Mahonia sagrana Harber & Bécquer, 2023

Harber, Julian & Bécquer, Eldis R., 2023, MAHONIA SAGRANA, A NEW SPECIES FROM CUBA, AND LECTOTYPIFICATION OF MAHONIA TENUIFOLIA AND BERBERIS FRAXINIFOLIA FROM MEXICO, Edinburgh Journal of Botany 80 (1961), pp. 1-17 : 4-14

Mahonia sagrana Harber & Bécquer , sp. nov.

Differs morphologically from Mahonia tenuifolia by its fewer leaflets (2–4 pairs vs 3–6 pairs), shorter distance to first pair of leaflets from base of petiole (2.7–4.5(–6) cm vs (3–)5.5–8(–10) cm), leaf texture coriaceous (vs papyraceous), shorter racemes (to 17 cm long vs to 37 cm long), and greater number of ovules ((3–)4–5 (vs 2)). – Type: Cuba, Pinar del Río Province, Sierra de los Organos, grupo del Rosario at Río Taco-taco, 12 x 1923, E.L. Ekman 17623 (holotype S [S16-43280!]; isotypes G [G00405854] image!, HAC!). Figures 1 , 2 , 3 , 4 .

Shrubs, evergreen, c. 3 m tall. Leaves imparipinnate, 16–29 cm long; leaflets 2–4 pairs, contiguous, lowest pair 2.7–4.5(–6) cm above base of petiole, abaxially and adaxially bright green, shiny, broadly ovate, elliptic-ovate or elliptic, 6–8.5(–9.5) × (2–)3.5–6(–7) cm, coriaceous; midvein raised abaxially, impressed adaxially, lateral veins inconspicuous on both surfaces, base attenuate; margin entire, slightly undulate; apex acute or acuminate. Inflorescence a single or 2–3(–5)-branched raceme to 17 cm, 25- to 40-flowered; pedicel 5–7 mm; floral bracts triangular ovate, 1.5 × 0.75 mm, apex acuminate. Flowers bright yellow, c. 5 mm in diameter; sepals in three whorls; outer sepals triangular ovate, 1 × 1 mm; median sepals broadly ovate, 2 × 1.75 mm; inner sepals broadly obovate to elliptic, 4–4.5

× 3–3.5 mm. Petals ovate-elliptic to elliptic, 4 × 2.5 mm, glands separate, apex obtuse. Stamens dentate, with teeth pointing upwards or downwards, 3 mm; anther connective not extended, truncate. Ovary 2.5 mm; ovules (3–)4–5. Fruit berry black, ellipsoid or oblong 6–12 × 6–7 mm, style persistent, short.

Distribution. Endemic to Cuba. Known from two separate mountainous areas: the Cordillera de Guaniguanico in the northwest and the Guamuhaya massif in the south centre of the island ( Figure 5 ).

Habitat and ecology. Mahonia sagrana has been found in karstic dense woodlands at or near hill and mountain tops at 440–1000 m. It has been collected in flower from July and November and in fruit from August to February.

Etymology. Named after Ramón de la Sagra (1798–1871), who collected the first specimen of the species and who was one of the first to warn about the consequences of deforestation in Cuba (Funes Monzote, 2007).

Proposed IUCN conservation category. Mahonia sagrana (as M. tenuifolia ) was not assessed in the Red List of Cuban Flora ( González Torres et al., 2016) but was recently considered Vulnerable (VU D2) by Bécquer & Rodríguez-Cala (2020). The assessment was based on its restricted geographical distribution (area of occupancy, 13 km 2), but its habitat has a low history of anthropogenic impact and is mostly within protected areas (i.e. Ecological Reserve ‘Lomas de Banao’, Natural Protected Landscape ‘Topes de Collantes’ and Biosphere Reserve ‘ Sierra del Rosario’). However, several studies project that species of these mountains will be affected in the coming decades by climate change; therefore, Bécquer & Rodríguez-Cala considered this a plausible threat based on the Red List Criteria, version 3.1, of the IUCN (2012).

Notes. In Gómez de la Maza (1889) there is the entry “ Berberis fraxinifolia, Hook. ? ( Mahonia cubensis Rich. )”, this synonymy being repeated in Gómez de la Maza (1890, 1897), where “Rich.” is expanded to “Richard”, presumably referring to Achille Richard. No source was given for this “ Mahonia cubensis ”. Richard’s work ( Richard, 1845) on the vascular plants of Cuba has no reference to any Berberidaceae , and the specimen sent to him by de la Sagra remained unidentified in his herbarium until 1960. Interestingly, neither Berberis nor Mahonia is listed in two earlier articles by Gómez de la Maza (1886a, 1886b), nor have we found any previous reference to a “ Mahonia cubensis ” by any other author. What prompted Gómez de la Maza in 1889 to refer to such a taxon thus remains unknown. Should any published description of a Mahonia cubensis from Cuba ever be found, then our M. sagrana would be a synonym. However, on current evidence such a discovery would seem quite unlikely.

Additional specimens examined. NB: collection numbers for specimens collected by Acuña, Bro. Leon, I. Arias et al., E. Bécquer et al., F. Felipe et al. and J. Natenson follow Gabancho et al. (2008).

CUBA. Cordillera de Guaniguanico : Sierra de los Órganos, Pinar del Río: Loma La Guira, ix 1935, J. Natenson SV-14323 ( HAC). Sierra Rosario , Artemisa: San Cristobal, 7 xii 1915, Roig 1188 ( HAC); San Cristobal, 8 xii 1915, Roig 8000 ( HAC); Sierra de los Organos, in forest on top of Peña Blanca, 750 m,

30 iii 1923, E. L. Ekman 16379 (G G00405854, NY 0512012, S S16-43276); Sierra de los Organos,

grupo del Rosario Sierra de Pendejeral , on top of the mountain, 750 m, 13 ix 1923, E. L. Ekman 17514

(S S16-43288); Sabicu, Rangel, Rosario Mts, 550 m, 1 vii 1926, Bro. Leon LS-12595 ( GH 01154528,

HAC, US 01078957); Rangel , Sierra del Rosario, on crest of Loma Sabicu, 440 m, viii 1926, Bro. Leon LS-12693 ( GH 01154526 , HAC, HAJB (3), NY 01512015-16 , US 01078958) ; Pan de Guajaibón y Pendejeral, 26 xii 1936, J. Acuña SV-10944 ( HAC) ; ibid., J. Acuña SV-11087 ( HAC) ; Pan de Guajaibón ,

La Palma, J. Acuña SV-23337 ( HAC) ; Rangel , Sierra del Rosario, viii 1941, Bro. Leon LS-20435 ( HAC) ; Sobre el paredón de la Jutía, Rangel, 500 m, i 1953, Bro. Alain 2713 ( GH 01154530 , HAC, US 01078959) ; Bosques : Rangel, Sierra del Rosario, viii 1954, Bro. Alain 6085 ( HAC) ; Rangel , Sierra del Rosario, viii 1954, Bro. Alain 6086 ( GH 01154529 , US 01078960) ; Municipio de Bahia Honda : Pan de Guajaibón, en el extremo oeste de la Sierra del Rosario, 22°47′23.6′′N, 83°21′59.5′′W, 699 m, 8 v 2004, J. R. Abbott & E. Bécquer 18893 ( FLAS) .

Guamuhaya massif: Sancti Spíritus: Edge of Río Tayaba , at San Juan de Letrán, 30 iii 1924, E. L. Ekman 18940 (S S16-43284) ; Trinidad Mts, Pico Sobrero , c. 1000 m, 9 vii 1953, G. L. Webster et al. 232 ( DAVH) ; Crest of Pico Potrerillo, Trinidad Mts , c. 1000 m, 16 vii 1957, Bro. Alain 6364 ( GH 01154525 ) ; Pico Potrerillo , 950 m, 21 vi 1970, A. Borhidi & O. Muñis s.n. ( BP) ; San Juan, Cuabales , near summit on steep slope, 6 ii 1986, G. Yelenevsky & A. Yelenevskaya s.n. ( MW MW0574605 ) ; Topes de Collantes, Mogote mi Retiro , 22.366667°N, 22°22′N, 79°50′W, 850 m, 2 vii 1993, P. Acevedo-Rdgz. et al. 5591 ( US 00681647) ; Lomas del Garrote, A/ P El Naranjal, Lomas del Banao , 700 m, 8 x 1995, E Bécquer 789 (Herbario Jardín Botánico, Sancti Spíritus) ; Base del farallón norte de Tetas de Juana , 21°51′46.8′′N, 79°35′56.4′′W, 600–650 m, 10–14 xii 2017, E. Bécquer et al. HFC-89693 ( HAJB) ; Reserva Ecológica Lomas de Banao, cabezadas del río Tayabacoa, alturas cársicas entre loma La Ventana y Puerta de la lechuza, 21°51′53.7′′N, 79°36′54′′W, 700–740 m, 21–25 vi 2022, F. Felipe et al. HFC-90958 ( HAJB). Cienfuegos: Vicinity of Soledad , viii 1941, R. A. Howard 6248 ( GH 01154527 ) ; Cumanayagua, Lomas alrededor de Los Tornos , 4 xi 1987, I. Arias et al. HFC-62849 ( HAJB (4)) ; Cumanayagua, Camino entre Los Tornos y El Sopapo, 6 xi, 1987, I. Arias et al. HFC-62931 ( HAJB (4)) ; Cumanayagua , subida al Pico San Juan, 7 xi 1987, I. Arias et al. HFC-62947 ( HAJB) ; Cumanayagua Lomas al sur del Pico San Juan, 9 xi 1987, I. Arias et al. HFC-63274 ( HAJB (2)) .

Without locality: s.d., R. de La Sagra 151 (P ex. Herb. Richard P02327264).

C. Wright 1855 specimens: s.d. ( BP, BRU ex Herb. Bennett, BRU ex Herb. Olneyannum, K K 001264493, P00752196, S S16-43286, US 01078961); 1860–1864 ( BM BM001209248 , G G00405936, G00405991, P P00752197, YU 244798 ) ; “ Retiro Sept. 1... San Christobel ” 1860–1864 ( GH 01154524 ) ; “ Mogot de Mono ” x 8, 1865 ( NY 01512010 ) ; sine ano ( NY 1512011 ) ; “ Cultivated , but indigenous on the northern coast” 1863 ( GOET GOET020003 ). “Cult but indigenous on the northern coast”, v 27 sine ano ( MO 1932676 ) .

Cultivated specimens: Cienfuegos: Cienfuegos Botanical Garden , 17 ii 1933, J. G. Jack 8676 ( NY 01512013 ) ; iv 1934, F. G. Walsingham s.n. ( NY 01512014 ) .

Living plants: Guamuhaya massif: Sancti Spíritus: Trinidad, Parc Codina, near mirador. Observed by Eldis Bécquer in ii 2015 and viii 2018 and Julian Harber in iv 2016. Two plants cultivated at the nurseries of National Botanical Garden, Havana (HAJB-201700057, from collection HFC-89693).

Mahonia tenuifolia (Lindl.) Czerw. & Warsz. Cat. Pl. Hort. Cracov 292 (1864). Basionym: Berberis tenuifolia Lindley , Edwards’s Bot. Reg. 24 (Misc.): 64 (1838); Odostemon tenuifolius Standl., Contr. U.S. Natl. Herb. xxiii: 270 (1922). – Type : Mexico, [near Veracruz, Zacuapam, xii 1836], K. T. Hartweg s.n. [262] (lectotype BM designated by Ahrendt, 1961, missing or non-existent; new lectotype designated here ( ICN Article 9.21: Turland et al., 2018) CGE 05241 ex Herb. Lindley!; isolectotypes K [ K000994123 !], LD [LD1420317] image!; probable isolectotype LE s.col., s.n., s.d., [01041505] image!) .

Berberis fraxinifolia Hook. Icon. Pl. 4: t. 329 (1841). – Type: Mexico, Xalapa , 3000 m,

i–x 1840, H. G. Galeotti s.n. [4640] (lectotype K ex. Herb Hook., designated here,

K [ K000407236 !] ; isolectotypes BR (3) [BR0000027930854, BR0000027930861, BR0000027930892] images!, G (2 sheets) [G00418680] images!, LE [01041505] image!, MEXU ex BR [1313357] image!, W not found; possible isolectotype E ex GL [E00318125!]).

Shrubs, evergreen, c. 5 m tall. Leaves imparipinnate, 30–35 cm long; leaflets 3–6 pairs, contiguous, lowest pair (3–)5.5–8(–10) cm above base of petiole, abaxially palish yellow green, adaxially dark green, shiny, ovate-lanceolate or lanceolate, rarely broadly ovate; margin entire; apex acute or acuminate, rarely obtuse, papyraceous; midvein raised abaxially, impressed adaxially, lateral veins inconspicuous on both surfaces, base attenuate; margin entire, slightly undulate; apex acuminate. Inflorescence a single or 2(–5)-branched raceme to 37 cm, 25- to 45-flowered; pedicel 4–15 mm; floral bracts triangular ovate, 1.5 × 0.75–1 mm, apex acuminate. Flowers yellow, c. 4 mm in diameter; sepals in three whorls; outer sepals elliptic orbicular or triangular ovate, 1 × 1 mm; median sepals elliptic, 5 × 5 mm; inner sepals elliptic, 5–6 × 5 mm. Petals elliptic, 4–5 × 2.5–3 mm, glands separate, apex rounded. Stamens dentate, with teeth horizontal or pointing downwards, 2.5 mm; anther connective extended, rounded. Ovary 3 mm; ovules 2. Fruit berry black, obovoid 7–8 × 4–5 mm, style persistent, conspicuous.

Distribution. Endemic to Mexico. Known from Veracruz and Oaxaca States.

Habitat and ecology. Found among secondary vegetation in deciduous and semi-evergreen forests including oaks, c. 900–1020 m.

Proposed IUCN conservation category. Data deficient (DD).

Notes. Determining the type specimens of Mahonia tenuifolia was not completely straightforward. Lindley’s protologue of Berberis tenuifolia stated that “seeds were sent by Mr Hartweg to the Royal Horticultural Society from the neighbourhood of Vera Cruz where it was found growing at a place called Zacuapam near the rancho of Mr Lavater”. A specimen at GCE (barcode 05241 (CGE)) that is ex Herb Lindley is annotated “ Mexico Hartweg” and “ Berberis tenuifolia ”, with a later additional annotation stating “ Berberis tenuifolia Bot Reg 1838 ”.

Subsequent to Lindley’s (1838) protologue, Bentham (1839), enumerating a small number of collections where there were insufficient specimens to distribute to all those who had subscribed to Hartweg’s Mexican expedition, included Hartweg 262 from Zacuapam near Vera Cruz, collected in October 1836, identifying this as Berberis tenuifolia and citing Lindley’s 1838 article.

Hartweg’s expedition was a long one, lasting from 1836 to 1843, during which time he collected not just in Mexico but in Guatemala and elsewhere in Central America. The account he gave on his return ( Hartweg, 1848) stated that he arrived in Mexico at Vera Cruz on 3 December 1836 (thus contradicting Bentham’s dating) and immediately took advantage of an offer to stay with a Mr Laveter on his farm at Zacuapam, which he reached after a two-day ride. He described Zacuapam as being at about 3000 ft “on the eastern declivity of the snow-clad Orizaba”. In this locality he recorded “ Berberis tenuifolia forming a shrub 10 to 12 feet high”. He left Zacuapam on 27 December, arriving at Jalapa [Xalapa] the following day. Although other species of Berberis (i.e. Mahonia ) are recorded for later in his expedition, this is the only reference to B. tenuifolia .

There would appear to be no published reference regarding where any specimen of Hartweg’s collection might have been until Ahrendt (1961) cited “ Mexico; Vera Cruz State, below Mt Orizaba, near Zacuapam, 3000 ft, 1836, Hartweg 262 (Type BM)”. Given that Lindley did not cite any herbarium, this undoubtedly constituted a lectotypification ( ICN Article 9.10: Turland et al., 2018). However, despite a thorough search, no such specimen has been found .

At Kew, however, there is an unnumbered specimen annotated as “ Berberis tenuifolia Lindl. Bot Reg Misc 1838 p 64 Zacuapam nr Vera Cruz Hartweg 1839”. The date, 1839, is clearly a mistake because in 1839 Hartweg was in Guatemala and elsewhere in Central America, but the Kew specimen is undoubtedly a type. However, the existence of this specimen raises a question as to whether there was ever a such specimen at BM, for Ahrendt was not always an accurate recorder and there is at least one instance of where he recorded a specimen at BM when it is actually at K. This is the type specimen of his Berberis orthobotrys var. rupestris from India ( Ahrendt, 1961) – for further information, see Harber (2020).

The fact that the Kew specimen of Mahonia tenuifolia does not have the collection details found in Ahrendt’s account is not necessarily important, in that it was sometimes Ahrendt’s practice to give collection details from protologues and elsewhere that are not from the specimen sheets he cites (see Harber, 2020). However, given the uncertainty as to whether or not the Kew specimen was actually the one Ahrendt was referring to, it would seem appropriate to designate the GCE Hartweg specimen that was originally in Lindley’s personal herbarium as a new lectotype and the Kew specimen as an isolectotype. Should the type specimen eventually be found at BM, the selection of the GCE specimen would be void.

The label of a probable isolectotype at LE simply states “ Bentham 1839 ” and “ Berberis tenuifolia Zacuapam nr Veracruz ”, the latter being in the same hand as the text on the label of the specimen at K. There would seem no reason for citing Bentham unless it was in relation to Hartweg’s collection.

Hooker’s (1841) protologue of Berberis fraxinifolia was accompanied by a line drawing and a brief description and gave the collection details simply as “Xalapa, Mexico, Galeotti”, without either date of collection or collector’s number. Given that it was in a publication he edited subtitled “with Brief Descriptive Characters and Remarks of New or Rare Plants selected from the author’s herbarium”, it seems reasonable to assume that the Galeotti specimen at Kew that is stamped “Herbarium Hookerianum”, and that has both a collector’s number and a date, is at least one of the specimens the protologue was based on. Whether it was the only one cannot be proved. Therefore, it is designated here as a lectotype rather than recognised as the holotype.

The Galeotti specimen at E cited above as a possible isolectotype is unnumbered and has no collection details beyond “ Jalapa ”. Fedde (1901) cited a specimen of Galeotti 4640 at W; its presence there is confirmed by an 1840 entry in their accessions register. However, the specimen cannot now be located. A large number of specimens of Berberidaceae were loaned by W to Schneider at B before the Second World War, and although most were returned, eleven sheets were not. It is probable that Galeotti 4640 was among these and therefore lost when the Berlin herbarium was destroyed (Christian Brauchler, personal communications, 23–25 August 2021).

Additional specimens examined. MEXico. Veracruz State: Mirador, 1839, J. Linden 991 (FI [FI018235],

G [G00426731, G00426733], GENT [ GENT10163672 ], K ex Herb. Hooker [ K000407234 ], LE [01041508] ); Mirador, viii 1841 F. M. Liebman 254.(C (2), O [V:2132516]); Mirador, x 1841, F. M. Liebman s.n. ( HBG [HBG-525597], M, NY [03091262]) ; Mirador, x 1841, F. M. Liebman 254 (C); Mirador ii 1842, F. M. Liebman s.n. ( LD [1974465]) ; Mirador, vii 1843, F. M. Liebman s.n. ( FI [ FI018236 ], UPS [V-865713]) ; Mirador, 1841–1843, F. M. Liebman s.n. (H [1348030], L [L.1746580], O, UPS [V:2132515], VT [UVMVT139773]); Mexico, s.d., F. E. Leibold 18 (B [10 0743100]); Mexico, 1848, A. Ghiesbreght 157

(G [G00426734], LE [01041509]); Mirador, c. 1000 m, ix 1906, H. Ross 640 (BP, M); Zacuapam, vii

1914, C. A. Purpus 7661 (GH (2), NY [03091267], UC [191111], US [00899927]); Zacuapam, 900 m,

22 iv 1922, C. A. Purpus 347 ( HBG [HBG-525596], GH) ; Zacuapam, viii 1929, C. A. Purpus 12056 ( GH,

NY [03091261]); Near Zacuapam, ix 1929, C. A. Purpus 13075 ( BM, CAS [0577428], K, S [S18-3175]) ; Zacuapam, ix 1930, C. A. Purpus 13076B (L [L.1746578-79], MICH [1550730]); Near Zacuapam, xi 1930, C. A. Purpus 13075B (C); Near San Martin Tlacotepec, xi 1932, H. W. von Rozynski 586 (F [V0083000], MICH [1550729], NY [03091260], US [00899926]) ; El Mirador, SE of Poblado, Mun. Totutla, 1000 m, 10

x 1970, F. Ventura 2553 ( CAS [0577429], CR [48357], ENCB, MICH [15507280], MO, TEX) ; Mata Obscura , Mun. Totutla, 900 m, 9 xii 1970, F. Ventura 2964 ( CAS [0577426], CR [48280], ENCB, MICH [1550727], MO, XAL [115376]) ; El Mirador Mun. Totutla , 1020 m, 8 x 1971, F. Ventura 4217 ( CAS [0577427], ENCB, MICH [1550726], XAL) ; Mun. Totutla , Mata Obscura, 850 m, 10 i 1973, F. Ventura 7685 ( CAS [0577430], ENCB, MEXU [260373], MO [6538501]) ; Xalapa , Potrero del Rancho La Palma, 1000 m, 10 iii 1973, M. Chazaro 41 ( ENCB, MEXU [606205]) ; Totutla , El Mirador, 13 vii 1976, 1000 m, F. Ventura 12993, ( MEXU [363516]) ; Totutla , Mirador, 900 m, 11 viii 1976, F. Ventura 13547 ( MEXU [363515]) ; Mata Obscura , Mun. Totutla, 850 m, 26 vii 1977, F. Ventura 14314 ( CAS [0577431], ENCB, IEB, MEXU [418658], XAL) ; Edo , Mun. Alto Lucero , camino del poblado Cruz Blanca a Sobrero, 19°45′N, 96°51′W, 1000 m, 9 ix 1979 ,

J. I. Calzada 5691 (F [1875193], XAL); Totutla , El Encinal, 750 m, 8 ix 1980, F. Ventura 17811 ( ENCB, MEXU [360454]) ; Edo , Mun. Totutla, 19°12′N, 96°51′W, 950–1000 m, 21 xi 1981, M. Nee & G. Cortés 23385 (F [1932649], K, NY [03091266]) ; Barranca 1 km S of Palmillas, 19°13′1.2′′N, 96°46′1.2′′W, 600 m, 23 v 1985, M. E. Medina & S. A. Contreras 134 ( ENCB, MEXU [691569]) ; Barranca de Palmillas , 2 km

SE de dicha poblacion, Municipo Puente National, 19°13′N, 96°46′W, 450 m, 24 x 1985, M. E. Medina & E. F. Vazquez 599 ( MEXU [771707], XAL [133759]) ; Totutla, Barranca 3 km SE of EL Mirador, 19°13′N, 96°52′W, 5 ii 1993, S. Avedaño & C. Durán 3052 ( MEXU [723140], XAL). Oaxaca State: Tehuantepec ,

“ El Ocotal”, Cerro Guiengola, 16°21′N, 95°19′1.2′′W, 950 m, 13 vi 1986, C. L Torres et al. 419 ( MEXU [1208344]) ; Tehuantepec Entrando a “El Ocotal”, lado derecho del arroyo por la ladera poniente

Cerro Guiengola, 14 xi 1986, C. L Torres et al. 268 ( MEXU [1208422]) ; Tehuantepec, “El Ocotal”, Cerro Guiengola, 16°21′N, 95°19′1.2′′W, 26 xi 1986, C. L Torres et al. 538 ( MEXU [1208347-48]) .

Figures

Figure 1. Mahonia sagrana Harber & Bécquer, sp. nov.: flower structure. A, Outer sepal; B and C, median sepals; D and E, inner sepals; F and G, petals; H, stamen; I, pistil; J, ovules; K, bract; L, pedicels. Drawn by Nicola Adams, based on a line drawing, by C. K. Schneider, attached to the sheet of holotype E.L. Ekman 17623 (S).

Figure 2. Mahonia sagrana Harber & Bécquer, sp. nov.: pistil and stamen of a plant in Parc Codina, collected by N. Arbelo. No voucher material made.

Figure 3. Mahonia sagrana Harber & Bécquer, sp. nov.: in flower, Jardín Botánico National, Havana. No voucher material made. Photograph: Luis M. Leyva.

Figure 4. Mahonia sagrana Harber & Bécquer, sp. nov.: leaf blade of a plant at Parc Codina, near Mirador, Trinidad, Santi Spiritus. No voucher material made. Photograph: Julian Harber.

Figure 5. Distribution of Mahonia sagrana in Cuba. Map created by Ernesto Testé.