Seven new species of Trichostema (Lamiaceae: Ajugoideae) from the North American Coastal Plain biodiversity hotspot Author Mcclelland, R. Kevan Schoonover 0000-0001-5353-4119 Department of Biology, Elon University, Elon, NC & kschoonovermcclell @ elon. edu; https: // orcid. org / 0000 - 0001 - 5353 - 4119 kschoonovermcclell@elon.edu Author Weakley, Alan S. 0000-0003-2093-3767 UNC-CH Herbarium (NCU), North Carolina Botanical Garden, and Department of Biology, University of North Carolina-Chapel Hill, Chapel Hill, NC & weakley @ unc. edu; https: // orcid. org / 0000 - 0003 - 2093 - 3767 weakley@unc.edu Author Poindexter, Derick B. 0000-0002-6446-7107 UNC-CH Herbarium (NCU), North Carolina Botanical Garden, and Department of Biology, University of North Carolina-Chapel Hill, Chapel Hill, NC & poindexter @ unc. edu; https: // orcid. org / 0000 - 0002 - 6446 - 7107 poindexter@unc.edu text Phytotaxa 2023 2023-07-18 603 2 95 149 http://dx.doi.org/10.1094/PDIS-04-22-0755-PDN journal article 56248 10.11646/phytotaxa.603.2.1 ebea7f61-a2fb-4618-88e2-1e9078b850ea 1179-3163 8157972 Trichostema fruticosum K. S. McClell. , sp. nov. ( Figures 9 and 10 ) Diagnosis: —Compact to diffuse bushy perennial to 1.25 m tall, morphologically most similar to Trichostema dichotomum . It differs from T. dichotomum by being a short-lived perennial ( vs. annual), its wider leaves of 0.74–1.6 cm ( vs. 0.59–1.25 cm ), smaller leaf length/ width ratio of 2.89–4.42 times long as wide ( vs. 3.38–4.92), lack of spreading trichomes more than 6–7 cells long on the stems and at nodes, ( vs. presence of spreading trichomes more than 6–7 cells long on the stems and at nodes; Figure 1 ), nutlets with short to mid-sized ridges and small to mid-sized alveoli ( vs. mid-sized to tall ridges and mid-sized to large alveoli; Figure 2 ), and year-long flowering time ( vs. July–November). Type: — U.S.A. Florida . Marion County . Restored Red Oak-Pine woodland, Mockernut Hill Botanical Garden , beside entrance gate, 16 October 2021 , R. Kevan Schoonover McClelland, Robert K. Schoonover, and Linda Duever 222a–w ( holotype NCU !; isotypes B !, BM !, F !, FLAS !, K !, G !, GH !, MEXU !, MO !, NCU !, NY !, OS !, P !, PH!, RSA!, UC !, US!, USF !) . Notes : Specimens R . Kevan Schoonover McClelland 222a–w represent a single gathering in accordance with Article 8.2 of the ICN ( Turland et al. 2018 ). Mockernut Hill Botanical Garden is a botanical garden oriented around land management training, showcasing trails through exemplary native vegetation rather than display gardens. The sampled population is one of many naturally occurring on the property and not of a horticultural origin. Bushy perennial to 1.25 m tall; branching throughout the plant; stem hairs downwardly curved, occasionally spreading to three, rarely to five, cells long ( Figure 1f ), with sessile and stalked glands to two cells long, rarely to three cells long (including gland); node hairs spreading, rarely downwardly curved, to four, rarely to six, cells long ( Figure 1f ); leaves widest towards the base, occasionally widest near the middle, rarely widest towards the tip, (1.6–)2.82–5.4(–7.4) × (0.33–)0.74–1.6(–2.6), average of 4.11 × 1.17, cm, (2.19–)2.89–4.42(–6.06), average of 3.65, times long as wide, base cuneate to attenuate, margin entire, apex obtuse, occasionally ±90°, rarely acute or rounded; inflorescence hairs spreading to upwardly curved, to three cells long, with sessile and spreading glands to two, rarely to three, cells long (including gland); bracts widest towards the tip or near the middle, rarely widest towards the base, rarely falcate (3.9–)6.41–12.03(–16.6) × (0.7–)1.21–2.71(–4.6), average of 9.22 × 1.96, mm, (2.65–)3.55–6.41(–11.3), average of 4.98, times long as wide, base cuneate to attenuate, margin entire, apex acute, rarely acuminate; fruiting calyx (5.6–)6.24–7.65(–9.2), average of 6.94, mm long; buds dark blue-purple to light blue-purple, corolla dark to light blue or blue-purple; anthers dark to light blue or blue-purple; style bifid, the longer arm 1–1.5, occasionally 2, times as long as the shorter; mature nutlets alveolate, (1.5–)1.71–2.06(–2.3), average of 1.88, mm long, hilum (0.4–)0.51–0.69(–0.9), average of 0.6, mm long, (0.24–)0.28–0.36(–0.44), average of 0.32, times the length of the nutlet, alveoli small to mid-sized, rarely large, ridges short, occasionally mid-sized ( Figure 2f ). FIGURE 9 . Type specimen of Trichostema fruticosum before pressing. R. Kevan Schoonover McClelland’s foot for scale (heel to toe ≈ 35 cm). Photo by R. Kevan Schoonover McClelland. FIGURE 10 . Holotype specimen of Trichostema fruticosum . Photo by R. Kevan Schoonover McClelland. Phenology: —Flowers and fruits April to December. Some plants will have a small number of flowers January to March. Etymology: —The epithet fruticosum (Latin, bushy) is given to this species based on its growth habit. It often has a woody base and forms a relatively large, much branched “bush” (to 1.25 m tall) compared to the typically smaller, more compact growth forms of T. floridanum and T. nesophilum and the more virgate branching of the members of the T. suffrutescens complex. The common name given to this species is Bushy Bluecurls. This is derived from the same feature as the specific epithet. The common name was suggested by Linda Duever, founder of Mockernut Hill Botanical Garden and co-collector of the type specimens. Habitat and distribution: — Trichostema fruticosum occurs in openings in a variety of mesic to semi-dry habitats with or without calcareous influences on the mainland and some barrier islands; 0–100 meters; USA : Alabama , Florida , Georgia , Mississippi . ( Figure 11 ). FIGURE 11 . Approximate range of Trichostema fruticosum based on field data and herbarium records. Conservation status: — Trichostema fruticosum , while having a more restricted range than T. dichotomum , is much more widespread than T. floridanum and T. nesophilum , most likely due to its habitat tolerances. It is common throughout its range, and it has also been introduced to the nursery trade under the species name of T. dichotomum . Although it is widespread, many of its populations are coastal or near coastal and under threat from rising sea levels and hurricanes. Based on the data collected for this study, this species should be recognized at a G3 ranking (NatureServe 2020). Most of the populations of this species occur in Florida and thus it was calculated to have an S3 ranking in Florida . The smaller numbers of populations in Alabama , Georgia , and Mississippi led those states having an S2 rank calculated for this species (NatureServe 2020). Notes: — Trichostema fruticosum was the only species not to form a clade in the BEAST analysis ( Figure 5 ). The sample from Estero Bay (EstBPu) is sister to the clade containing T. floridanum and T. nesophilum and the sample from Mississippi (NCU 667421) diverges at the base of the perennial T. dichotomum complex clade in the MCC tree. The reason for the incongruence in the placement of the Estero Bay specimen is likely due to its low number of high-quality reads mapped to the reference. This would explain why this specimen, which is geographically and morphologically distinct from T. nesophilum , is not in the clade with the rest of T. fruticosum . Thus, it is possible that the missing data would have aligned this specimen with the larger clade. This sample, though, based on its morphology, ecology, and biogeography still falls within the concept of T. fruticosum . The Mississippi sample poses a different issue. Unlike the Estero Bay sample, this sample had a high number of high-quality reads mapped to the reference. However, like the Estero Bay specimen, this specimen was collected at the edge of the species range, this time at the far western edge. It could be that this specimen represents a separate taxon. If this were true, its potential extent is west of the Tombigee River and Mobile Bay, with the T. fruticosum-T. floridanum-T. nesophilum clade all occuring east of that demarcating line, a biogeographic pattern shown in other taxa ( Soltis et al. 2006 ). Larger amounts of sampling would be needed to determine this. If it is recognized as a separate taxon, it would likely be morphologically cryptic relative to T. fruticosum . To increase taxonomic and nomenclatural stability, the type specimen for this species was selected from a population that always fell within the strongly supported clade of T. fruticosum near the center of the species’ range. Additional specimens examined ( paratypes ): — U.S.A. Alabama , Baldwin County : Montrose , 9 October 1967 , Kral 29784 ( NCU !) ; Florida : Florida, 1897, Chapman s.n. ( NCU !); Alachua County : San Felasco Hammock State Park , 29.715116 N , 82.460713 W , 7 September 2019 , Abair 424 ( NCU !); outer zone-Devil’s Millhopper ; Gainesville , Florida , 4 October 1931 , Arnold s.n. ( FLAS !); Devil’s Millhopper Geologic State Park , 29.705578 N , 82.39521389 W , 29 June 2018 , McClelland & Schoonover 150 ( NCU !); 5 miles N of Alachua in Section 22, T7 S, R18 E on Townsend Branch of Mill Creek. , 29.865622 N , 82.50574 W , 3 November 1983 , Simons & Hall s.n. ( FLAS !); Bay County : St. Andrews , s.d., Beck 154 ( FLAS !); Charlotte County : 4.5 mi SE of Sarasota / Charlotte Co. line on US 41. Coco Plum Waterway ; canal and surrounding area., 22 August 1985 , Thompson & Nishida 2794 ( CM !, USCH !); Citrus County: “The Cove”, on Lake Tsala Apopka , 7 mi. n.e. of Floral City , 25 October 1966 , Beckner 1576 ( NCU !); Fort Cooper State Park : E edge of lake; SE 1/4 SW 1/4 S27 T19 S R20 E , 28.802238 N , 82.299104 W , 20 November 1999 , Hattaway FC 0050 ( USF !); McGregor Smith Scout Reservation , 15 September 1983 , Hopkins 1859 ( FTU !); Dixie Shores Trail , Crystal River , 28.878477 N , 82.614741 W , 16 October 2021 , McClelland & Schoonover 219 ( NCU !); Hollins zone 16. Crystal River State Buffer Preserve , 6 October 2000 , Scanlon 17 ( FLAS !); Collier County : in pastureland near seasonal lake, Kissimmee Billy , north of Alligator Alley. , 2 May 1976 , Smiley s.n. ( FTG !); Duval County : Vicinity of cabin; Roosevelt Wildlife Preserve , 7 October 1975 , Harrison & Garren 65 ( FLAS !); Roosevelt Wildlife Preserve -- 200 yards inside main gate., 25 September 1976 , Harrison & Garren 73 ( FLAS !); Little Talbot Island State Park , 30.457417 N , 81.41862778 W , 29 September 2018 , McClelland 183 ( NCU !); Timucuan Ecological and Historic Preserve - Theodore Roosevelt, 30.369639 N , 81.48503611 W , 30 September 2018 , McClelland 186 ( NCU !); Escambia County : At the E edge of Pensacola , off the E side of Scenic Highway , at the Bayview Memorial Cemetery , 20 November 1983 , Wilhelm 11857 ( USF !); Franklin County : near Apalachicola , Florida , 11 August 1876 , Anonymous s.n. ( US !); St. Vincent Island , Tahiti Point ( NE end)., 22 September 1983 , Christman s.n. ( FLAS !); St. James Bay Golf Course , just N of US 98, ca. 6 mi E of Carrabelle. , 10 October 2005 , Hansen 13067 ( USF !); near the Gulf of Mexico , at Alligator Point , 22 August 1967 , Henderson 67-1477 ( GA !, NCU !, USCH !) ; St. Vincent Island , 30 October 1910 , McAtee 1744 A ( US !) ; St. George Island State Park , 29.692044 N , 84.78745833 W , 14 July 2018 , McClelland & Schoonover 158 ( NCU !); Intersection of Alligator Drive and Bald Point Road , 29.901653 N , 84.348771 W , 17 October 2021 , McClelland & Schoonover 223 ( NCU !); Bald Point State Park , 29.910328 N , 84.36224444 W , 14 July 2018 , McClelland & Schoonover 157 ( NCU !); ca. 1/ 2 mi. E . of St Teresa and next to Camp Weed on US 98., 18 October 1969 , Sullivan 1302 ( NCU !); Gulf County : on road to Cape San Blas. , 11 September 1977 , Crewz 1194 ( FLAS !, USF !); Hardee County : Charlie Creek. , ca. 7 miles E of Zolfo Springs , 23 July 1949 , Brass 20612 ( FLAS !, US !); Cattle Range Station , Limestone , 1 July 1942 , Kirk s.n. ( FLAS !); CF Properties , USF Archaeological Site #8 Hr 11. Orchard-fenceline site, 10 mi W of Bowling Green. , 24 July 1977 , Kuczynski s.n. ( USF !); in Payne Creek State Historic Site. 2 mi. SE of center of Bowling Green , on S side of Rd. 664 A , W of Peace River , 15 June 1978 , Willson 213 ( FTU !); Hendry County : Rt. 80, LaBelle , 8 March 1950 , Hood 3442 ( FLAS !); Hernando County : s.w. side of Chinsegut Hill , ca. 3 mi. ne.e. of Brooksville , ne.e. corner of S3 , T22 S , R19 E , 28.605294 N , 82.385651 W , 25 September 1965 , Beckwith s.n. ( FLAS !); McCarty Woods , ca. 0.5 mi. N of Brooksville. , June 1989 , Krofta s.n. ( USF !); McCarty Woods , ca. 0.5 mi. N of Brooksville. , 2 October 1986 , Krofta s.n. ( USF !); northwest of Brooksville , off US 98., 22 September 1963 , Lakela 26478 A ( USF !); Hillsborough County : Wilderness Park , on Morris Bridge Road , near the Hillsborough River , 3 September 1989 , DeVore 1013b ( MU !); Along Patterson Rd. near Odessa. , 2 July 1963 , Ducey 49 ( USF !); Hillsborough River State Park : N 1 /2, SW 1/4, SE 1/4, NW 1/4, S8 , T27 S , R21 E ; Burn Zone HR-14; along Baynard Nature Trail , 28.15075 N , 82.231469 W , 7 July 1994 , Graham HR0049 ( USF !); Hillsborough River State Park ., 2 August 1966 , Lakela & Almeda 30121 ( USF !); Beacon Key study site #12. Sec. 34 T32 S R18 E … east of U.S. 41, 27.67392 N , 82.53593 W , 31 July 1974 , Todd 282 ( USF !); Lake County : 4 miles south of Leesburg , 20 October 1968 , Baltzell 697 ( FLAS !); Between Eustis & Leesburg Florida , 1 October 1933 , Buswell s.n. ( FTG !); S . edge Lake Dora , 1 November 1978 , Daubenmire & Daubenmire s.n. ( USF !); Eustis, Trout Lake Nature Center, 26 August 1989 , Daubenmire & Daubenmire s.n. ( USF !); Eustis, Trout Lake Nature Center, 19 November 1989 , Daubenmire & Daubenmire s.n. ( USF !); 3 mi. SSW of Leesburg. , 15 September 1990 , Daubenmire & Daubenmire s.n. ( USF !); Leesburg-Flat Island, 9 January 1991 , Daubenmire & Daubenmire s.n. ( USF !); Flat Island , SW of Leesburg , 15 November 2017 , Daubenmire & Daubenmire s.n. ( USF !); Entrance road to Flat Island Reserve , 28.781050 N , 81.892782 W , 16 October 2021 , McClelland & Schoonover 215 ( NCU !); Lee County : [Fort] Myers , August 1900 , Hitchcock 279 ( US !); Estero Bay Preserve , Estero River Scrub Unit , near Mullock Creek , 26.46727788 N , 81.8360699 W , 14 October 2021 , Horn s.n. ( NCU !); Salvista , Lee Co. , Fla. , 23 April 1930 , Moldenke 1075b ( DUKE !); Levy County : Waccasassa Bay State Preserve; Dewey Allen Park Road , northwest of junction with C-40 & C-40 A ; roadside., 29.036667 N , 82.733333 W , 30 September 1996 , Abbott 9600 ( FLAS !); Vicinity of Cedar Key and along highway SR . 24 between Cedar Key and Otter Creek , 15 September 1971 , Carlton , Sreemadhavan , Lakela & Long 3348 ( USF !); second island off the mouth of the Withlacoochee River ., 16 September 1959 , Cooley, Eaton & Ward 6958 ( USF !); Sea Horse Key near Cedar Keys, 16 November 1952 , Davis s.n. ( NCSC !); Cedar Key Train Trestle Nature Trail, 29.14357 N , 83.032334 W , 16 October 2021 , McClelland & Schoonover 220 ( NCU !); North Key : an island in Cedar Keys National Wildlife Refuge ; 2 mi W of Cedar Key. , 29.12672 N , 83.08562 W , 8 April 2004 , Sprenkle & Judd 318 ( FLAS !); Gulf Slope of Seahorse Key , 2 mi. s.w. of Cedar Key , 1 December 1962 , Ward 3254 ( FLAS !); s. side of Fla. 40, w. of Yankeetown , ca. 1 mi. from end of road, 26 March 1962 , Ward & Tomlinson 2986 ( FLAS !, USF !); Liberty County : at clearing above the Apalachicola River at Alum Bluff overview (ca. 2.0 mi N of Bristol )., 30.46389 N , 84.98693 W , 12 November 2012 , Anderson 26803 ( NCU !); at top of Alum Bluff , ca. 3 mi. n. of Bristol , S24 , T1 N , R8 W , Liberty County. , 10 November 1963 , Ward & Ford 3731 ( FLAS !, NCU !); Manatee County : Bradentown , 25 June 1923 , Cuthbert s.n. ( FLAS !); At Palmetto , 21 August 1895 , Nash 2452 ( FLAS !, US! ); Marion County : near lake, Fla 42, 0.5 mi. e. of Linadale , 30 September 1973 , Baltzell 5360 ( NCU !); Silver Springs State Park , 29.201458 N , 82.0389 W , 7 October 2018 , McClelland 191 ( NCU !); Mockernut Hill Botanical Garden , 29.454400 N , 82.30558889 W , 28 June 2018 , McClelland , Schoonover & Duever 148a ( NCU !); Mockernut Hill Botanical Garden , 29.454958 N , 82.30566944 W , 28 June 2018 , McClelland , Schoonover & Duever 148b ( NCU !); Nassau County : Amelia City , 20 November 1941 , Knight s.n. ( FLAS !); Fort Clinch State Park , 30.702975 N , 81.45328889 W , 30 September 2018 , McClelland 185 ( NCU !); Okaloosa County : Fred Gannon Rocky Bayou State Park , 30.498422 N , 86.42638333 W , 4 August 2018 , McClelland 166 ( NCU !); Henderson Beach State Park , 30.384908 N , 86.44789722 W , 4 August 2018 , McClelland 165 ( NCU !); Orange County : At Clarcona , 25 September 1899 , Meislahn 63 ( US !); Pasco County : Werner-Boyce Salt Springs State Park , 28.291022 N , 82.71776944 W , 24 June 2018 , McClelland & Schoonover 137 ( NCU !); Werner-Boyce Salt Springs State Park : Directly N of park shop, 5 September 2002 , van Hoek & Petty WB128 ( USF !); Polk County : Colt Creek State Park ; ca. 2.5 km SSE of FL 471 bridge over the Withlacoochee River , ca 5.1 km NNE jct. FL 471 and US 98., 28.293972 N , 82.045306 W , 26 September 2008 , Kunzer , Dauphin & Eckert 2530 ( USF !); Colt Creek State Park , 28.277592 N , 82.03106944 W , 24 June 2018 , McClelland & Schoonover 139 ( NCU !); Santa Rosa County : Gulf Islands National Seashore - Naval Oaks , 30.364872 N , 87.13128889 W , 5 August 2018 , McClelland 167 ( NCU !); Sarasota County : Sarasota , off Linwood Avenue just south and east of junction with Vinson Street. , 25 November 1995 , Abbott 8108 ( FLAS !); Oscar Scherer State Park : Head of Green Trail near W edge of campground, 6 August 2003 , Bishop OS0059 ( USF !); 3210 Old Oak Drive , Sarasota, FL , 23 September 1978 , Dodson 4836 ( USF !); Churchill Ranch on Border Rd. , part of Deer Prairie Creek Preserve, ca. 2.3 km ENE of jct. I-75 and C-777 ( River Rd ), 27.1175 N , 82.327778 W , 23 May 2007 , Franck 151 ( USF !); Off Bahia Vista Rd. down a dirt road., 11 February 1976 , Gordon & Mack 21 ( USF !); Oscar Scherer State Park , 27.170161 N , 82.47661667 W , 8 June 2018 , McClelland 131 ( NCU !); Seminole County : my home on Hermits Trail ; Altamante Springs , 25 October 1963 , Schallert 521 ( FTU !); St. Johns County : T6 S , R29 E , S39 ; 13 miles N. of St. Augustine and 15 miles S . of jacksonville Beach off SR A1 A in Guana River State Park . Approximately. 25 mile W . of park gate at Guana River Dam. North side of road., 23 September 1990 , Herring & Herring 138 ( FLAS !); south of Moses Creek and west of Matanzas River ; Moses Creek Conservation Area ;. 28 miles north of SR 206 and 2.2 miles east of US 1, 16 September 2003 , Slaughter 14440 ( FTG !); Sumter County : 6 mi. w. of Leesburg , 1 November 1967 , Baltzell 177 ( FLAS !); Walton County : Eden Gardens State Park , 30.370883 N , 86.12180278 W , 3 August 2018 , McClelland 164 ( NCU !) ; Georgia , Camden County : Cumberland Island National Seashore , 30.909086 N , 81.41468611 W , 1 October 2018 , McClelland & Neilan 188 ( NCU !); Little Cumberland Island : 1.15 miles from the lighthouse on the north end of the island on a 166 degree azimuth., 1 October 1971 , Richardson 45 ( MU !); Mississippi , Harrison County : Biloxi , Miss. , 19 September 1899 , Tracy 4905 ( CM !, NCU !); Jackson County : P . O . Ocean Springs , 21 August 1952 , Demaree 32960 ( NCU !); P . O . Ocean Springs , 17 October 1954 , Demaree 36229 ( NCU !, WILLI !); Gulf Islands National Seashore - Ocean Springs , 30.391514 N , 88.79065278 W , 6 August 2018 , McClelland 169 ( NCU !); about 2 miles east of Ocean Springs , coast road., 7 October 1950 , Whitehouse 23907 ( NCSC !) .