Fridericia craterophora (Mart. ex DC.) L. G. Lohmann

2. Fridericia craterophora (Mart. ex DC.) L. G. Lohmann in Ann. Missouri Bot. Gard. 99: 436. 2014 ≡ Bignonia craterophora Mart. ex DC., Prodr. 9: 147. 1845 ≡ Arrabidaea craterophora (Mart. ex DC.) Bureau in Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 6, 6(3): 422. 1892. – Holotype: Brazil, Provinciae M. Geraes (= Minas Gerais), deserto Serro Frio versus Contendas, Feb, Herb. Fl. Bras. 335 (= C. F. P. von Martius s.n.) (M barcode M0086376!). – Fig 1F–H, 2.

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= Bignonia craterophora var. acutifolia Mart. ex DC., Prodr. 9: 147. 1845 ≡ Arrabidaea craterophora var. acutifolia (Mart. ex DC.) Bureau in Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 6, 6(3): 422. 1892. – Holotype: Brazil, in campis editis Provinciae M. Geraes (= Minas Gerais), s.d., Herb. Fl. Bras. 336 (= C. F. P. von Martius s.n.) (M barcode M0086377!).

Arrabidaea craterophora subvar. glabrescens Bureau & K. Schum., Fl. Bras. 8(2): 67. 1896. – Lectotype (designated here): Brazil, Minas Gerais, Lagoa Santa, 1867, E. Warming Bignon 33 (BR barcode BR0000008802637 [image!]).

Notes — Four syntypes were listed in the protologue of Arrabidaea craterophora subvar. glabrescens . Among these specimens, Pohl 1376 (only found in W) is a sterile specimen and Gardner 3899 (only found in K) has a label indicating the locality and date of collection. However, the protologue cites “loco haud accuratius addicto” indicating that Bureau and Schumman likely did not see the specimen deposited in K. Out of the two Warming specimens from Lagoa Santa deposited in BR, Warming s.n. (barcode BR0000008802965 ) has two distinct determinations of the subvarieties annotated by Bureau, making this specimen confusing. The other specimen, Warming Bignon 33 (barcode BR0000008802637 ), has a detailed habitat description that matches the habitat cited in the protologue; furthermore, this is a good-quality specimen, bearing a single identification handwritten by Bureau that matches the species identity. The specimen Warming Bignon 33 is selected here as the lectotype .

Arrabidaea craterophora subvar. velutina Bureau & K. Schum., Fl. Bras. 8(2): 67. 1896. – Lectotype (designated here): Brazil, Goiás [habitat in ejusdem provinciae campis elevatis], A. Glaziou 21861 (K barcode K000402725!; isolectotypes: P barcode P02895999!, R barcode R000011030 !).

Notes — In the Flora brasiliensis, Bureau & Schumann (1896: 67) listed a single gathering of Arrabidaea craterophora subvar. velutina made by Martius (“habitat in ejusden provincieae campis elevatis”). However, this gathering was not located. In the same treatment, Bureau & Schumann (1897: 406) added two more examined gatherings of this subvariety in the Mantissa at the end of the Flora , including a specimen from Goiás ( Glaziou 21861) and a specimen from Santa Luzia do Rio das Velhas in Minas Gerais ( Schwake 11421). Of these, we were unable to locate Schwake 11421, but we found three specimens of Glaziou 21861, deposited in K, P and R. The specimen in P (barcode P02895999) has a handwritten label with a specific location (“Paranauá près des Cascades, Goyaz”) and the date of collection is 15 February 1898. Furthermore, information handwritten on the typed label indicates that this material was donated to P from Glaziou’s private herbarium =

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– by Glaziou’s daughter after his death in 1907. The specimen in K includes less detailed locality information (“Chiefly Province of Goyaz”) that matches the locality information provided in the protologue. Furthermore, the specimen K000402725 was identified by Bureau and was received by K in 1896, the same year of the publication of A. craterophora subvar. velutina ; this specimen is selected here as the lectotype.

Arrabidaea lenticellosa Bureau & K. Schum., Fl. Bras. View in CoL 8(2): 64. 1896 [non Arrabidaea lenticellosa Pittier View in CoL in J. Wash. Acad. Sci. 18: 340. 1928]. – Lectotype (designated as “type” by Gentry 1982a: 101): Brazil, Mato Grosso, L. Riedel 1469 (K barcode K000402704!; isolectotypes: LE barcode LE 01072038 [image!], LE barcode LE 01072039 [image!], OXF barcode OXF00085420 [image!]).

Note — In the protologue of Arrabidaea lenticellosa, Bureau & Schumann (1896) View in CoL did not indicate where the holotype was deposited. Gentry (1982a) indicated the specimen in K as the “type”, which before 2001 is sufficient to designate a lectotype according to Art. 7.11 and 9.23 of the Code ( Turland & al. 2018).

“ Arrabidaea craterophora var. obtusifolia ” Mart ex. Bureau & K. Schum, Fl. Bras. 8(2): 67. 1896. Designation not validly published (Art. 26.2)

Note — Arrabidaea craterophora var. obtusifolia was described based on the type of Bignonia craterophora DC. View in CoL

“ Fridericia craterophora ” (Mart. ex DC.) L. G. Lohmann in Monogr. Syst. Bot. Missouri Bot Gard. 107: 1605. 2008. Designation not validly published (Art. 41.1).

“ Fridericia craterophora ” (Mart. ex DC.) L. G. Lohmann in Forzza & al., Cat. Pl. Fung. Brasil 1: 765. 2010. Designation not validly published (Art. 41.1).

Morphological description — Lianas, shrubs or scandent shrubs, evergreen, up to 10 m high. Branches terete to subquadrangular, striate and lenticellate, brown or grey when dry, glabrescent to velutinous, with simple trichomes; without interpetiolar ridge, with interpetiolar glandular fields; prophylls of axillary buds triangular or narrowly triangular, not apiculate, 1.7–2.1 mm long. Leaves 1- or 2-foliolate; petiole 0.1–0.8(–1.1) cm long, villous, with simple trichomes; petiolules with equal lengths, 0.2–0.4(–0.7) cm, villous, with simple trichomes; blades chartaceous, smooth (sometimes slightly bullate), margin revolute, central leaflet lanceolate, ovate or obovate, base rounded, apex acute or acuminate, lateral leaflets elliptic to ovate, 2–9.7 × 1.1–4.2 cm, base cuneate or rounded, apex acuminate (sometimes retuse or rounded), adaxially not vernicose, glabrescent to puberulous, with simple trichomes throughout (sometimes hispid), abaxially glabrescent to velutinous, with simple trichomes throughout (sometimes only pubescent at veins), venation pinnate, secondary veins raised, tertiary veins raised, without pocket and with or without trichome tuft domatia. Inflorescence terminal and axillary, thyrse with abortion of lateral buds, resembling a raceme, with 1 or 2 orders, first-order peduncles 0.8–4.8 cm long, second-order 0.3–1.7 cm, glabrescent to velutinous, with simple trichomes; bracts linear, 1.3–3 mm long, caducous; bracteoles linear, 0.5–0.8 mm long, caducous; pedicels 0.2–0.4 cm long. Calyx tubular or obconic, not costate, truncate and laterally or irregularly split, 1.5–1.8(–2) × 0.4–0.6(–0.9) cm, glabrescent with rim pubescent to velutinous throughout, with simple trichomes or lepidote throughout, chartaceous, green with pinkish apex, with glandular areas evenly distributed. Corolla infundibular, not curved, slightly zygomorphic, furrowed, 4.1–6.4 cm long, 0.9–2.5 cm wide at tube mouth, externally villous, without glands, lobes 0.7–0.9 × 0.6–1.1 cm, margin rounded and undulate, dark red, lilac, pink or purple. Androecium with all stamens included; longer filaments 1.6–2 cm long, shorter filaments 1.2–1.6 cm; staminode 1.3–2 mm long; anthers 3–4.4 mm long, connectives protruding 0.5 mm. Gynoecium with ovary cylindric, 2–4 × 0.9–1.3 mm, furrowed, lepidote; style 2.8–3.4 cm long; stigma lanceolate or orbicular; nectar disk annular and smooth under ovary, c. 0.8 × 1.2 mm. Fruit linear, flat, margins not raised or slightly raised, central ridge not raised (sometimes slightly raised), valves coriaceous, rough, 9.5–28.5 × 0.6–1 cm, glabrous; septum coriaceous. Seeds oblong, body oblong, 0.4–0.7 × 1.5–2 cm, wings hyaline, 0.1–0.3 cm wide, margins crisped.

Phenology — Produces flowers from November to June. Fruits were collected from June to January.

Distribution and habitat — Fridericia craterophora is mainly distributed in the Cerrado and Chaco, reaching ecotone areas between the Amazon and the Atlantic forest. It is distributed through Bolivia ( Santa Cruz), Brazil ( Bahia, Distrito Federal, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, PiauÍ, Rio de Janeiro, Rondônia, São Paulo and Tocantins) and Paraguay ( Amambay).

Conservation status — Fridericia craterophora is categorized as Least Concern (LC) based on its Extent of Occurrence ( 3,195,212 km 2) and Area of Occupancy ( 160,000 km 2).

Remarks — Fridericia craterophora is one of the most variable species in the “ Neomacfadya ” clade due to the variation in leaflet number, shape, texture and indumentum. This species is easily recognized by the villous and short petioles 0.1–0.8(–1.1) cm. The petioles are commonly so short that the leaf appears to be sessile, with a thickened area at the base. In fact, the pair of 2-foliolate leaves located on the opposite side of branches can be confused with four verticillate, simple leaves ( Fig 1G), as described by Gentry (1980). However, a scar is visible between the two lateral leaflets, and a central tendril can also be detected between the leaflets [see Britto 33 (MBM) and Glaziou 21861 (K, P)].

Fridericia craterophora can be confused with the sympatric F. resinosa and F. simplex . The villous and shorter petiole [0.1–0.8(–1.1) cm], petiolules [0.2–0.4(– 0.7) cm] and non-vernicose adaxial surface of the leaflets of F. craterophora contrast with the glabrous and longer petioles ( 1–3.3 cm), petiolules [(0.5–) 1–1.8 cm] and vernicose adaxial surface of the leaflets of F. resinosa . The characters that distinguish F. craterophora from F. simplex are the branches subquadrangular and lenticellate (vs branches terete and sparsely or not lenticellate in F. simplex ), the leaflets with pinnate venation (vs leaflets with actinodromous venation in F. simplex ) and the calyx glabrous to villous (vs calyx lepidote in F. simplex ). In ecotone areas between Chaco and Amazonian rainforests F. craterophora is often confused with F. japurensis . These two species can be distinguished by the longer petioles [(1.1–) 1.4–2.8 cm] and petiolules [(1.1–)1.8–2(–2.7) cm] of F. japurensis , contrasting with the shorter petioles [0.1–0.8(–1.1) cm] and petiolules [0.2–0.4(–0.7) cm] of F. craterophora . Furthermore, F. japurensis has fruits with valves woody and sticky, whereas the fruit valves of F. craterophora are coriaceous and not sticky. Fridericia craterophora can also be confused with F. podopogon due to the short petiole and petiolules and similar shape of the leaflets. However, F. podopogon is restricted to the N portion of Central America and Cuba, whereas F. craterophora is restricted to C South America.

Gentry (1982a) treated Arrabidaea lenticellosa Bureau & K. Schum. as a synonym of A. japurensis [ ≡ Fridericia japurensis ], although no comments about this synonymization were provided. However, a careful analysis of the type material of A. lenticellosa indicat- ed a number of differences with F. japurensis , namely: the shrubby habit described in the label of the specimen LE 01072038 (vs liana in F. japurensis ), the leaves 1- or 2-foliolate (vs 2-foliolate in F. japurensis ), small petioles ( 0.4–1 cm long vs 1.7–2 cm long in F. japurensis ) and leaflets with pinnate venation (vs actinodromous basal in F. japurensis ). All these features are common in F. craterophora . Based on these morphological traits, we treat A. lenticellosa as a synonym of F. craterophora .

The name Arrabidaea lenticellosa Pittier was published for another species from Venezuela. Sandwith (1968) synonymized this name in A. corallina (Jacq.) Sandwith (= Tanaecium dichotomum (Jacq.) Kaehler & L. G. Lohmann ), which was subsequently accepted by Gentry (1977b, 1982 a, 1982b) and by Lohmann & Ulloa Ulloa (2022 +). We follow this same circumscription here.