Fridericia is a conspicuous genus of neotropical lianas, with 60 species distributed among seven lineages that are broadly distributed through the neotropics in both dry and wet environments. As part of ongoing taxonomic studies of the genus, we provide taxonomic revisions of two clades: (1) the “Acrodromous venation” clade and (2) the “Piriadacus” clade. The former is characterized by 1- or 2-foliolate leaves with acrodromous venation, tubular calyces and coriaceous fruits with slightly raised edges. The latter is recognized by the red, pink-reddish or orange, tubular corolla and exserted or sub-exserted anthers. We recognize two species in each lineage and provide identification keys, morphological descriptions, illustrations, distribution maps and comments on the distribution, phenology, conservation status and taxonomy. We propose the lectotypification of 13 names: Arrabidaea brachypoda var. acuminata, A. brachypoda var. attenuata, A. brachypoda var. firma, A. brachypoda var. induta, A. brachypoda var. rigida, A. platyphylla var. lasiorrhachis, A. platyphylla var. puberula, Bignonia brachypoda var. firmula, B. erubescens var. breviflora, B. erubescens var. subtruncata, B. regnelliana, Cuspidaria erubescens var. glabrescens and Petastoma simplicifolium. We further correct the typification of B. erubescens var. breviflora and B. erubescens var. subtruncata and propose two new synonyms for F. platyphylla (i.e. A. celastroides and B. violacea).
Citation: Kaehler M. & Lohmann L. G. 2021: Taxonomic revisions in Fridericia (Bignonieae, Bignoniaceae) I: the “Acrodromous venation” and “Piriadacus” clades. – Willdenowia 51: 181–194.
Version of record first published online on 21 June 2021 ahead of inclusion in August 2021 issue.
Introduction
Fridericia Mart. emend. L. G. Lohmann is one of the largest and most complex genera in the plant family Bignoniaceae (Kaehler & al. 2019). This genus belongs to tribe Bignonieae, which includes lianas and is broadly distributed throughout the neotropics, where they occur in both wet and dry habitats (Lohmann & Taylor 2014). The generic classification of Bignonieae remained problematic for more than a century due to overlapping patterns of morphological variation and a lack of diagnostic generic characters. Molecular phylogenetic data (Lohmann 2006) allowed for a re-circumscription of monophyletic genera diagnosed by morphological synapomorphies (Lohmann & Taylor 2014). Under this new generic classification system, Fridericia is much more broadly circumscribed, including many species of the previously recognized Arrabidaea DC.
Arrabidaea originally included four Brazilian species (Candolle 1838). Its circumscription was later expanded to accommodate 21 species (Candolle 1845). By the time the monumental treatment of Flora brasiliensis was published, Arrabidaea included 70 species and an incredible range of morphological variation (Bureau & Schumann 1896). The Flora brasiliensis remains the most comprehensive treatment of Arrabidaea to date. To help organize the broad range of morphological features included in the genus, three sections distinguished by the number and arrangement of ovules per locule were proposed (Bureau & Schumann 1896), as follows: (1) A. sect. Microcarpaea K. Schum., with fewer than five ovules in each of the two locule rows; (2) A. sect. Macrocarpaea K. Schum., with more than five ovules in each of the locule rows; and (3) A. sect. Paracarpaea K. Schum., with ovules affixed in four rows (Bureau & Schumann 1896). This infrageneric system was followed by subsequent authors who described new species within the genus (e.g. Kränzlin, Pittier, Rusby and Sprague). The circumscription of Arrabidaea continued to grow in subsequent years, especially due to the inclusion of the species-rich genera Cremastus Miers, Panterpa Miers, Paramansoa Baill. and Petastoma Miers (Sandwith 1968). The most-comprehensive treatment of the Bignoniaceae prior to the availability of phylogenetic data recognized an Arrabidaea with c. 100 species (Fisher & al. 2004). Despite a large number of species in the genus, the infrageneric classification system proposed by Bureau & Schumann (1896) was subsequently refuted by morphology (Sandwith 1968) and molecular characters (Lohmann 2006; Kaehler & al. 2019).
The first molecular phylogeny that sampled multiple members of Arrabidaea (Lohmann 2006) indicated that the genus was polyphyletic, leading to a revised circumscription (Lohmann & Taylor 2014). Namely, A. rego (Vell.) DC. [= Fridericia rego (Vell.) L. G. Lohmann], the type of Arrabidaea, was placed within the same clade as F. speciosa Mart., the type of the monotypic Fridericia, indicating that these taxa were best treated within a single genus (Lohmann 2006). Because Fridericia is the older name, 66 species of Arrabidaea were transferred into Fridericia, leading to a broadly circumscribed Fridericia with 67 species (Lohmann & Taylor 2014). The remaining species of Arrabidaea were shown to be more closely related to Cuspidaria DC., Tanaecium Sw. and Xylophragma Sprague and were transferred into those genera instead. While Fridericia is clearly monophyletic, the genus remained morphologically variable (Lohmann & Taylor 2014; Kaehler & al. 2019). The 60 species currently recognized in Fridericia are distributed in seven lineages characterized by morphological traits in an informal classification system (Kaehler & al. 2019).
Most species of Fridericia are lianas, although some are shrubs or treelets; the prophylls of the axillary buds are minute, triangular and distichous, sometimes concentrically disposed or foliaceous; the leaves are usually ternate with the terminal leaflet often replaced by a simple tendril, although 1-foliolate leaves are also found; the corolla is usually in shades of pink, less commonly white, red, orange or yellow; the ovary is lepidote, with two lines of ovules per locule; the fruits are linear, usually coriaceous, rarely woody, bearing winged seeds, with two hyaline or papyraceous wings (Lohmann & Taylor 2014; Kaehler & al. 2019).
As part of ongoing taxonomic studies of Fridericia, we provide taxonomic revisions of the “Acrodromous venation” and “Piriadacus” clades, both of which are exclusively South American and centred in Brazil. The “Acrodromous venation” clade is the first diverging lineage in Fridericia, whereas the “Piriadacus” clade is a small lineage within the following polytomy. Due to their small size, these two clades are treated together in this study. We provide an identification key, typifications, morphological descriptions, illustrations, distribution maps and comments on the geographical distribution, phenology, conservation and taxonomy for each clade.
Material and methods
Almost 700 gatherings and several duplicates, including all nomenclatural types deposited in HAL, K, M, MBM, MO, NY, P, R, RB, S, SPF, TEPB, UB and UPCB were analysed (acronyms following Thiers 2021+). Additionally, we consulted images in electronic repositories such as Global Plants ( https://plants.jstor.org/), JACQ ( https://www.jacq.org/), Reflora ( http://reflora.jbrj.gov.br/) and Species Link ( http://splink.org.br/) as well as photographs from C and LE provided by the staff of these two herbaria. The complete list of specimens examined is provided in Appendix 1 (wi.51.51202_Appendix_1.pdf), sorted by collector.
Types are listed alphabetically and herbarium accession numbers are provided when available. Whenever herbarium accession numbers were lacking, specimen barcodes were provided instead. Taxonomic headings include type information provided in the labels of each specimen and the phenological condition of each specimen. Whenever the data provided in the label differed from those included in the protologue, the additional information provided in the protologue was added between square brackets.
The terminology of leaf venation follows Ellis & al. (2009), trichome types follow Nogueira & al. (2013) and inflorescence morphology follows Weberling (1981). All other terms follow Lohmann & Taylor (2014).
Distribution maps were prepared using QGis 3.14.1 Pi (QGis Development Team 2020) using the Americas Base Map (Bletter & al. 2004). The conservation status of each species was estimated using the GeoCAT (Bachman & al. 2011). Because all species treated here have wide Extent of Occurrence (more than one million km2), we opted to estimate the Area of Occupancy using the auto-value option for the cell width (50 km of cell width for all species). The 2 km cell width recommended by the IUCN (2019) is too small for broadly distributed species with relatively small density of collections within this large area (Lughadha & al. 2018).
Results and Discussion
Key for the identification of the “Acrodromous venation” and “Piriadacus” clades and included species
1. Leaves 1- or 2(or 3)-foliolate, venation acrodromous; corolla pink or white “Acrodromous venation” clade: 2
– Leaves (2 or)3-foliolate, venation pinnate; corolla red, pinkish red or orange “Piriadacus” clade: 3
2. Inflorescence branches pubescent to velutinous; calyx cupular, costate, pubescent, chartaceous; corolla infundibuliform; stamens inserted. Broadly distributed through South America (Amazonian and Atlantic rainforests, Caatinga, Chaco and savannahs) 1. Fridericia platyphylla
– Inflorescence branches lepidote; calyx tubular, not costate, lepidote, coriaceous; corolla narrowly infundibuliform; stamens at same height as tube mouth. Restricted to lowland Amazonian rainforests 2. Fridericia prancei
3. Prophylls of axillary buds broadly triangular, arranged as a bromeliad, 1.8–2.1 mm long; blade of lateral leaflets 2.4–5.8(–10.6) × 1.2–5.7 cm; corolla zygomorphic. Caatinga, Cerrado and ecotone areas of Atlantic rainforest 3. Fridericia erubescens
– Prophylls of axillary buds triangular, arranged in 2 rows (sometimes concentrically disposed at inflorescence peduncles), c. 1 mm long; blade of lateral leaflets 9.1–15.2 × 4.5–11 cm; corolla actinomorphic. Lowland oriental Amazonian rainforests 4. Fridericia ornithophila
Taxonomic treatments
The “Acrodromous venation” clade
The species of the “Acrodromous venation” clade are characterized by 1- or 2-foliolate leaves with acrodromous venation, tubular calyces and smooth, coriaceous fruits, without prominent midrib and with edges slightly raised. These morphological features are consistent between the two species of this group and are not found concomitantly in any other species of Fridericia. The morphological complexity of this clade is mainly due to the wide morphological variation of F. platyphylla (Cham.) L. G. Lohmann.
1. Fridericia platyphylla (Cham.) L. G. Lohmann in Ann. Missouri Bot. Gard. 99: 442. 2014 ≡ Bignonia platyphylla Cham. in Linnaea 7: 679. 1832 ≡ Bignonia brachypoda var. platyphylla (Cham.) DC. in Candolle, Prodr. 9: 146. 1845 ≡ Arrabidaea brachypoda var. platyphylla (Cham.) Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 98. 1894 ≡ Arrabidaea platyphylla (Cham.) Bureau & K. Schum., Fl. Bras. 8(2): 38. 1896, nom. illeg. [non Arrabidaea platyphylla DC. in Candolle, Prodr. 9: 186. 1845] ≡ Arrabidaea platyphylla var. stricta Bureau & K. Schum., Fl. Bras. 8(2): 41. 1896. – Holotype: Brazil, fl., s.d., F. Sellow Herb. Reg. Berolinense 1841 [s.n.] (LE [image!]; isotypes: B destroyed [image negative 18452 at F!], HAL barcode HAL098686!, K barcode K000449124!, US barcode US282538 [image!]). – Fig. 1, 2.
≡ Bignonia brachypoda DC. in Candolle, Prodr. 9: 145. 1845, nom. illeg. (Art. 52.1) ≡ Arrabidaea brachypoda Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 98. 1894. – Lectotype (designated here): Brazil, fl., s.d., F. Sellow Herb. Reg. Berolinense 1841 [s.n.] (LE [image!]; isolectotypes: B destroyed [image negative 18452 at F!], HAL barcode HAL098686!, K barcode K000449124!, US barcode US282538 [image!]).
Note — Bignonia brachypoda DC. is illegitimate under Art. 52.1 of the International Code of Nomenclature for algae, fungi, and plants (Turland & al. 2018, hereafter called “Code”) because Candolle (1845) included under the simultaneously published B. brachypoda var. platyphylla (Cham.) DC. the earlier species name B. platyphylla Cham. Because var. platyphylla is a subordinate taxon that did not include the intended type of B. brachypoda, the type of the latter name is not automatically that of B. platyphylla (Art. 7.6). Arrabidaea brachypoda Bureau (1894) is a legitimate replacement name based on the illegitimate B. brachypoda (Art. 6.12 and 58.1). Bureau's name is not likewise illegitimate under Art. 52.1 because in 1894 the epithet platyphylla was not available for a new combination in Arrabidaea, having already been used in the heterotypic A. platyphylla DC. in 1845 (Art. 58 Note 1). We have selected as the lectotype of B. brachypoda the specimen Sellow s.n. deposited in LE, which is also the holotype of B. platyphylla, the two names thereby becoming homotypic synonyms.
= Bignonia brachypoda var. acutifolia DC. in Candolle, Prodr. 9: 145. 1845. – Holotype: Brazil, Mato Grosso, Cuiabá, fl., 1832, da Silva Manso 70A [s.n.] (G-DC barcode G00133313 [image!]).
= Bignonia brachypoda var. cuneata DC. in Candolle, Prodr. 9: 145. 1845 ≡ Arrabidaea brachypoda var. cuneata (DC.) Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 98. 1894 ≡ Arrabidaea platyphylla var. cuneata (DC.) Bureau & K. Schum., Fl. Bras. 8(2): 40. 1896. – Holotype: Brazil, Mato Grosso, Cuiabá, fl., 1832, da Silva Manso s.n. (G-DC barcode G00133320 [image!]).
= Bignonia brachypoda var. elliptica DC. in Candolle, Prodr. 9: 145. 1845 ≡ Arrabidaea platyphylla var. elliptica (DC.) Bureau & K. Schum., Fl. Bras. 8(2): 40. 1896. – Holotype: Brazil, Mato Grosso, Cuiabá, fl., 1832, da Silva Manso s.n. (G-DC barcode G00133316 [image!]).
= Bignonia brachypoda var. firmula DC. in Candolle, Prodr. 9: 146. 1845 ≡ Arrabidaea platyphylla var. firmula (DC.) Bureau & K. Schum., Fl. Bras. 8(2): 40. 1896. – Lectotype (designated here): Brazil, Minas Gerais, Serro Frio, fl., 1833, M. Vauthier 250 (G-DC barcode G00133322 [image!]; isolectotypes: G-DC barcode G00008804 [image!], P barcode P03586005!, W herbarium accession number W-0057543!).
Fig. 1.
Fridericia platyphylla – A: habit of a sclerophyllous plant with 1-foliolate leaves with orbicular leaflets; B: branch of sclerophyllous plant with 2-foliolate leaves and elliptic leaflets; C: leaflet of a mesophyllous plant showing elliptic shape and shortly acuminate apex; D: leaflet of a sclerophyllous plant showing elliptic shape and retuse apex; E: calyx cupular, costate near rim; detail showing 5-acuminate calyx rim; F: open flower in longitudinal section (staminode is omitted for clarity); G: ovary and disk; H: mature fruit; I: seed. – A, E, F, G from Hatschbach & Koczicki 33052 (MBM); B from Irwin & al. 23926 (UB); C from Hage & al. 2284 (MBM); D, H, I from Queiroz & al. 4712 (MBM). – Drawn by Diana Carneiro.
Note — Four syntypes were listed in the protologue of Bignonia brachypoda var. firmula DC. Two of these are gatherings from Minas Gerais (Vauthier 250 [s.n.] and Martius s.n.), one is a gathering from Bahia (Martius s.n.) and the other is another gathering by Martius s.n. with fruits deposited in Munich. We have selected one of the two specimens of Vauthier's gathering deposited in G-DC (G00133322) as the lectotype because it is well preserved and was identified by Candolle.
= Bignonia brachypoda var. heterophylla DC. in Candolle, Prodr. 9: 146. 1845 ≡ Arrabidaea platyphylla var. heterophylla (DC.) Bureau & K. Schum., Fl. Bras. 8(2): 40. 1896. – Holotype: Brazil, São Paulo, São Carlos, fl., 1834, P. W. Lund 1879 (G-DC barcode G00133364 [image!]).
= Bignonia violacea DC. in Candolle, Prodr. 9: 156. 1845, syn. nov. – Holotype: Bolivia, Santa Cruz, Concepción, fl., Jul, A. C. V. D. d'Orbigny 669 [s.n.] (P barcode P03586052!).
Note — We synonymize the previously unplaced name Bignonia violacea DC. under Fridericia platyphylla. We agree with Bureau (in sched.), who identified the holotype in Paris (P03586052) as Bignonia brachypoda var. elliptica, here considered as a synonym of F. platyphylla.
= Bignonia regnelliana Sond. in Linnaea 22: 558. 1849. – Lectotype (designated here): Brazil, Minas Gerais, fl., s.d., A. F. Regnell I-295 (S herbarium accession number S04-3501!; isolectotypes: BM barcode 000848555 [image!], P barcode P03585983!).
Note — The protologue of Bignonia regnelliana Sond. does not provide any information about where the type was deposited, nor about where the holotype was collected. Four specimens of Regnel I-295 were located, three of which do not contain any evidence that Sonder studied them. The first specimen is deposited in P and includes Regnell's handwritten note “Ad Caldas in Provincié Mina Ger. Brasl [sic]” in the label. The second is deposited in BM and includes the same locality of the specimen deposited in P and the species name handwritten by Regnell. The third is deposited in MO (MO-083287) and includes two labels. One of those labels includes two notes handwritten by Regnell using two different pen colours as follows: (1) “Entre M C Lopes e a Capella – illegible – Sta Rita” and (2) “Ad Beau 1867 I.295.” The second label was typed within a printed label from the Uppsala herbarium that describes the collection locality as “entre Lopes Jaguary e St. Rita” and the collection date as “24 Apr 1862”. The fourth specimen held at S includes an identification label handwritten by Regnell and a handwritten note by Sonder indicating the collection locality as “Provincia Minas Geraes [sic] Am. [America] Regnell”. Because the latter specimen is well preserved and is the only one that clearly indicates that it was analysed by Sonder, we have selected it (S04-3501) as the lectotype. Following Art. 9.4(a) of the Code, the specimen deposited in MO (MO-083287) does not represent an isolectotype because the collection date handwritten by Regnell is subsequent to the publication of B. regnelliana, indicating that this specimen does not represent original material.
= Petastoma simplicifolium Miers in Proc. Roy. Hort. Soc. London 3: 194. 1863. – Lectotype (designated here): Brazil, São Paulo and Rio [São Paulo], fl., 1861-2 [s.d.], J. Weir 204 (K barcode K000990553!; isolectotype: K barcode K000990524! [i.e. only the branch on the left of the sheet]).
Note — The protologue of Petastoma simplicifolium Miers does not indicate the herbarium where the type material was deposited. Two specimens of Weir 204 were found at K. Because one of them (K000990524) is mounted on the same sheet as a specimen from different gathering, Correa de Mello 2 (K000990533), we have selected the other specimen (K000990553) as the lectotype. The selected material is well conserved allowing for accurate species identification.
= Arrabidaea brachypoda var. acuminata Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 98. 1894 ≡ Arrabidaea platyphylla var. acuminata (Bureau) Bureau & K. Schum., Fl. Bras. 8(2): 39. 1896. – Lectotype (designated here): Brazil, Minas Gerais, Lagoa Santa, fl., 16 Feb [“floret Februario ad Aprilem”], E. Warming s.n. (C barcode C10021771 [image!]; isolectotype: C barcode C10021772 [image!]).
Note — Bureau (1894) indicated two syntypes for Arrabidaea brachypoda var. acuminata Bureau in the protologue: Warming s.n., collected at Lagoa Santa in Minas Gerais, and Glaziou 7775, collected in Rio de Janeiro. We have selected the specimen Warming s.n. (C10021771) as the lectotype because the description of the flowers in the label matches perfectly the description provided in the protologue.
= Arrabidaea brachypoda var. attenuata Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 98. 1894. – Lectotype (designated here): Brazil, Minas Gerais, Lagoa Santa, fl., s.d., E. Warming 1961/3 [s.n.] (C barcode C10021767 [image!]).
Note — Bureau (1894) did not indicate where the type of Arrabidaea brachypoda var. attenuata Bureau was deposited. We found three gatherings of Warming with this determination made by Bureau deposited in C. We have selected the specimen Warming 1961/3 (C10021767) as the lectotype because the habitat and plant description match the protologue. This gathering has a collecting number very similar to the number of the lectotype of A. brachypoda var. rigida Bureau (Warming 1961/2, C10021764). Because the specimen C10021764 includes a distinct collection date, this material is not a duplicate. Among the two remaining specimens identified by Bureau as A. brachypoda var. attenuata, C10031766 does not include the collecting number and C10021765 has a different number (Warming 1965). Therefore, these specimens are also not isolectotypes.
= Arrabidaea brachypoda var. firma Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 98. 1894. – Lectotype (designated here): s. loc. [Lagoa Santa], fl., s.d., A. Glaziou 12970 (C barcode C10021773 [image!]; isolectotypes: P barcodes P03586027!, P03586028!).
= Note — Bureau (1894) listed Arrabidaea brachypoda var. firmula DC. “(partim)” as a synonym of A. brachypoda var. firma Bureau. In the protologue, he cited two syntypes, the gatherings Warming s.n. and Glaziou 12970; neither was cited by Candolle (1845). Following Art. 52.1 of the Code, because Bureau indicated “partim”, the type of A. brachypoda var. firmula was not definitely included in A. brachypoda var. firma, resulting in his varietal name being legitimate. Among the two gatherings mentioned by Bureau (1894), we found only three specimens of Glaziou 12970, one in C without locality and date, and two in P, one of them with Glaziou's handwritten indication that the gathering was made in Ipanema on 23 April 1881. The specimens in P were donated to the herbarium after Glazious's death in 1907 (Bureau 1908) and, therefore, after the publication of the new varietal name. Despite lacking detailed gathering information, we have selected C10021773 as the lectotype because it is a well-preserved specimen and was annotated by Bureau.
= Arrabidaea brachypoda var. induta Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 98. 1894 ≡ Arrabidaea platyphylla var. induta (Bureau) Bureau & K. Schum., Fl. Bras. 8(2): 39. 1896. – Lectotype (designated here): s. loc. [“Brasilia australiore”], fl., Feb 1882 [s.d.], A. Glaziou 12992 (C barcode C10021775 [image!]; isolectotypes: BR barcode 880546 [image!], C barcode C10021774 [image!], K [barcode K000449128!], MO herbarium accession numbers MO-083153!, MO-083247!, P barcodes P03586029!, P03586030!).
Note — Bureau (1894) did not mention where the type of Arrabidaea brachypoda var. induta Bureau was deposited. We were able to locate eight specimens of Glaziou 12992, one deposited in each of BR and K and two deposited in each of C, MO and P. The specimens deposited in BR and K do not have Bureau's handwriting, while the two deposited in MO are sterile. The two deposited in P have the varietal name annotated by Bureau in a label that indicates that these specimens were donated to P by Glaziou's daughter in 1907, after Glaziou's death (Bureau 1908), subsequent to the publication of A. brachypoda var. induta. We opted to select as the lectotype the only specimen from Warming's herbarium held in C (C10021775), which has Bureau's handwriting and is very well preserved.
= Arrabidaea brachypoda var. rigida Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 98. 1894. – Lectotype (designated here): Brazil, Minas Gerais, Lagoa Santa, fr., 16 Dec 1863 [s.d], E. Warming 1961/2 [s.n.] (C barcode C10021764 [image!]).
Note — Bureau (1894) did not indicate where the type of Arrabidaea brachypoda var. rigida Bureau was deposited. We were only able to locate one specimen of a Warming gathering determined by Bureau with this name (C10021764). This material is a fruiting specimen collected in December. Even though the protologue describes corolla colour and not the fruit, it indicates “October to February” as the flowering period and “December to January” as the fruiting period. Because this was the only Warming specimen annotated by Bureau as A. brachypoda var. rigida, we have selected it as the lectotype.
= Arrabidaea celastroides Bureau ex K. Schum. in Engler & Prantl, Nat. Pflanzenfam. 4(3b): 213. 1894, syn. nov. – Lectotype (designated by Lohmann in Lohmann & Taylor 2014: 434): Brazil, Bahia, Jacobina, fl., 1841 [s.d.], J. S. Blanchet 3419 (G-DC barcode G00008903 [image!]; isolectotypes: BR barcode BR8805393 [image!], FI barcode FI009859 [image!], G-DC barcode G00133799 [image!], P barcodes P00468568!, P00468569!).
Note — Arrabidaea celastroides Bureau ex K. Schum. is here proposed as a new synonym of Fridericia platyphylla. The description provided in the protologue of A. celastroides is very concise, mentioning only the glabrous, orbicular or oblong leaflets. The protologue has no mention of any specimen (Schumann, 1894). In the Flora brasiliensis, Bureau & Schumann (1896) segregated this species based on the shrubby habit, “herbacea” leaves, flowers smaller than 2 cm and denticulate calyx rim. On the other hand, A. platyphylla (Cham.) Bureau & K. Schum. (non A. platyphylla DC.) is characterized by the shrubby or scandent habit, coriaceous leaves, flowers larger than 2.5 cm and almost denticulate calyx rim. Even though the type materials of both species represent the extremes of their variation, the pattern of morphological variation is continuous. Molecular phylogenetic data (Kaehler & al. 2019) provide additional support for the recognition of a single taxon.
= Arrabidaea platyphylla var. lasiorrhachis Bureau & K. Schum., Fl. Bras. 8(2): 40. 1896. – Lectotype (designated here): Brazil, São Paulo, Minas Gerais and Goiás [“Brasilia australiore”], fl., Jul–Oct 1877 [s.d.], Vic. P. Seguro 59 [s.n.] (W barcode W0057598!).
Note — Bureau & Schumann (1896) did not specify where the type of Arrabidaea platyphylla var. lasiorrhachis Bureau & K. Schum. (Vic. P. Seguro s.n.) was deposited. We have selected as the lectotype the only specimen located (W0057598) with an identification clearly made by Schumann.
= Arrabidaea platyphylla var. puberula Bureau & K. Schum., Fl. Bras. 8(2): 41. 1896. – Lectotype (designated here): s. loc. [“in Brasilia occidentali”], fl., s.d., Tamberlik s.n. (NY barcode NY720025!; isolectotypes: MO herbarium accession number MO-085876!, W herbarium accession number W-57550!). Note — Four syntypes were listed in the protologue of Arrabidaea platyphylla var. puberula Bureau & K. Schum. (the gatherings Pohl 1105, Vic. Porto Seguro s.n., Sellow 5511 and Tamberlik s.n.). Of these, only three specimens of Tamberlik s.n. were located, all of which included Schumann's handwriting in the label. The specimen with the highest quality (NY720025) has been selected as the lectotype.
– “Fridericia platyphylla” (Cham.) L. G. Lohmann in Monogr. Syst. Bot. Missouri Bot Gard. 107: 1606. 2008. Designation not validly published (Art. 41.1).
– “Fridericia platyphylla” (Cham.) L. G. Lohmann in Forzza & al., Cat. Pl. Fung. Brasil 1: 766. 2010. Designation not validly published (Art. 41.1).
Morphological description — Lianas, shrubs (sometimes small trees), up to 20 m tall. Branches terete, lenticellate, grey or brown when dry, glabrescent to velutinous with simple trichomes (sometimes branched) and sparsely lepidote with peltate glandular trichomes, without interpetiolar ridges, with interpetiolar glandular fields; prophylls of axillary buds triangular, arranged in 2 rows, 1–2 mm long. Leaves 1- or 2-foliolate; petiole 0.4–2 cm long, glabrescent to velutinous with simple trichomes (rarely branched); petiolules of equal lengths, when 1-foliolate 0.5–1(–1.3) cm, when 2-foliolate 1–2.2 cm, glabrescent to velutinous with simple trichomes (rarely branched); blade chartaceous to coriaceous, margin entire or revolute, central leaflet elliptic, orbicular or obovate (sometimes ovate), lateral leaflets elliptic or orbicular (sometimes ovate), 4–16 × 1.7–13.5 cm, base rounded or cuneate, apex rounded, shortly acuminate, acute or emarginate, surface adaxially lepidote with glandular peltate trichomes, glabrescent to velutinous with simple trichomes, abaxially lepidote with glandular peltate trichomes, glabrescent to velutinous with simple trichomes, venation acrodromous, secondary veins raised, tertiary veins flat or slightly raised, domatia as vestigial pockets and trichome tufts. Inflorescence terminal (rarely axillary), a thyrse, with 2 or 3 orders; peduncle of first order 2–8.3 cm long, of second order 1.2–3.4 cm long, of third order 0.3–0.7 cm long, pubescent to velutinous with simple trichomes (sometimes branched); bracts linear, 1–2.5 mm long, caducous; bracteoles linear, 0.6–0.8 mm long, caducous; pedicels 0.3–0.6 cm long. Calyx cupular, costate throughout (sometimes only near rim), 5-acuminate (sometimes shortly split), 0.3–0.6 × 0.2–0.5 cm, pubescent with simple trichomes (rarely with branched trichomes), chartaceous, white, pink or purplish, with glands close to rim. Corolla infundibuliform, zygomorphic, slightly furrowed, 2.4–5.1 cm long, 0.9–1.3 cm wide at tube mouth, externally pubescent with simple and glandular peltate trichomes, without glands, internally glabrous, villose at base and close to base of stamens, with glandular capitate trichomes, pubescent at lobes with simple and glandular capitate trichomes, lobes 0.7–1 × 0.8–1.2 cm, pink. Androecium with stamens included; longer filaments 1–1.3 cm long, shorter filaments 0.8–1 cm long; staminode c. 0.3 cm long; anthers c. 2 mm long, connectives not protruding. Gynoecium with ovary cylindric, 1.8–2.7 × 0.7–1 mm, smooth, lepidote; style 1.6–2.3 cm long; stigma elliptic to rhomboid; nectar disk plate-shaped, 0.6 –1 × 1.1–2 mm. Fruit linear, flat, margin not raised, with or without a central ridge; valves coriaceous, smooth, 11–33 × 0.8–1.4 cm, lepidote with glandular peltate trichomes; septum coriaceous. Seeds oblong or elliptic, 0.7–1.3 × 2–4 cm, body oblong or elliptic, wings hyaline, 0.2–0.4 cm wide, margin entire.
Phenology — Produces flowers and fruits all year long.
Distribution and habitat — Fridericia platyphylla is broadly distributed through South America. This species occupies dry and wet Amazonian forests, Atlantic rainforest, savannahs (including Cerrado, Llanos and Guiana Highlands), Chaco and Caatinga. It is distributed through Bolivia (El Beni, La Paz and Santa Cruz), Brazil (Acre, Amazonas, Bahia, Ceará, Distrito Federal, Espírito Santo, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Paraíba, Paraná, Piauí, Rondônia, Roraima, São Paulo, Rio de Janeiro and Tocantins), Guyana (Upper Takutu-Upper Essequibo), Paraguay (Amambay, Caazapá, Central, Canindeyú and Paraguarí), Peru (Cusco, Huánuco, Junín, Loreto, San Martín and Ucayali) and Venezuela (Amazonas, Bolívar, Guárico and Portuguesa).
Conservation status — Fridericia platyphylla is categorized as Least Concern (LC) based on its Extent of Occurrence (29 120 298 km2) and Area of Occupancy (502 500 km2).
Remarks — Fridericia platyphylla is easily recognized by its 1- or 2-foliolate leaves and cupular calyx, costate when dry. An infraspecific classification was proposed by Candolle (1845) based on leaf morphology. Indeed, the leaves show wide variation, usually being 2-foliol-ate and chartaceous in moist and forested environments, where the plants are usually lianas, but 1-foliolate and coriaceous in dryer and open vegetation, where the plants are shrubs, scandent shrubs or treelets. Despite the variation found in the leaf morphology, the flower and fruit morphologies are constant, leading us to disregard the infraspecific classification. Additionally, some specimens (e.g. Giulietti & al. 1845, MBM; Hassler 10576, NY, P; Krapovickas & al. 14077, MBM) were shown to bear sclerophyllous leaves in older branches and mesophyllous leaves in younger branches, indicating that leaf morphology can vary even within the same individual.
Two distinct species were described in different genera with the specific epithet “platyphylla”: Bignonia platyphylla DC. and Arrabidaea platyphylla Cham. The correct name of the species treated here was carefully explained by Lohmann & Taylor (2014). However, the name Arrabidaea platyphylla DC. [= Xylophragma platyphyllum (DC.) L. G. Lohmann] is often mistakenly applied to Fridericia platyphylla (Cham.) L. G. Lohmann. Xylophragma platyphyllum can be easily identified by the leaves with venation darker than the blade when dry (vs. never darker when dry in F. platyphylla), corolla tubular (vs. corolla infundibular in F. platyphylla), smaller than 1.2 cm (vs. 2.2–5.1 cm in F. platyphylla), nectar disk surrounding the ovary (vs. nectar disk under the ovary in F. platyphylla) and branched trichomes (vs. simple trichomes in F. platyphylla; only the specimen Irwin 13141 seen in MO, NY, P and UB has branched trichomes on the inflorescence axis).
2. Fridericia prancei (A. H. Gentry) L. G. Lohmann in Ann. Missouri Bot. Gard. 99: 443. 2014 ≡ Arrabidaea prancei A. H. Gentry in Ann. Missouri Bot. Gard. 63: 50. 1976. – Holotype: Brazil, Amazonas, km 67 E of Manaus–Itacoatiara road, fl., Nov 1974, A. H. Gentry 12882 (INPA not seen; isotypes: MO herbarium accession number MO-2251712!, RB barcodes RB00536870!, RB00538142!, RB00538143!, TEPB herbarium accession number TEPB-4488!). – Fig. 2, 3A–C.
– “Fridericia prancei” (A. H. Gentry) L. G. Lohmann in Forzza & al., Cat. Pl. Fung. Brasil 1: 766. 2010. Designation not validly published (Art. 41.1).
Morphological description — Lianas, up to 25 m tall. Branches terete, lenticellate, brown or black when dry, lepidote with glandular peltate trichomes; without interpetiolar ridges, with interpetiolar gland fields; prophylls of axillary buds triangular, arranged in 2 rows, 1–2.3 mm long. Leaves 2-foliolate; petiole 1.2–2.7(–3.5) cm long, glabrescent with simple trichomes and lepidote with glandular peltate trichomes; petiolules of equal lengths, (0.8–)2–3.6 cm long, glabrescent with simple trichomes and lepidote with glandular peltate trichomes; blade elliptic, coriaceous, margin revolute, 10.5–19 × 5.5–9 cm, base rounded or cuneate, apex acuminate, surface adaxially glabrous or lepidote with glandular peltate trichomes, abaxially glabrous or pubescent with simple trichomes only on veins and lepidote with glandular peltate trichomes, venation acrodromous, secondary veins raised, tertiary veins slightly raised, domatia as vestigial pockets and trichome tufts. Inflorescence axillary or terminal, a thyrse, with 3 or 4 orders; peduncle of first order 0.7–1.1(–4.2) cm long, of second order 0.5–1.4 cm long, of third order 0.4–0.6 cm long, of fourth order 0.3–0.4 cm long, lepidote with glandular peltate trichomes; bracts linear, 1.2–2.5 mm long, persistent; bracteoles linear, 0.9–1.3 mm long, caducous; pedicels 0.6–1.1 cm long. Calyx tubular, not costate, truncate to shortly 5-acuminate, 0.8–1.1 × 0.3–0.5 cm, lepidote with glandular peltate trichomes (sometimes pubescent with simple trichomes at rim), coriaceous, white or cream, with glands clustered close to acumens. Corolla narrowly infundibuliform, zygomorphic, not or slightly furrowed, 5–6.2 cm long, 0.6–1.2 cm wide at tube mouth, externally pubescent with simple and glandular peltate trichomes, without glands, internally glabrous, villose at base and close to base of stamens with glandular capitate trichomes, pubescent at lobes with simple and glandular capitate trichomes, lobes 1.2–1.6 × 0.8–1.3 cm, white, pinkish white or lavender outside, white inside. Androecium with stamens at same height as tube mouth; longer filaments 1.7–1.9 cm long, shorter filaments 1.3–1.6 cm long; staminode 0.7–0.9 cm long; anthers c. 3 mm long, connectives not protruding. Gynoecium with ovary cylindric, 3.1–3.5 × c. 1 mm, furrowed, lepidote; style 2.8–3 cm long; stigma elliptic to rhomboid; nectar disk plate-shaped, c. 0.8 × 1.4 mm. Fruit linear, flat, margin not raised, without a central ridge; valves leathery, smooth, 34–40 × 1.3–1.7 cm, glabrous and sparsely lepidote with glandular peltate trichomes; septum coriaceous. Seeds oblong or elliptic, 1.1–1.5 × 4.2–4.9 cm, body oblong or elliptic, wings hyaline, 0.1–0.2 cm wide, margin entire.
Fig. 2.
Distribution of the species of the “Acrodromous venation” clade: Fridericia platyphylla (black circles); F. prancei (open diamonds).
Phenology — Produces flowers all year long. Fruiting specimens were collected in July, August and November.
Distribution and habitat — Fridericia prancei is distributed in lowland Amazonian rainforest in Venezuela (Amazonas), Guyana (Upper Takutu-Upper Essequibo) and Brazil (Acre, Amazonas, Pará, Rondônia and Roraima).
Fig. 3.
A–C: Fridericia prancei; A: habit; detail showing densely lenticellate stem; B: flower; C: fruit. – D–H: Fridericia erubescens; D: habit; E: leaf; F: flower; G: fruit; H: fruit cross-section showing large wings. – A from Steward & al. 7 (NY); B from Gentry 13055 (MO); C from Oliveira 4665 (NY); D–F from Queiroz & al. 7023 (UPCB); G–H from Anderson 9151 (MO). – Drawn by Diana Carneiro.
Conservation status — Fridericia prancei is categorized as Least Concern (LC) based on its Extent of Occurrence (1 876 438 km2) and Area of Occupancy (47 500 km2).
Remarks — Fridericia prancei can be easily identified because of the large and coriaceous leaves with marked suprabasal acrodromous venation, tubular, not costate calyx, long and narrowly infundibuliform, white or whitish pink corolla, and stamens at the same height as the tube mouth.
Al Gentry annotated “12822” over his collector's number typed (12882) on the isotype deposited in MO. However, he did not mention the differential notation in the protologue or in any other published work. Lohmann and Taylor (2014) cited the number handwritten by Gentry (in sched.). Since the protologue and all remaining isotypes found have the number 12882, we consider this to represent the correct collecting number.
The “Piriadacus” clade
The “Piriadacus” clade is composed of two species that share axillary buds with concentrically disposed prophylls (seen at the base of the inflorescences in Fridericia ornithophila (A. H. Gentry) L. G. Lohmann), inflorescences and calyx with capitate glandular trichomes, tubular, red, pink-reddish or orange corollas and exserted or sub-exserted stamens. The name of the clade is based on the mono-specific genus Piriadacus Pichon, which has Bignonia erubescens DC. (= F. erubescens (DC.) L. G. Lohmann) as the type (Kaehler & al. 2019).
3. Fridericia erubescens (DC.) L. G. Lohmann in Ann. Missouri Bot. Gard. 99: 437. 2014 ≡ Bignonia erubescens DC. in Candolle, Prodr. 9: 157. 1845 ≡ Cuspidaria erubescens (DC.) Bureau in Vidensk. Meddel. Dansk. Naturhist. Foren. Kjøbenhavn 1893: 102. 1894 ≡ Piriadacus erubescens (DC.) Pichon in Bull. Soc. Bot. France 92: 225. 1945 ≡ Alsocydia erubescens (DC.) J. C. Gomes in Revista Brasil. Biol. 11: 49. 1951. – Lectotype (designated by Lohmann in Lohmann & Taylor 2014: 437): Brazil, Minas Gerais, Serro Frio, fl., Jul [s.d.], C. F. P. von Martius s.n. [herb. 342] (M barcode M0088919!). – Fig. 3D–H, 4.
= Bignonia erubescens var. breviflora DC. in Candolle, Prodr. 9: 158. 1845. – Lectotype (designated here): Brazil, Minas Gerais [in Sylvis Capoes deserti], fl., Jul 1818 [s.d.], C. F. P. von Martius s.n. [herb. 341] (M barcode M0088918!).
= Bignonia erubescens var. subtruncata DC. in Candolle, Prodr. 9: 157. 1845. – Lectotype (designated here): Brazil, Minas Gerais, fl., s.d., C. F. P. von Martius s.n. [herb. 343] (M barcode M0088920!).
Note — Bignonia erubescens DC. includes three varieties, but the types of the species and varietal names were not directly designated by Candolle (1845). All three types are Martius's gatherings conserved in Munich. Lohmann (in Lohmann & Taylor 2014) chose the specimen M0088919 as the lectotype of B. erubescens with M0088918 and M0088920 as isolectotypes. However, the two varietal names also need lectotypification. The specimen M0088918 has the same locality description provided in the protologue of B. erubescens var. breviflora DC., as well as a label with a comment from Candolle on the small size of the flowers. Therefore, we have chosen M0088918 as the lectotype of this varietal name. Lastly, the label of M0088920 includes Candolle's handwriting and the specimen matches well the description of B. erubescens var. subtruncata DC. provided in the protologue. Therefore, we have chosen this specimen as the lectotype of this varietal name. Specimens M0088918 and M0088920 are not isolectotypes of B. erubescens, but lectotypes of the two varietal names described in the same treatment.
= Cuspidaria erubescens var. glabrescens Bureau in Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1893: 103. 1894. – Lectotype (designated here): Brazil, s. loc. [Minas Gerais], fl., s.d., A. Glaziou 14106 (C barcode C10021781 [image!]; isolectotypes: K barcode K000449476!, P barcode P02885020!).
Note — Bureau (1894) did not cite where the type of Cuspidaria erubescens var. glabrescens Bureau was deposited. We were only able to locate three specimens, none of which was collected in the locality provided in the protologue. We have chosen as the lectotype the only specimen annotated by Bureau that does not have a locality description but whose morphology matches the protologue (leaves 1- or 2-foliolate, whereas the specimens K000449476 and P02885020 have leaves 3-foliolate).
– “Fridericia erubescens” (DC.) L. G. Lohmann in Forzza & al., Cat. Pl. Fung. Brasil 1: 765. 2010. Designation not validly published (Art. 41.1).
Morphological description — Lianas or scandent shrubs, up to 5 m tall. Branches terete, striate and sparsely lenticellate, brown or black when dry, glabrescent to villose with simple and glandular capitate trichomes, with or without inconspicuous interpetiolar ridges, with or without interpetiolar glandular fields; prophylls of axillary buds broadly triangular, arranged as a bromeliad, 1.8–2.1 mm long. Leaves 2- or 3-foliolate; petiole 0.8–2.4 cm long, glabrescent to villose with simple trichomes (sometimes glandular capitate); petiolules in two sizes, central ones 0.7–2.2 cm long, lateral ones 0.2–0.5 cm long, glabrescent to villose with simple trichomes (sometimes glandular capitate); blade chartaceous to coriaceous, margin revolute, central leaflet elliptic, (2.1–)3–7.1(–11.5) × 3–6.4 cm, base rounded or cuneate, apex rounded or acuminate, lateral leaflets elliptic or obovate, 2.4–5.8(–10.6) × 1.2–5.7 cm, base rounded or cuneate, apex rounded or shortly acuminate, surface adaxially lepidote with glandular peltate trichomes, glabrescent to velutinous with simple trichomes, abaxially lepidote with glandular peltate trichomes, glabrescent to velutinous with simple trichomes, venation pinnate, secondary veins raised, tertiary veins raised, without domatia (sometimes with trichome tufts). Inflorescence terminal or axillary, thyrsoid, with 3 orders; peduncles of first order (1.6–)3–5.9 cm long, of second order 1.8–2.9 cm long, of third order 0.9–1.5 cm long, glabrescent to villose with simple and glandular capitate trichomes; bracts linear, caducous or persistent, 0.5–1 cm long; bracteoles linear, 0.3–0.4 cm long, caducous or persistent; pedicels 0.2–0.8 cm long. Calyx tubular, not costate, irregularly split or truncate, 0.6–1.3 × 0.3–0.4 cm, glabrescent to villose with simple and glandular capitate trichomes, membranous or chartaceous, green, pinkish or purplish, without glands. Corolla tubular, zygomorphic, not furrowed, (2.4–)3–5(–5.7) cm long, 0.5–0.6 cm wide at tube mouth, externally pubescent with simple and glandular peltate trichomes, without glands, internally glabrous, villose at base and close to base of stamens with glandular capitate long-stalked trichomes, pubescent at lobes with simple and glandular capitate trichomes, lobes 0.4–1.1 × 0.3–0.5 cm, orange, reddish or pinkish. Androecium with stamens exserted; longer filaments 2.9–4.3 cm long, shorter filaments 1.7–3.3 cm long; staminode 0.3–0.4 cm long; anthers 3–4 mm long, connectives protruding 0.6–0.9 mm. Gynoecium with ovary cylindric, 3–4 × 1–1.2 mm, furrowed, lepidote (sometimes glabrous); style 3.1–4.9 cm long; stigma elliptic to rhomboid; nectar disk plate-shaped, c. 1.2 × 2.6 mm. Fruit linear, flat, margin winged, with or without a central ridge; valves woody, smooth, 13–22(–27) × 2.9–3.5(–5.3) cm, glabrous; septum coriaceous. Seeds oblong to semicircular, 0.4–1 × 1.9–4 cm, body of seeds semicircular, wings hyaline, 0.3–0.4 cm wide, margin entire.
Fig. 4.
Distribution of the species of the “Piriadacus” clade. Fridericia erubescens (black circles); F. ornithophila (open diamonds).
Phenology — Produces flowers and fruits all year long.
Distribution and habitat — Fridericia erubescens occurs in Caatinga, Cerrado and ecotone areas of Atlantic rainforest. It is endemic to Brazil (Bahia, Ceará, Espírito Santo, Minas Gerais, Pernambuco and Piauí).
Conservation status — Fridericia erubescens is categorized as Least Concern (LC) based on its Extent of Occurrence (1 206 403 km2) and Area of Occupancy (307 500 km2).
Remarks — Fridericia erubescens is easily recognized by the tubular calyx, frequently covered by glandular capitate trichomes giving a sticky feel, the tubular and long corolla, usually red, orange or pinkish orange, the exserted stamens and winged fruit. Some species of Cuspidaria and Tynanthus Miers also have winged fruits, but the mature fruits of F. erubescens are longer (reaching 30 cm) and the leaves are often 3-foliolate. On the other hand, Cuspidaria and Tynanthus have shorter fruits (around 20 cm) and the leaves are usually 2-foliolate.
The placement of this species has long been confusing. Candolle (1845) described it in Bignonia based on flower morphology, because the fruit was unknown at that time. Bureau (1894) transferred it into Cuspidaria without providing any justification. The species was treated in Cuspidaria until Pichon (1945) described the new genus Piriadacus Pichon (an anagram of the name Cuspidaria), with Fridericia erubescens as the type. According to Pichon (1945), the tubular calyx, glabrous anthers, pollen organized in monads with a granulose exine pattern and fruits with large wings were not shared with the remaining species of Cuspidaria. Gomes (1951) described the genus Alsocydia Mart. ex J. C. Gomes with B. erubescens DC. as type, possibly unaware of Pichon's work. His classification was based on “simple pollen” (i.e. in monads), which is distinct from Cuspidaria with its pollen organized in tetrads. Finally, molecular evidence led Lohmann (in Lohmann & Taylor 2014) to transfer this species into Fridericia.
4. Fridericia ornithophila (A. H. Gentry) L. G. Lohmann in Ann. Missouri Bot. Gard. 99: 441. 2014 ≡ Arrabidaea ornithophila A. H. Gentry in Ann. Missouri Bot. Gard. 64: 312. 1977 [“1978”]. – Holotype: Brazil, Pará, Tomé Açú, Pão Vermelho, fl.-bud, fl., 3 Aug 1931, Y. Mexia 6041 (MO herbarium accession number MO-1068874!; isotypes: A barcode 00091926 [image!], GH barcode 00091927 [image!], NY barcode 00346064!, S herbarium accession number S09-21565 [image!], US barcode 00170304 [image!]). Fig. 4, 5.
– “Fridericia ornithophila” (A. H. Gentry) L. G. Lohmann in Forzza & al., Cat. Pl. Fung. Brasil 1: 766. 2010. Designation not validly published (Art. 41.1).
Morphological description — Lianas or scandent shrubs, up to 20 m tall. Branches terete, striate and sparsely lenticellate, grey to brown when dry, puberulous to velutinous with simple trichomes, with inconspicuous or without interpetiolar ridges, with interpetiolar glandular fields; prophylls of axillary buds triangular, arranged in 2 rows (sometimes concentrically disposed at inflorescence peduncles), c. 1 mm long. Leaves 3-foliolate; petiole (2.5–)4.2–6 cm long, puberulous with simple trichomes; petiolules in two sizes, central ones 1.4–4(–4.9) cm long, lateral ones 0.6–2.8 cm long, puberulous to velutinous with simple trichomes; blade chartaceous, margin entire, central leaflet elliptic, 10.2–16.5(–26) × 4–9.4(–16.9) cm, base rounded or cuneate, apex acuminate (sometimes retuse), lateral leaflets elliptic, oblong to slightly ovate, 9.1–15.2 × 4.5–11 cm, base rounded (sometimes cuneate), apex acuminate, surface adaxially glabrescent to puberulous with simple trichomes (sometimes only at mid vein) and lepidote with peltate trichomes, abaxially puberulous to velutinous throughout with simple trichomes, venation pinnate, secondary veins raised, tertiary veins raised, without domatia. Inflorescence terminal or axillary, thyrsoid, with 2–4(or 5) orders; peduncles of first order (3.2–)9–15 cm long, of second order 1.6–3.9 cm long, of third order 0.6–1.8 cm long, of fourth order 0.4–0.8 cm long, of fifth order c. 0.8 cm long, velutinous with simple and glandular capitate trichomes; bracts linear, 1–1.2 mm long, caducous; bracteoles linear, 0.8–1 mm long, caducous; pedicels 0.5–0.9 cm long. Calyx tubular, not costate, bilabiate (sometimes irregularly split), 1–1.6 × 0.3–0.5 cm, pubescent with simple and glandular capitate trichomes, thick, red, without glands. Corolla tubular, actinomorphic, not furrowed, 2.4–4 cm long, 0.4–0.6 cm wide at tube mouth, externally glabrescent to pubescent with simple and peltate trichomes, without glands, internally glabrescent with glandular capitate long stalked trichomes, villose at base with capitate glandular long-stalked trichomes, villose at lobes with simple trichomes, lobes 0.3–0.7 × 0.3–0.5 cm, red outside and white or yellow inside. Androecium with stamens at same height as tube mouth or slightly below; longer filaments c. 2.7 cm long, shorter filaments c. 2.5 cm long; staminode c. 0.4 mm long; anthers 3–4 mm long, connectives protruding 0.4–1 mm. Gynoecium with ovary cylindric, c. 2 × 0.8 mm, smooth, lepidote; style 2–2.6 cm long; stigma orbicular; nectar disk plate-shaped, c. 1 × 2–3.2 mm. Fruit not seen.
Fig. 5.
Fridericia ornithophila – A: habit; details showing interpetiolar gland fields (centre right), prophylls concentrically disposed at base of inflorescence (bottom left) and abaxial surface of leaves densely pubescent with protruding secondary venation (bottom right); B: second-order branch of inflorescence; C: flower showing narrowly tubular corolla and actinomorphic disposition of lobes; D: old flower without corolla showing bilabiate calyx and orbicular stigma. – A–D from Pires 605 (MO). – Drawn by Diana Carneiro.
Phenology — Produces flowers from March to November. Fruits unknown.
Distribution and habitat — Fridericia ornithophila occupies lowland oriental Amazonian rainforests. A single gathering (Teixeira & al. 803, INPA) was collected in ecotone with Cerrado. This species is distributed in Bolivia (El Beni) and Brazil (Acre, Amapá, Maranhão, Mato Grosso, Pará and Rondônia).
Conservation status — Fridericia ornithophila is categorized as Least Concern (LC) based on its Extent of Occurrence (1 820 317 km2) and Area of Occupancy (52 500 km2).
Remarks — Among the paratypes of Fridericia ornithophila, Gentry (1978) listed the sterile gathering Prance 58902 deposited in MO and NY. We found a fertile specimen of this gathering in UB that allowed us to identify it as F. trailii (Sprague) L. G. Lohmann. When sterile, this species is very similar to F. ornithophila due to the leaf size, shape and discolorous pattern. However, F. trailii has denser inflorescences with 4–6 branching orders without glandular capitate trichomes, the calyx rim is truncate and shortly 5-cuspidate, also lacking the glandular capitate trichomes, and the corolla is slender. On the other hand, F. ornithophila has a lax inflorescence with 2–4 branching orders covered with glandular capitate and simple trichomes, the calyx rim is bilabiate (sometimes irregularly split) with glandular capitate trichomes and the corolla is thick.
Acknowledgements
We thank the staff and curators of the herbaria HAL, K, M, MBM, MO, NY, P, R, RB, S, SPF, TEPB, UB and UPCB for specimen loans and assistance during herbarium visits; Olof Ryding (from C) and Larissa Orlova (from LE) for locating and providing high-quality photographs of selected specimens; Maria Cláudia M. P. Medeiros for photographs from Bignoniaceae specimens deposited in LE; Annelise Frazão, Jéssica Francisco and Ricardo da Silva Ribeiro for photographs of living plants; Diana Carneiro for the line drawings; Fabián A. Michelangeli for hosting M.K. as a visiting scholar at the New York Botanical Garden (NY); Jefferson Prado for assistance with nomenclature; and Maria Cláudia M. P. Medeiros, Richard Olmstead and Nicholas Turland for helpful comments on this manuscript. This work was supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) through a postdoctoral fellowship to M.K. (232621/2014-4) and a Pq-1B grant to L.G.L. (30778/2013-5); and the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) through two grants to L.G.L. (2011/50859-2 and 2012/50260-6).
References
Appendices
Supplemental content online
Appendix 1 (wi.51.51202_Appendix_1.pdf). Specimens examined (see https://doi.org/10.3372/wi.51.51202).
