Sapindus standleyi A.R. Franck

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 : 47-49

9. Sapindus standleyi A.R. Franck , sp. nov. (LSID: 77340323-1) ( Figures 2 and 15 ).

Type:— NICARAGUA. Jinotega: vicinity of Finca San Roque, sierra east of Jinotega, 5 July 1947, P. C . Standley 10855 (holotype, EAP53400 !; isotype, F1263981 !) .

Diagnosis:— Sapindus standleyi is similar to S. saponaria subsp. saponaria and S. thurstonii . The distinctive characters of S. standleyi are the unwinged to scarcely winged rachises, at least on mature trees (vs. unwinged to winged in S. saponaria subsp. saponaria and unwinged except for seedlings in S. thurstonii ), usually subsessile leaflets with at least some blade tissue decurrent to the rachis (vs. often distinctly petiolulate in S. saponaria subsp. saponaria and often subsessile in S. thurstonii ), leaflet blades lanceolate to ovate-lanceolate, typically straight to lightly falcate, 2.2–4 times longer than wide with more acute to gradually acuminate apices (vs. usually lanceolate, ovate, elliptic, to obovate, 1.8– 3.5 times longer than wide, often strongly falcate-arcuate, with often more abruptly acuminate to gradually acuminate apices in S. saponaria subsp. saponaria and ovate to ovate-lanceolate, 2.5–3.5 times longer than wide with often falcate, gradually acuminate apices in S. thurstonii ), drying dark green, dark purplish green, to dark brown (vs. mostly light green to green in S. saponaria subsp. saponaria and mostly brown-green, purplish or reddish brown, to pale brown in S. thurstonii ), adaxially the midrib brownish pale green, reddish brown, to dark brown (vs. usually pale green to pale yellow in S. saponaria subsp. saponaria and pale yellow to brownish yellow in S. thurstonii ), abaxially the veins drying pale yellow to slightly brownish (vs. pale green to pale yellow in S. saponaria subsp. saponaria and pale yellow to yellowish green in S. thurstonii ), flowering usually late-spring–summer, rarely October (vs. flowering late summer–early spring in S. saponaria subsp. saponaria in the northern hemisphere and flowering late summer–early fall in S. thurstonii ), and restricted to 670+ m (vs. from 0–2000 m, more common below 1000 m, infrequent above 1000+ m for S. saponaria subsp. saponaria and 900–1400 m in S. thurstonii )

Etymology:—The species epithet honors Paul C. Standley, who provisionally named it as a new species on a specimen label. Over 20 years before he collected the type of S. standleyi, Standley (1923) had published the opinion that S. saponaria was a variable species, and that S. drummondii and S. marginatus were synonyms of it. Standley ( Williams 1963) was certainly familiar with S. saponaria , having collected it at least twelve times from 1922–1949 in Costa Rica, El Salvador, Guatemala, Honduras, Panama, and Florida, USA (see specimens cited under S. saponaria subsp. saponaria ) and he had identified the species from the collections of others. On 5 July 1947, he made two collections of S. standleyi , one sterile and one with flower buds. On his labels, Standley gave it the unpublished name “ Zanthoxylum collinum”, nom. inval. On the label of the sterile specimen at F (Standley 10869), the authorship of this unpublished Zanthoxylum name was credited as “Standl. & Steyerm., sp. nov.” and on the other specimens at EAP and F the labels bore the same but with “Steyerm.” scratched out and replaced with the handwritten “L. Wms.” (L.O. Williams). The one with flower buds at EAP (Standley 10855) bears the note “ Type collection” and the duplicate at F is identified as a “ Type ”. Another sterile specimen assigned here to S. standleyi was collected by Standley in 1940 (Standley 77387), and it was originally identified as S. saponaria . Williams had also collected S. standleyi in Nicaragua, identified as S. saponaria by his colleague J.A. Molina Rosito, and another collection a day later was misidentified as Alfaroa williamsii (Molina R. 1968: 357) .

Description:—Tree, to 21 m tall, bark finely roughened, becoming irregularly platy and slightly flaky. Petiole 3–8 cm long, pubescent to glabrate, unwinged to scarcely winged, mostly uniformly pale yellow to pale brown, sometimes with brown speckling; rachis pubescent to glabrate, unwinged; leaflets 6–17; petiolule 0–3 mm long, pubescent to glabrate; leaflet blade lanceolate to ovate-lanceolate, asymmetric to symmetric, usually falcate, the apex acuminate with an acute to obtuse tip, 5–14 cm long, 1.5–4 cm wide, 2.5–4 times longer than wide, adaxially drying dark green, dark purplish green, to dark brown, glabrous, midrib brownish pale green, reddish brown, to dark brown, occasionally pale yellow towards the base, at mid-blade the ridge 0.1–0.3 mm tall, 0.1 mm wide, the blade often somewhat sunken along the midrib, secondary veins pale yellow to reddish pale yellow or brownish pale green, tertiary and quaternary venation prominent, pale yellow to brownish pale green, discolorous from the blade surface, quaternary venation becoming obscure, blade abaxially drying light green to light brown, glabrous to densely pubescent or tomentose, midrib drying reddish brown to pale orange-yellowish, secondary veins pale yellow to slightly brownish, tertiary and quaternary venation prominulous to nearly obscure, pale yellow, discolorous to nearly concolorous with the blade surface, quinternary venation obscure to unapparent; foveolae inconspicuous to conspicuous on the abaxial leaflet blade surface, exudate clear. Petal lacking appendages. Mature mericarp 19–23 cm wide, pericarp 0.5–2 mm wide, seed 13–17 mm wide.

Distribution, habitat, and phenology:—Eastern Mexico (Chiapas, Guanajuato, Michoacan, Oaxaca, Queretaro, Tamaulipas, and Veracruz) to northwestern Nicaragua (Esteli, Jinotega, Matagalpa, and Nueva Segovia) ( Fig. 2 ); 670–1800 m. Cloud forest, montane forest, and pine-oak forest. Flowering May–July (rarely October, also August, October, November, and January in cultivated specimens in southern California).

Notes:— Sapindus standleyi is restricted to high elevations (670+ m, usually above 1000 m), and it is distinguished by its falcate leaflets that dry to a dark coloration adaxially with usually reddish to brownish venation and prominent reticulate veins on the upper surface. This species predominantly flowers in the late spring–early summer (supported by 15 collections), yet two collections from October in southern Mexico are in early inflorescence (Jerónimo 31 & Cedillo & Calzada 42). Given the stark difference in habitat and flowering period, Sapindus standleyi is probably completely reproductively isolated or nearly so from S. saponaria .

The distribution of S. standleyi is consistent with an area of endemism in montane habitats identified by SánchezGonzález et al. (2008) from the Sierra Madre Oriental, Mexico to northern Nicaragua. This area is also similar to a particular distribution pattern of Neotropical cloud forests (pattern F in Luna-Vega et al. 2001) and the distribution for the Mexican and Central American populations of Liquidambar styraciflua ( Linnaeus 1753: 999) ( Martin & Harrell 1957, Luna-Vega 2008, Stull 2023). Montane frugivores that are large enough to feed on Sapindus fruits and have somewhat similar distribution patterns as S. standleyi include the emerald toucanet ( Wheelwright 1985), cracids ( Muñoz & Kattan 2007), and the Central American red brocket deer (Gallino-Tessaro et al. 2019).

The flowering specimen of Karwinski collected from Zimapan, Mexico in July that was cited by Radlkofer (1932a) under S. marginatus may belong to S. standleyi . The description of Sapindus in Rzedowski & Rzedowski (2006) concerning the Bajío region of central Mexico pertains mostly to S. standleyi . Although Rzedowski & Rzedowski (2006) report the elevation as low as 250 m, this more likely corresponds to specimens of S. saponaria subsp. saponaria . It was also reported by Rzedowski & Rzedowski (2006) that the petals bear appendages, but I have only seen tufts of hairs consistent with the illustration in Rzedowski & Rzedowski (2006).

Unexpectedly, this species is in cultivation in southern California, and the leaflets are strongly tomentose, very similar to specimens from the area of San Sebastián de las Grutas, Oaxaca, Mexico. According to one label (Moran 2452), Franceschi planted this species at a private residence in Montecito, Santa Barbara County. This could refer to Francesco Franceschi, who resided in California from 1893–1913, or perhaps his sons who oversaw his nursery after 1913 ( Tucker 1943). Francesco Franceschi had introduced numerous species to southern California, corresponding with people from numerous countries, including Oaxaca, Mexico ( Franceschi 1904 –1918). These cultivated specimens in southern California flowered in June, August, October, November, and January (based on SBBG herbarium specimens cited below).

Conservation Status:—The species has a relatively large extent of occurrence, distributed at high elevations from western Nicaragua to northeastern Mexico. The species should be classified as Least Concern ( IUCN 2012). However, the potential for continuing deforestation and fragmentation could threaten the loss of local subpopulations.

Specimens examined:— EL SALVADOR. Guatemala: Guatemala, Zona 1, Cerrito del Carmen , 14.646164°, - 90.505398°, 19 August 2022, Eufragio et al. AEB310 ( USCG) . Santa Ana: San José Ingenio, P. N. Montecristo , 1600 m, 14° 25’ N, 89° 21’ W, 3 October 2001, Martínez 245 ( MEXU) . GUATEMALA. Chimaltenango: volcán Acetenenago, Aldea Quisache , 19 May 2004, Véliz 15269 ( MEXU) . Escuintla: 11 km N of Palin , 4 July 1970, Harmon & Dwyer 2889 ( BRIT [ex NLU]) . Huehuetenango: between Democracia and Santa Ana Huista, Sierra de los Cuchumatanes , 25 August 1942, Steyermark 51303 ( F) . Jalapa: vicinity of Jalapa, elev. 1360 m, 7–18 November 1940, Standley 77387 ( US) . Quiché: Sacapulas , 15° 16’ 51” N, 91° 5’ 59” W, 18 June 2006, Véliz & Méndez 16966 ( F) . Sacatepéquez: Antigua Guatemala, Hotel Antigua , 22 May 1996, Véliz 96.5539 [cultivated] ( MEXU) . MEXICO. Chiapas: near Siltepec , 2000 m, 6 July 1941, Matuda 4344 ( F, MO, US) ; Zacatonal, Acacoyagua , 10 September 1948, Matuda 18345 ( F) . Guanajuato: Carricillo, 10 km al SE de La Joya , Atarjea Mpio. , 1800 m, 5 Jan 1989, Ventura & López 6531 ( MEXU) . Michoacan : cerca de Santiago, mpio. de Tzintzuntzan, 2100 m, 24 June 1986, Díaz Barriga 2334 ( MEXU) ; ibid., 21 June 1988, Díaz Barriga 4709 ( MEXU) . Nuevo Leon: Sierra de la Silla, frente a El Mezcal , Cadreyta Jiménez , N. L., 670 m, 28 May 1978, Moya Rdz. 113 ( MEXU) . Oaxaca: Mpio de San Sebastian Las Grutas, km 10 de la carretera Vado-Tlacuache , 20 May 1978, Solano 158 ( F) ; El Rincón Tlapacaoyán, Distr. Zimatlán , 1350 m, 23 June 1979, Sousa 10451 ( MEXU) ; San Sebastian de las Grutas, Niñero La Cadena , 1750 m, 13 November 1981, Avila s.n. ( MEXU) ; San Sebastián, 11 km al W del entronque de la carr. Oaxaca-Pt. Escondido con la brecha à San Sebastián , distr. de Sola de Vega , alt. 1650 m, 16 May 1982, Torres C. et al. 447 ( MO) ; Cañada de San Jerónimo Coatlán, Mpio San J. Coatlán , Dto. Miahuatlán , 28 June 1990, Campos V. & Toriz 3184 ( MEXU) ; San Cristóbal Amatlán , 28 October 1994, Jerónimo 31 ( USF) . Queretaro : La Venta, 21 April 1979, Arguelles 1235 ( LAF) ; 9 km al NE de El Humo, Mpio. Landa de Matamoros , 850–1000 m, 24 May 1989, Zamudio R. 7312 ( MEXU) ; La Curva, 1.5 km al N de el Barrioa de la Luz, Mun. de Landa , 1220 m, 6 June 1989, González 651 ( MEXU) ; 2 km al SE de Neblinas, Mpio. Landa , 900 m, 23 June 1989, Rubio 828 ( MEXU) ; 2–3 km al norte de La Parada, municipio de Jalapan (1100 m), 2 May 1990, Servín 174 ( F) . Tamaulipas: Mpio. Gomez Farias, Sierra de Guatemala , Rancho Cielo Biological Stations , 4 June 1971, Sullivan 293 ( TEX) ; Mpio Gómez Farías, Rancho del Cielo , 1150-1200 m, 18 September 1971, González-Medrano et al. 3579 ( MEXU) ; 7 mi. NW of Gomez Farias, Rancho del Cielo , 23 July 1973, Lonard 3766 ( PAUH) ; 4 km al sur de Torrecillas, 660 m, 30 May 1974, Medrano 7244 ( MEXU) ; Sierra de Gomez Farias , 27 June 1988, Cházaro B. et al. 5507 ( MICH) ; Reserva de la Biósfera Rancho del Cielo , 27 June 1988, Vázquez et al. 4607 ( F, USF) . Veracruz: Carretera a Sontecomapan Ejido Xovi , Rancho Cuencamo , 23 October 1972, Cedillo & Calzada 42 ( F, MEXU) ; Mpio. Las Minas, entre Las Minas de Somelahuacan y Molinillos , 22 August 1985, Chagaro & Acosta 3698 ( F) . NICARAGUA. Esteli: Municip. de Esteli, Reserva Natural Miraflor, Comarca Puertas Azules , 800–1300 m, 14 May 1999, Rueda & Velásquez 11157 ( MO, P) ; Esteli, Reserva Natural Miraflor, sur de El Cebollal , 1200-1350 m, 12 June 1999, Rueda et al. 11402 ( MO) . Jinotega: vicinity of Finca San Roque , sierra east of Jinotega, 5 July 1947, Standley 10869 ( EAP, US) ; Ocotillo near Sta. Lastenia , 17 January 1965, Williams et al. 27851 ( EAP, F, US) ; 10 km al SE de Tepesomoto camino Tepesomoto Pueblo Nuevo en el Potaste , 28 August 1982, Martínez S. & Grijalba 1771 ( MEXU) . Matagalpa: Matagalpa , 3500 ft., 10 April 1957, Salas & Taylor 2386 ( EAP) ; Finca Sta. María de Ostuma, Cordillera Central de Nicaragua , 1400 m, 18 January 1965, Williams et al. 27983 ( EAP) ; NW slope of Cerro El Picacho, 1420–1520 m, 25 May 1983, Stevens 22126 ( BRIT, MEXU, MO, US) . Nueva Segovia: Loma Fria-ridge west of Rio Las Manos , 20 km NW of Ocotal, 1200 m, 16 June 1977, Neill 2203 ( MO) . USA. California. Santa Barbara County: 9 High Rd., Montecito , 25 February 1945, Ryon 224 ( SBBG) [cultivated]; Harrison Ryon place, 9 High Road, Montecito , 15 August 1947, Moran 2452 ( SBBG) [cultivated, allegedly planted by Francesco Franceschi]; West Victoria Street, between San Andreas and Chino , 4 June 1957, Hopkins & Smith s.n. ( SBBG) [cultivated]; Victoria Street nera Chino Street, Santa Barbara, 22 October 1957, Hopkins s.n. ( SBBG) [cultivated]; along Alisos between Cota and De la Guerra and between Montecito and Gutierrez Streets , 7 January 1966, Muller & Broder M2208 ( SBBG) [cultivated]; 700 block of West Victoria, Santa Barbara , 29 November 1971, Muller s.n. ( SBBG) [cultivated]; 700 block of Alisos Street , 24 June 2004, Muller 04-428 ( SBBG) [cultivated]; Ortega Street , 3 November 2023, Guilliams 8537 ( FLAS) .

Figures

FIGURE 2. Morphology of various species of Sapindus. Top-left: seedlings of S. mukorossi (Franck 5932, FLAS), top-middle: seedling of S. saponaria subsp. saponaria (Wilder et al. 31737, SWF; photo by K. Relish), top-right: foliaceous cataphyll at base of newly emerging shoot of S. drummondii var. glabratus (Trelease 60, MO), bottom-left: exudates (e.g. white arrow) of glandular foveolae on abaxial leaflet surface of S. standleyi (Neill 2203, MO), and bottom-right: exudates (e.g. white arrow) of glandular foveolae on abaxial leaflet surface of S. mukorossi (Yonekura 96562, A).

FIGURE 15. Sapindus standleyi, clockwise from top-left: fertile branch (Stevens 22126, image courtesy of The Missouri Botanical Garden Herbarium, MO, accessed on Tropicos® [10 August 2023]), petal adaxial surface (Torres C. et al. 447, MO), leaflet adaxial surface (Véliz & Méndez 16966, F), leaflet abaxial surface (Véliz & Méndez 16966, F), and petiole base (Torres C. et al. 447, MO).