Sapindus marginatus Willdenow (1809: 432)

Franck, Alan R., 2024, Revision of Sapindus sect. Sapindus (Sapindeae, Sapindoideae, Sapindaceae), including the description of three new species, Phytotaxa 648 (1), pp. 1-71 : 22-25

3. Sapindus marginatus Willdenow (1809: 432) ( Figure 8 ). Lectotype (designated here, or perhaps holotype):— USA. “Carolina” (lectotype, B -W07740 -01 0!). = Sapindus falcatus Rafinesque (1830: 261) . Neotype (designated here):— USA. “Carolina” (lectotype, B -W07740 -01 0!). = Sapindus acuminatus Rafinesque (1838: 22) , nom. illeg. (non S. acuminatus Wall. ex Royle 1835 ). Lectotype (designated here):— USA. Florida: ex herb. Durand, ex herb. Darlington (DWC!). = Sapindus manatensis Shuttlew. ex Radlkofer (1878: 318 , Tab. II: nos. 35 & 38, 400). Lectotype (designated here):— USA. Florida: Manatee County: Ad ostia fl. Manate, Florida austr. occ., July 1845, F.I.X. Rugel 115

( lectotype, G00096035 !; isolectotypes, BM000838047 !, BRIT548367 ! [ex VDB], BRIT548368 ! [ex VDB], GH00050889 !,

K000701891!, MEXU1003156!, NA0217604!, P02297184!, P02297185!, PH 00022285!, PH 00022286!)

Etymology:—For the usually marginate or unwinged rachis, probably to distinguish it from S. saponaria which can sometimes have distinctly winged rachises.

Description:—Tree, to 24 m tall, bark texture appearing somewhat smooth but finely roughened on the surface, developing shallow fissures, becoming somewhat flaked or platy into longitudinal strips. Petiole 1–8 cm long, glabrous to sparsely pubescent, usually unwinged or scarcely winged, rarely the wing to 3 mm wide on one side; rachis glabrous to sparsely pubescent, unwinged or wing to 3 mm wide on one side; leaflets 6–14(16); petiolule 0–3 mm long, glabrous to sparsely pubescent; leaflet blade lanceolate to ovate-lanceolate, asymmetric to nearly symmetric, straight to falcate, the apex subtly acuminate with an acute to obtuse tip, 4–16 cm long, 1.5–4 cm wide, 3–6 times longer than wide, adaxially drying green to brownish green, glabrous or the midrib glabrous to sparsely pubescent, midrib pale green, 0.1–0.2 mm high, 0.1–0.2 mm wide, the blade usually not sunken along the midrib or only slightly so, secondary veins pale green, tertiary venation pale green, slightly discolorous from the blade surface, quaternary venation prominulous, nearly concolorous with the blade surface, quinternary venation obscure, abaxially drying pale green to somewhat glaucescent, glabrous to sparsely pubescent, midrib pale green to pale orange, secondary veins pale green, prominent, tertiary venation pale green, slightly discolorous from the blade surface, quaternary venation prominulous, nearly concolorous with the blade surface, quinternary venation obscure; foveolae inconspicous on the abaxial leaflet blade surface, exudate clear. Petal with 2 appendages, the appendage 0.5–1.5 mm long. Mature mericarp subglobose, 14–18 mm wide, pericarp 0.5–2 mm wide, seed 9–12 mm wide.

Distribution, habitat, and phenology:—Southeastern USA (Alabama [ Lelong 1988], Georgia, Mississippi, South Carolina, and Florida) ( Fig. 3 ), 0– 50 m. Calcareous woodlands, coastal hammocks, and shell mounds. Flowering April–June.

Notes:— Sapindus marginatus is characterized by its winter-deciduous leaves, foliaceous cataphylls, glabrous to sparsely pubescent and elongate leaflets, its petals with appendages (illustrated in Sargent 1902), epigeal and phanerocotylar cotyledons, and its spring–summer flowering season. This species can form clonal patches of numerous above-ground stems connected through horizontal roots. It is often found in calcareous areas, and an analysis of soil for one specimen (Laessle M-57) from Florida showed a pH reading of 7.7–8.2 ( Laessle 1942). From observations in 2022–2024 in Alachua and Gulf counties, Florida, mature plants were nearly completely leafless by mid-January after experiencing freezing or near-freezing temperatures, and new leaves arose in March.

Sapindus marginatus shares some similarities with S. drummondii ( Brizicky 1963) and S. mukorossi , both of which are winter-deciduous, have foliaceous cataphylls, petals with appendages, and a spring–summer flowering period. The leaflets of S. marginatus are generally larger and more glabrous than those of S. drummondii var. drummondii , and the fruits are larger than S. drummondii . Epigeal germination is known for S. marginatus , and hypogeal and cryptocotylar germination occurs in S. drummondii . Sargent (1891) and Torrey & Gray (1840: 685) considered S. drummondii to be a synonym of S. marginatus , but later Sargent (1902) recanted and recognized them as separate species. In Georgia, USA, the name S. saponaria var. drummondii was misapplied to plants of S. marginatus (Jones, Jr. & Coile 1988) . Compared to S. mukorossi , S. marginatus has more narrowly elongate leaflets which lack the reddish leaf exudates, the leaflet midrib ridge is usually not as high, and the leaflet venation is usually not as whitish.

Many treatments had placed S. marginatus as a synonym of S. saponaria ( Pursh 1814, Nutall 1818, Radlkofer 1878, 1900, Standley 1923, Little 1979, Wunderlin et al. 1985, Wunderlin 1998, Wunderlin & Hansen 2011, Acevedo-Rodríguez 2014), while some kept them distinct ( Chapman 1878, 1889, 1897, Sargent 1902, Small 1903, West & Arnold 1946, Kurz & Godfrey 1962, Wunderlin 1982, Clewell 1985). LaMotte (1935) proffered that these two species “appear to intergrade” but offered no evidence. The flowering period of S. marginatus (April–June) is effectively separate from that of S. saponaria (September–March, rarely April, July, or August in the northern hemisphere). Even plants of S. saponaria in Florida cultivated within the range of S. marginatus still flowered in the fall (e.g. Spence s.n. and Williams s.n.). The cotyledons of S. marginatus are epigeal and phanerocotylar, and they are hypogeal and cryptocotylar in S. saponaria . In seedlings, the first true leaves of S. marginatus are compound, whereas they are simple in S. saponaria . Appendages are present on the petals of S. marginatus , and they are lacking in S. saponaria . The leaflets of S. marginatus are 3–6 times longer than wide, usually nearly glabrous, lack conspicuous foveolae, and usually the rachis is wingless or only narrowly winged. In S. saponaria , the leaflets are usually 1.8–3.5 times longer than wide, glabrous to pubescent, often with conspicuous foveolae, and the rachis can range from wingless to strongly winged. The southernmost native populations of S. marginatus are in central Florida (Sarasota and Highlands counties), USA, ca. 100 km north of the nearest native population of S. saponaria in Lee County, Florida. The allopatric ranges ( Brizicky 1963), nearly opposite flowering times, and distinctive morphologies fully support recognizing S. marginatus as a distinct species. This species was accepted by Radklofer (1932a).

Pursh (1814) and Little (1979) supposed the species was partly spread by humans, which is certainly plausible. The species has been found on several different shell mounds. Like nearly everywhere else where Sapindus occurs, S. marginatus was probably used by indigenous peoples, e.g. for soap and jewelry ( Austin 2004). Its distinctiveness as a species would favor it being present long before humans arrived within its distribution.

An early written account of S. marginatus comes from William Bartram (1791, Harper 1958) where he noted the species at Colonel’s Island (Georgia) in 1773, and in 1774 along the St. Johns River near Palatka (Florida) and the Alachua Sink (Florida). In May 1788, Michaux sighted this species in Florida at Mount Royal (Putnam County) and Silver Glen Spring (Marion County) ( Taylor & Norman 2002) and in Michaux’s flora he gives the locality as near the shore in Georgia ( Michaux 1803: 242, as S. saponaria ).

As explained in the preface of Willdenow’s Enumeratio Plantarum, S. marginatus was in cultivation in the summer of 1808 at the Berlin botanical garden. In the protologue of S. marginatus , the “T.” indicated it was cultivated in the open air in summer, and in a warm room in the winter. No explicit type is indicated in the protologue, and thus a lectotype from Willdenow’s herbarium is designated here. The lectotype chosen here has also been interpreted as the holotype ( Acevedo-Rodríguez 2014), although Willdenow also cited Michaux’s (1803) account.

Nuttall probably saw S. marginatus , perhaps with Baldwin’s advice, in the fall of 1815 along the “sea-coast” of Georgia ( Nuttall 1818, Pennell 1936, Graustein 1967: 100). Baldwin might have collected flowering material in southeastern Georgia or northeastern Florida in the 1810s ( Small 1921), as there is a specimen at PH from the Schweinitz herbarium but lacks additional data; Schweinitz had purchased the Baldwin herbarium but discarded the original labels ( Stuckey 1971). On this same sheet is a label “Florid. Nuttall” which possibly refers to the lone leaf at the bottom of the sheet and could have been collected during Nuttall’s visit to the Florida panhandle in March 1830 ( Nuttall 1834, 1841, Pennell 1936). Williams (1827) must have observed this species of Sapindus in Franklin County, Florida.

The 1828 catalogue of Bartram’s Garden (Philadelphia, PA) listed S. saponaria under the “Green-House Plants” ( Carr 1828), but this was more likely S. marginatus , since the Bartrams only explored northern Florida. In the protologue of S. acuminatus , nom. illeg. (= S. marginatus ), Rafinesque mentioned it was “seen alive in Bartram’s garden”. In the protologue of S. falcatus, Rafinesque cited Elliott (1817) which clearly refers to S. marginatus and cited Schoepf (1787: 59), who listed Carolina and the Bahamas for the distribution, but no Sapindus is known from the Bahamas. No original material of S. falcatus is known, and a neotype is designated. The type of S. manatensis was collected by Rugel in 1845 near the mouth of the Manatee River, probably on Snead’s Island ( Nash 1896). The lectotype of S. manatensis at G bears the annotation “! Radlk.” in Radlkofer’s hand.

Sapindus marginatus once occurred in coastal South Carolina. In the winter of 1784, in the vicinity of Charleston, South Carolina, Schoepf ( Morrison 1911: 172) mentioned the occurrence of Sapindus saponaria , which must refer to S. marginatus . There is one herbarium specimen (USCH) from South Carolina collected by Mellichamp from Bluffton (Beaufort County), ca. 50 km north of extant populations in Chatham County, Georgia. Further, the provenance of the type specimen was given as “ Carolina ” (though this could refer to an antiquated definition that included parts of Georgia and northern Florida). Thus, S. marginatus is presumed extirpated from South Carolina.

A tree at the Alachua Sink (Alachua County, Florida) was declared the State of Florida and USA National Champion, being 22 m tall with a trunk circumference of 2.2 m ( Ward & Ing 1997), but in 2014 a tree on the University of Florida campus in Gainesville (Alachua County, Florida) was declared the State Champion being 23.8 m tall with a trunk circumference of 1.8 m ( FDACS 2023, as S. saponaria ).

Conservation Status:—Although the extent of occurrence is rather large (> 50,000 km 2), the area of occupancy is quite small (<2,000 km 2). Each subpopulation is very localized, creating a fragmented population. With continuing deforestation within its extent of occurrence, the species should be categorized as Vulnerable ( IUCN 2012).

Specimens examined:— USA. Alabama: Baldwin County: SE shore of Tensaw River at confluence with Apalachee River , Blakeley Historic Site , 21 October 1977, Lelong 9992 ( USAM). Florida : East Florida, [probably along St. Johns River in Volusia County in June 1843, with flower buds and flowers], Buckley s.n. ( BRU, GH); [probably Jul–August 1843 with immature fruits], ex A. Gray , Buckley s.n. ( M); S. Florida, [1800s], Chapman s.n. ( FLAS). Alachua County : small rocky knoll at Great Sink , 20 June 1919, Harbison 98 ( A); Levy Lake , 19 September 1932, West 844 ( FLAS); Haufler’s Dairy pasture, Paynes Prairie , 3 June 1936, Pasture Survey 1936 s.n. ( DUKE, FLAS); NE of Alachua Sink , T10 S, R20 E, Sec. 22, S ½, 23 November 1981, Easterday 786 ( FLAS, GA, LAF); Paynes Prairie Preserve, 4 April 2009, Schlegel 278 ( BUF); University of Florida campus, 11 May 2022, Franck 5673 ( FLAS). Brevard County : Merritt’s Island, 11 July 1893, Baldwin 58 ( NY); Merritt’s Island near Cocoa, [s.d.], McFarlin 4929 ( MICH); 3 mi. N of Tropic, 5 October 1943, Rhoads s.n. ( FLAS); North Merritt’s Island, 24 May 1973, Shuey M0064 ( FTU, USF). Citrus County: about a mile NE of Homosassa, 24 May 1909, Harper 43 ( A, NY, US); Crystal River , 21 April 1922, Small s.n. ( FLAS, GH); north of Homosassa Springs , 10 April 1938, Baker s.n. ( FLAS); Pineola Fern Grotto, 18 May 1974, Lakela et al. 5309 ( FSU, USF); Pineola Grotto, 1 June 1975, Ward et al. 8831 ( FLAS, FSU). Columbia County : SE corner of Boston Rd & Montana Rd, T 6 S, R15 E, Sec. 26, NE ¼ of ¼, 27 December 1982, Burckhalter 98 ( UNA); O’Leno State Park , 5 June 1990, Tan 678 ( FLAS); Ichetucknee Springs State Park , 12 June 1993, Herring & Herring 1159 ( FLAS). Dixie County : Suwannee, 27 July 1962, Kral & Godfrey 15164 ( BRIT [ex VDB], FSU). Duval County : vicinity of Mayport and Jacksonville, 1870-1876, Keeler s.n. ( BRU); near Jacksonville, 6 April 1905, Moulié s.n. ( US); first island W of the inland waterway on the N side of the St Johns River , 25 December 1965, Blake s.n. ( GA, VSC); Fort Caroline National Monument and Timucuan Ecological and Historical Preserve, Willie Brown Trail, 1 July 2005, Giannasi et al. 1566 ( GA). Franklin County : St. Vincent’s Isle , West Florida, Chapman s.n. ( NY); N shore of St Vincents Island , 16 December 1975, Clewell 4856 ( FSU); just E of Thank You Ma’am Creek on left bank of East River , 9 October 1986, Anderson 10055 ( FSU); Apalachicola River , N side of East River , 16 April 1994, Nelson et al. 15410 ( FSU). Gulf County : T. H. Stone Memorial St. Joseph Peninsula State Park , 26 March 2024, Franck 6379 ( FLAS). Hernando County : Brooksville , 7 June 1938, Phillips et al. s.n. ( FTG); 1.5 mi. N of Chinsegut Hill , 18 May 1965, Ward & Beckner 4700 ( FLAS); along US 1, 5 mi. N of Brooksville , 21 November 1966, Clewell 2464 ( FSU); S side of US 98, W of junction of route 491 A, entrance to Brooksville Quarry, 31 December 1982, Leonard 7938 ( FSU). Highlands County: Highlands Hammock State Park , Ancient Hammock Trail , 9 May 1982, Popenoe & Roberts 2215 ( FTG). Hillsborough County : W of US 41, S of 74th Street, on the Manatee County line, 7 November 1986, Shuey 1803 ( USF). Lake County: Juniper Hunt Club, N of Juniper Run , 25 August 1982, Hopkins 1261 ( FTU). Levy County : Cedar Key, November 1898, Sargent s.n. ( A); Cedar Keys, 2 June 1901, Tracy 7482 ( GH, NY, US); Chambers Island , 4 May 1959, Wood, Jr. et al. 9221 ( A, GA, NCU, USF); Punkin Island , W of Yankeetown , 21 August 1971, Godfrey 70713 ( FSU); Cedar Key, 19 May 1984, Godfrey & Baker 81310 ( FLAS, FSU); Waccasassa Bay State Preserve, 21 June 1994, Judd 7034 ( FLAS); ibid., 22 September 1996, Abbott 9516 ( FLAS); North Key , Cedar Keys National Wildlife Refuge, 8 April 2004, Sprenkle & Judd 366 ( FLAS). Manatee County : Manatee, [ca. 1882-1890s], J. H. Simpson s.n. ( AUA); Sneads Island , 21-23 August 1895, Nash 2460 ( GH, MICH); ibid., 14 May 1900 [flowering], Tracy 6741 ( DUKE, GH, NY, US); Terra Ceia Island , 23 May 1921, Cuthbert s.n. ( FLAS); south of McMullen Creek , 5 November 1976, Shuey 1798 ( USF). Marion County : Silver River , S of US 40, 3 October 1984, Godfrey 81643 ( FSU); Silver Glenn Springs, Ocala National Forest, 11 July 1994, Miller s.n. ( FLAS); Silver River State Park , Silver Springs, 23 June 2006, Hubbard 555 ( FLAS). Okaloosa County: near Buccaroo Point , mouth of Boggy Bayou , 18 July 1968, Ward 6739 ( FLAS). Orange County : ( FLAS). Polk County : Green Swamp , T25 S, R23 E, Sec. 14, 22 November 1983, Post & Lopez s.n. ( USF). Putnam County : Conservation Reserve , Welaka, 16 June 1939, DeVall s.n. ( FLAS); University of Florida Conservation Reserve , Orange Point, 29 June 1939, Laessle M-57 ( FLAS); edge of St. Johns River , Orange Point, Welaka Conservation Preserve, 7 October 1961, Ward & Myint 2823 ( BRIT [ex VDB], FLAS, GH, NCU, US). Sarasota County : Historic Spanish Point, 13 June 2002, Holst 8042 ( SEL, USF). St. Johns County : Guana River State Park , 14 November 1993, Harrison & Harrison 624 ( FLAS, FTG); ibid., 14 May 1994, Harrison & Harrison 684 ( FLAS). Sumter County : hunting preserve 8 mi. NW of Bushnell, 24 April 1965, Wiggins 19811 ( FLAS, NCU); W of Lake Panasoffkee , 28 June 1984, Wunderlin & Hansen 9779 ( FTG, USF). Volusia County : Hontoon Island , 15 January 1971, Smith 17 ( FSU); ibid., July 1986, Newsom s.n. ( FLAS). Georgia: [Chatham /Liberty County: Ossabaw /St. Catherines] Island in the mouth of Midway [Medway] River, 1910, Julia King s.n. ( A); Hurricane Island, King s.n. ( A). Camden County: N end of Cumberland Island at Abraham Point , 23 April 1990, Krakow & Schinhofen 574 ( FTG, GA). Chatham County : Ossabaw Island , 6 July 1979, Tatum s.n. ( GA). Liberty County : St. Catherines Island , 23 August 1983, Jones et al. 23849 ( GA, TENN). McIntosh County : Sapelo Island along Duplin River , 16 July 1956, Duncan 20296 ( NCSC); Sapelo Island , 14 June 1974, Duncan 29065 ( FLAS). Mississippi: Jackson County : North Rigolets Island , 24 September 1981, McDaniel et al. 25518 ( FSU, GSW, MMNS); Bangs Island near Pt. Aux Chenes Bay , 24 September 1981, McDaniel et al. 25539 ( MMNS); Bangs Island at point which Bayou Cumbest enters into Pt Aux Chenes Bay , 15 May 1992, Wieland & Ruple 6637 ( MMNS); Bangs Island , Grand Bay National Estuarine Preserve, 23 April 1998, Sorrie & Wieland 9667 ( NCU); Bangs Island , 4.4 air mi. S of Orange Grove , 24 April 1998, Wieland et al. 8022 ( MMNS); Grand Bay NERR, 1 May 2004, Sullivan et al. 04-547 ( MMNS). South Carolina : Beaufort County : Bluffton, [1850s–1903], Mellichamp s.n. ( USCH) .

Figures

FIGURE 3. Native distribution of cited specimens of Sapindus in the Americas and eastern Oceania (cultivated or naturalized occurrences are not mapped). Longer names of some taxa are abbreviated, i.e. S. drummondii var. drummondii (abbreviated as drum. var. drum.), S. drummondii var. glabratus (drum. var. glabratus), S. saponaria subsp. jardinianus (sap. subsp. jard.), and S. saponaria subsp. saponaria (sap. subsp. sap.), The symbol for the occurrence of S. saponaria subsp. jardinianus in Hawaii is placed beside Hawaii on the map.

FIGURE 8. Sapindus marginatus, clockwise from top-left: fertile branch (Easterday 786, FLAS), adaxial side of petal with white arrow pointed at appendage (Franck 5673, FLAS), leaflet adaxial surface, leaflet abaxial surface, and petiole base (Pasture Survey 1936 s.n., FLAS).