and fleshy-fruit gladecress. increases in flooding in the Midwest. vulnerable to localized extinction Below we state which of the five factors Based on our review of the best because of the small number of are determined to be threats to these available scientific and commercial occurrences and the small population species and summarize the severity, information, we conclude that adverse sizes within the species’ limited range. timing, and significance of those threats. effects associated with small and often Small population sizes decrease the isolated populations, as described in the resilience of individual fleshy-fruit Short’s Bladderpod Factor E analysis, both alone and in gladecress occurrences to recover from The most significant threats to this conjunction with the widespread threats effects of other threats affecting the species are described under Listing described under Factor A, constitute species’ habitat. There are only six Factors A (the present or threatened significant threats to Short’s remaining flesh-fruit gladecress destruction, modification, or bladderpod. As discussed under Factor occurrences, and only one of these is curtailment of its habitat or range) and D, no regulatory mechanisms exist that protected. The loss of any occurrences E (other natural or manmade factors would prevent or restrict activities would significantly impact the species’ affecting its continued existence). Based described under Factor A that constitute viability by reducing its redundancy on on the Factor A analysis, we conclude significant threats to the species. the landscape, which would increase its that the loss and degradation of habitat Therefore, on the basis of best available vulnerability to stochastic represents the greatest threat to Short’s scientific and commercial information environmental stressors and reduce the bladderpod. Road construction has we have determined that Short’s species’ resilience to recover from caused the loss of habitat and all Short’s bladderpod is in danger of extinction

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throughout all or a significant portion of A that constitute significant threats to habitat and reduce genetic variation its range and that a proposed the species. Therefore, on the basis of among populations. There are only six determination as an endangered species best available scientific and commercial remaining flesh-fruit gladecress is appropriate. information we have determined that occurrences, and only one of these is whorled sunflower is in danger of protected. The loss of any occurrences Whorled Sunflower extinction throughout all or a significant would significantly impact the species’ The most significant threats to this portion of its range and that a proposed viability by reducing its redundancy on species are described under Listing determination as an endangered species the landscape, which would increase its Factors A (the present or threatened is appropriate. vulnerability to stochastic destruction, modification, or environmental stressors and reduce the Fleshy-fruit Gladecress curtailment of its habitat or range) and species’ resilience to recover from E (other natural or manmade factors The most significant threats to this effects of threats discussed in the above affecting its continued existence). Based species are described under Listing sections. on the Factor A analysis, we conclude Factors A (the present or threatened Based on our review of the best that the loss and degradation of habitat destruction, modification, or available scientific and commercial represents the greatest threat to whorled curtailment of its habitat or range) and information, we conclude that adverse sunflower. Past and ongoing risk of E (other natural or manmade factors effects associated with limited adverse effects from mechanical or affecting its continued existence). Based distribution and small population size, chemical vegetation management for on the Factor A analysis, we conclude as described in the Factor E analysis, industrial forestry, right-of-way that the loss and degradation of habitat both alone and in conjunction with the maintenance, or agriculture is a threat to represents the greatest threat to fleshy- threats described under Factor A, three of the four extant populations of fruit gladecress. The threats to fleshy- constitute significant threats to fleshy- this species. Modification of the fruit gladecress from habitat destruction fruit gladecress. As discussed under remnant prairie habitats that the species and modification are occurring Factor D, no regulatory mechanisms occupies due to shading and throughout the entire range of the exist that would prevent or restrict competition resulting from vegetation species. These threats include activities described under Factor A that succession also threatens these three agricultural conversion for use as constitute significant threats to the populations, limiting growth and pasture or incompatible practices, species. Therefore, on the basis of best reproductive output of whorled maintenance of transportation rights-of- available scientific and commercial sunflower. These threats are expected to way (including mowing and herbicide information we have determined that continue in the foreseeable future. A treatment prior to seed set along fleshy-fruit gladecress is in danger of conservation easement and suitable roadsides), the impacts of off-road extinction throughout all or a significant habitat management currently alleviates vehicles, dumping, residential and portion of its range and that a proposed these threats that otherwise would industrial development, and shading determination as an endangered species adversely affect the Georgia population. and competition. Conservation efforts of is appropriate. The Factor E analysis demonstrated the U.S. Forest Service have removed that whorled sunflower is vulnerable to threats associated with off-road vehicle Significant Portion of the Range localized extinction because of its use and encroachment of invasive The Act defines an endangered extremely restricted distribution and species at one site; however, species as ‘‘any species which is in small population sizes at most known maintenance of transportation right-of- danger of extinction throughout all or a locations. Small population size could ways and use of off-road vehicles could significant portion of its range.’’ A major be affecting reproductive fitness of adversely affect the remaining five part of the analysis of ‘‘significant whorled sunflower by limiting extant populations. portion of the range’’ requires availability of compatible mates or by Shading due to natural forest considering whether the threats to the causing higher rates of inbreeding succession and competition from species are geographically concentrated among closely related individuals. Both nonnative invasive plants presents a in any way. If the threats are essentially of these could be contributing to significant threat to fleshy-fruit uniform throughout the species’ range, reduced seed production and viability gladecress, and has been implicated in then no portion is likely to warrant rates, which limit the species’ ability to the loss of five historic occurrences. One further consideration. Based on the recovery from acute demographic effects site, reported to be widely open in 1968, threats to Short’s bladderpod, whorled of habitat loss or modification. The is now partially shaded due to closing sunflower, and fleshy-fruit gladecress species’ dependence on remnant prairie of the canopy and the presence of throughout their entire known ranges, habitats, which are isolated on the nonnative plants, including Ligustrum we find that these species currently are landscape, limits the potential for vulgare (common privet) and Lonicera in danger of extinction throughout all of recolonization in the event that maackii (bush honeysuckle), and these their ranges, based on the severity and localized extinction events occur. are significant threats in many glades scope of the threats described above. As Based on our review of the best due to the ever present disturbances that discussed above, these species are available scientific and commercial allow for nonnative plant colonization. proposed for listing as endangered information, we conclude that adverse These threats are expected to continue species, rather than threatened species, effects associated with extremely into the foreseeable future. because the threats are occurring now or restricted distribution and small and The Factor E analysis demonstrated will in the near term, and their potential isolated populations, as described in the that fleshy-fruit gladecress is vulnerable impacts to the species would be severe Factor E analysis, both alone and in to localized extinction because of the given the limited known distribution of conjunction with the threats described small number of occurrences and the the species, the small population sizes under Factor A, constitute significant small population sizes within its limited at many of the remaining sites, and the threats to whorled sunflower. As range. Small population sizes decrease small area occupied by many of these discussed under Factor D, no regulatory the resilience of individual fleshy-fruit populations, putting these species at mechanisms exist that would prevent or gladecress occurrences to recover from risk of extinction at the present time. As restrict activities described under Factor effects of other threats affecting its these threats extend throughout their

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entire ranges, it is unnecessary to progress. Recovery plans also establish Section 7(a) of the Act requires determine if they are in danger of a framework for agencies to coordinate Federal agencies to evaluate their extinction throughout a significant their recovery efforts and provide actions with respect to any species that portion of their ranges. Therefore, on the estimates of the cost of implementing is proposed or listed as an endangered basis of the best available scientific and recovery tasks. Recovery teams or threatened species and with respect commercial data, we propose listing (comprised of species experts, Federal to its critical habitat, if any is Short’s bladderpod, whorled sunflower, and State agencies, nongovernment designated. Regulations implementing and fleshy-fruit gladecress as organizations, and stakeholders) are this interagency cooperation provision endangered species throughout their often established to develop recovery of the Act are codified at 50 CFR part ranges in accordance with sections 3(6) plans. When completed, the recovery 402. Section 7(a)(4) of the Act requires and 4(a)(1) of the Act. outline, draft recovery plan, and the Federal agencies to confer with the Service on any action that is likely to Available Conservation Measures final recovery plan will be available on our Web site (http://www.fws.gov/ jeopardize the continued existence of a Conservation measures provided to endangered), or from the Service’s species proposed for listing or result in species listed as endangered or Tennessee Ecological Services Field destruction or adverse modification of threatened species under the Act Office (see FOR FURTHER INFORMATION proposed critical habitat. If a species is include recognition, recovery actions, CONTACT). listed subsequently, section 7(a)(2) of requirements for Federal protection, and Implementation of recovery actions the Act requires Federal agencies to prohibitions against certain practices. generally requires the participation of a ensure that activities they authorize, Recognition through listing results in broad range of partners, including other fund, or carry out are not likely to public awareness and conservation by Federal agencies, States, Tribal, jeopardize the continued existence of Federal, State, Tribal, and local nongovernmental organizations, the species or destroy or adversely agencies; private organizations; and businesses, and private landowners. modify its critical habitat. If a Federal individuals. The Act encourages Examples of recovery actions include action may affect a listed species or its cooperation with the States and requires habitat restoration (e.g., restoration of critical habitat, the responsible Federal that recovery actions be carried out for native vegetation), research, captive agency must enter into formal all listed species. The protection propagation and reintroduction, and consultation with the Service. required by Federal agencies and the outreach and education. The recovery of Federal agency actions within the prohibitions against certain activities many listed species cannot be species’ habitat that may require are discussed, in part, below. accomplished solely on Federal lands conference or consultation or both as The primary purpose of the Act is the described in the preceding paragraph conservation of endangered and because their range may occur primarily or solely on non-Federal lands. To include federally funded or permitted threatened species and the ecosystems actions occurring within habitat for achieve recovery of these species upon which they depend. The ultimate Short’s bladderpod, whorled sunflower, requires cooperative conservation efforts goal of such conservation efforts is the or fleshy-fruit gladecress (e.g., on private, State, and Tribal lands. recovery of these listed species, so that management and any other landscape they no longer need the protective If these species are listed, funding for altering activities on Federal lands measures of the Act. Subsection 4(f) of recovery actions will be available from administered by the U.S. Army Corps of the Act requires the Service to develop a variety of sources, including Federal Engineers or U.S. Forest Service; and implement recovery plans for the budgets, State programs, and cost share issuance of section 404 Clean Water Act conservation of endangered and grants for non-Federal landowners, the (33 U.S.C. 1251 et seq.) permits by the threatened species. The recovery academic community, and U.S. Army Corps of Engineers; planning process involves the nongovernmental organizations. In construction and management of gas identification of actions that are addition, pursuant to section 6 of the pipeline and power line rights-of-way necessary to halt or reverse the species’ Act, the States of Indiana, Kentucky, by the Federal Energy Regulatory decline by addressing the threats to its and Tennessee would be eligible for Commission; construction and survival and recovery. The goal of this Federal funds to implement maintenance of roads or highways process is to restore listed species to a management actions that promote the funded or carried out by the Federal point where they are secure, self- protection or recovery of Short’s Highway Administration; and Federal sustaining, and functioning components bladderpod. The States of Georgia and Emergency Management Agency-funded of their ecosystems. Tennessee would eligible for Federal actions). Also subject to consultation Recovery planning includes the funds to implement management would be provision of Federal funds to development of a recovery outline actions that promote the protection or State and private entities through shortly after a species is listed and recovery of whorled sunflower. Federal programs such as the Service’s preparation of a draft and final recovery Information on our grant programs that Partners for Fish and Wildlife Program, plan. The recovery outline guides the are available to aid species recovery can State Wildlife Grant Program, and immediate implementation of urgent be found at: http://www.fws.gov/grants. Federal Aid in Wildlife Restoration recovery actions and describes the Although Short’s bladderpod, Program. process to be used to develop a recovery whorled sunflower, and fleshy-fruit The Act and its implementing plan. Revisions of the plan may be done gladecress are only proposed for listing regulations set forth a series of general to address continuing or new threats to under the Act at this time, please let us prohibitions and exceptions that apply the species, as new substantive know if you are interested in to endangered plants. All prohibitions information becomes available. The participating in recovery efforts for this of section 9(a)(2) of the Act, recovery plan identifies site-specific species. Additionally, we invite you to implemented by 50 CFR 17.61, apply. management actions that set a trigger for submit any new information on this These prohibitions, in part, make it review of the five factors that control species whenever it becomes available illegal for any person subject to the whether a species remains endangered and any information you may have for jurisdiction of the United States to or may be downlisted or delisted, and recovery planning purposes (see FOR import or export, transport in interstate methods for monitoring recovery FURTHER INFORMATION CONTACT). or foreign commerce in the course of a

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commercial activity, sell or offer for sale taxa at least 100 years old, as defined by participate in a public hearing should in interstate or foreign commerce, or section 10(h)(1) of the Act; contact the Tennessee Ecological remove and reduce the species to (2) Unauthorized removal, damage, or Services Field Office at (931) 528–6481, possession from areas under Federal destruction of Short’s bladderpod or as soon as possible. To allow sufficient jurisdiction. In addition, for plants fleshy-fruit gladecress plants from time to process requests, please call no listed as endangered, the Act prohibits populations located on Federal land later than one week before the hearing the malicious damage or destruction on (lands owned by the U.S. Army Corps date. Information regarding this areas under Federal jurisdiction and the of Engineers or on which they hold proposed rule is available in alternative removal, cutting, digging up, or easements, or U.S. Forest Service lands); formats upon request. and damaging or destroying of such plants Required Determinations in knowing violation of any State law or (3) Unauthorized removal, damage or regulation, including State criminal destruction of Short’s bladderpod, Clarity of the Rule trespass law. Certain exceptions to the whorled sunflower, or fleshy-fruit Executive Order 12866 requires each prohibitions apply to agents of the gladecress plants on private land in agency to write regulations that are easy Service and State conservation agencies. violation of any State regulation, to understand. We invite your The States of Georgia, Indiana, including criminal trespass. comments on how to make this rule Kentucky, and Tennessee have Questions regarding whether specific easier to understand including answers regulations authorizing the activities would constitute a violation of to questions such as the following: promulgation of lists of endangered section 9 of the Act should be directed (1) Are the requirements in the rule plants; however, with the exception of to the Service’s Tennessee Ecological clearly stated? Georgia, these regulations create no Services Field Office (see FOR FURTHER (2) Does the rule contain technical obligations on the part of landowners, INFORMATION CONTACT). Requests for language or jargon that interferes with public or private, to protect State-listed copies of the regulations concerning its clarity? plants. The Georgia Environmental listed species and general inquiries (3) Does the format of the rule Policy Act requires that impacts to regarding prohibitions and permits may (grouping and order of sections, use of protected species be addressed for all be addressed to the U.S. Fish and headings, paragraphing, etc.) aid or projects on State-owned lands, and for Wildlife Service, 105 West Park Drive, reduce its clarity? all projects undertaken by a Suite D, Athens, GA 30606; telephone (4) Would the rule be easier to municipality or county if funded half or 706–613–9493; facsimile 706–613–6059. understand if it were divided into more (but shorter) sections? more by State funds, or by a State grant Peer Review of more than $250,000. The Act will, (5) Is the description of the rule in the therefore, offer additional protection to In accordance with our joint policy SUPPLEMENTARY INFORMATION section of these species. published in the Federal Register on the preamble helpful in understanding July 1, 1994 (59 FR 34270), we will seek We may issue permits to carry out the emergency rule? What else could we the expert opinions of at least three otherwise prohibited activities do to make the rule easier to appropriate and independent specialists involving endangered and threatened understand? regarding this proposed rule. The Send a copy of any comments that plant species under certain purpose of peer review is to ensure that concern how we could make this rule circumstances. Regulations governing our listing determination for these easier to understand to Office of permits are codified at 50 CFR 17.62 for species is based on scientifically sound Regulatory Affairs, Department of the endangered plants, and at 17.72 for data, assumptions, and analyses. We Interior, Room 7229, 1849 C Street NW., threatened plants. With regard to have invited these peer reviewers to Washington, DC 20240. You also may endangered plants, a permit must be comment during the public comment email the comments to this address: issued for the following purposes: for period. [email protected]. scientific purposes or to enhance the We will consider all comments and propagation or survival of the species. information received during the National Environmental Policy Act (42 It is our policy, as published in the comment period on this proposed rule U.S.C. 4321 et seq.) Federal Register on July 1, 1994 (59 FR during preparation of a final We have determined that 34272), to identify, to the maximum rulemaking. Accordingly, the final environmental assessments and extent practicable at the time a species decision may differ from this proposal. environmental impact statements, as is listed, those activities that would or defined under the authority of the would not constitute a violation of Public Hearings National Environmental Policy Act of section 9 of the Act. The intent of this The Act provides for one or more 1969, need not be prepared in policy is to increase public awareness of public hearings on this proposal, if connection with listing a species as an the effect of a proposed listing on requested. Requests must be received endangered or threatened species under proposed and ongoing activities within within 45 days after the date of the Act. We published a notice outlining the range of the species proposed for publication of this proposal in the our reasons for this determination in the listing. The following activities could Federal Register. Such requests must be Federal Register on October 25, 1983 potentially result in a violation of sent to the address shown in the FOR (48 FR 49244). section 9 of the Act; this list is not FURTHER INFORMATION CONTACT section. comprehensive: We will schedule public hearings on References Cited (1) Unauthorized collecting, handling, this proposal, if any are requested, and A complete list of all references cited possessing, selling, delivering, carrying, announce the dates, times, and places of in this rule is available on the Internet or transporting of Short’s bladderpod, those hearings, as well as how to obtain at http://www.regulations.gov under whorled sunflower, or fleshy-fruit reasonable accommodations, in the Docket No. FWS–R4–ES–2013–0087 or gladecress, including import or export Federal Register and local newspapers upon request from the Field Supervisor, across State lines and international at least 15 days before the hearing. Tennessee Ecological Services Field boundaries, except for properly Persons needing reasonable Office (see FOR FURTHER INFORMATION documented antique specimens of these accommodations to attend and CONTACT section).

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Authors Proposed Regulation Promulgation ■ 2. In § 17.12 paragraph (h), add entries for Helianthus verticillatus, The primary authors of this proposed Accordingly, we propose to amend rule are the staff members of the Leavenworthia crassa, and Physaria part 17, subchapter B of chapter I, title globosa, in alphabetical order under Tennessee Ecological Services Field 50 of the Code of Federal Regulations, FOR FURTHER INFORMATION FLOWERING PLANTS, to the List of Office (see as set forth below: CONTACT) and the Alabama Ecological Endangered and Threatened Plants, to Services Field Office. PART 17—[AMENDED] read as follows: List of Subjects in 50 CFR Part 17 § 17.12 Endangered and threatened plants. ■ 1. The authority citation for part 17 Endangered and threatened species, * * * * * Exports, Imports, Reporting and continues to read as follows: (h) * * * recordkeeping requirements, Authority: 16 U.S.C. 1361–1407; 1531– Transportation. 1544; 4201–4245, unless otherwise noted.

Species Historic range Family Status When listed Critical Special Scientific name Common name habitat rules

FLOWERING PLANTS

******* Helianthus whorled sunflower .. U.S.A. (AL, GA, TN) Asteraceae ...... E ...... NA NA verticillatus.

******* Leavenworthia fleshy-fruit U.S.A. (AL) ...... Brassicaceae ...... E ...... NA NA crassa. gladecress.

******* Physaria globosa ..... Short’s bladderpod U.S.A. (IN, KY, TN) Brassicaceae ...... E ...... NA NA

*******

* * * * * Dated: July 18, 2013. Stephen Guertin, Acting Director, U.S. Fish and Wildlife Service. [FR Doc. 2013–18213 Filed 8–1–13; 8:45 am] BILLING CODE 4310–55–P

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