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Published In: Histoire des Plantes de la Guiane Françoise 1: 102, pl. 40, f. 1. 1775. (Jun-Dec 1775) (Hist. Pl. Guiane) Name publication detailView in BotanicusView in Biodiversity Heritage Library

Project Name Data (Last Modified On 11/13/2017)
Acceptance : Accepted
Note : Tribe Coussareeae
Project Data     (Last Modified On 6/20/2021)

Faramea Aubl. includes approximately 170 species of tropical shrubs and small trees found in moist to wet vegetation in the Antilles and central Mexico through tropical South America. These species are characterized by generally opposite leaves in a distichous arrangement; sheathing or sometimes calptrate, triangular, aristate stipules; cymose, terminal or rarely axillary inflorescences of varied arrangement, from capitate and sessile to branched to several orders, and with bracts reduced or rarely well developed; distylous, 4-merous, fragrant flowers; white to blue or occasionally yellow, salverform, generally showy corollas with the lobes valvate in bud; a generally unilocular ovary with one basal ovule; and fleshy, oblate, blue or purple-black fruits with a single, relatively large seed or pyrene that has a papery to firm wall. Faramea is usually recognizable even without flowers or fruits, by the stipules with their well developed aristas crossed at the stem apex and in many species also leaves with a well developed, nearly straight submarginal vein. The peduncles are are often articulated, with a joint bearing two small stipules borne near its middle; presumably this indicates the inflorescence is a reduced cyme. The fruits of Faramea are generally fleshy, juicy, or leathery and have a characteristic shape, depressed-globose (i.e., oblate) and often also laterally flattened. The stems in Faramea are usually flattened and costate. Some species are characterized by well developed acarodomatia on the leaves. The most widespread and commonly collected species are Faramea occidentalis and Faramea multiflora, and other particularly common widespread species are Faramea anisocalyx and Faramea capillipes. The identity and circumscription of Faramea multiflora and some other widespread species were clarified by Taylor & Jardim (2020). 

Faramea is notable for its broad geographic range and its wide range of morphological variation. The plants range from subshrubs to trees and the leaves from small to quite large, and the inflorescences vary from solitary flowers to involucrate subcapitate groups and small to large, lax cymes. A few species show unusual features such as large calyx limbs and relatively large, laterally flattened, ellipsoid fruits. The inflorescences also apparently may be produced in variously terminal or axillary positions, though the axillary inflorescence could also be regarded as terminal inflorescences borne on short axillary stems as in some other genera (e.g., Ixora). Most authors have considered inflorescence position variable within species, though a few (e.g., Borhidi 2012) considerd this a consistent character and used it to separate species within otherwise morphologically indistinguishable populations. Several species of wet montane forests have triangular to rounded stipules with the aristas reduced, for whatever reason (e.g., Faramea flavicans, Faramea neblinae). Most Faramea species have distylous flowers or the common form in Rubiaceae, with the anthers and stigmas reciprocally place in the two forms with one of them exserted in each form. Taylor & Jardim (2020) detailed apparently another distylous arrangement in Faramea kampauicola, with the anthers positioned near the middle of the corolla tube in both forms and the stigmas either exserted or positioned below the anthers. Similar variation in the forms of distylous flowers is found in Palicourea (Taylor, 2017). 

One group of Faramea species shares leaves with weakly brochidodromous venation, oblate (but not laterally flattened) black fruits, and white flowers that are at lesat usually noctural. and a habitat in somewhat seasonal vegetation; this group includes Faramea occidentalis. Another distinctive group of species has leaves with well developed submarginal veins, fruits that are markedly laterally flattened and often blue, and blue diurnal flowers, and a habitat in very wet vegetation; this group includes Faramea multiflora. Many other Faramea species do not correspond to either of these groups. Hooker [Gen. Pl. 2(1): 121-122, 1873] separated several species groups with distinctive morphology but did not name them, but they did separate two Brazilian species of Faramea with well developed calyx limbs into Homaloclados. Later Mueller in his treatment for the Flora brasiliensis separated four sections, and included Homaloclados within the circumscription of Faramea. The genus Faramea currently appears to comprise a monophyletic diagnosable group, but it has not yet been studied in detail with molecular data (Cabral, Jardim, Löfstrand, et al. in prep.).

Faramea has not been studied comprehensively, but a number of floristic treatments are available. This genus has centers of species diversity in southern Central America (Lorence et al., 2012), the Guianas and Guayana Highlands region (Steyermark, 1967; Taylor et al, 2004), the northern and central Andes (Taylor, 1999), and eastern Brazil (Jardim, in prep.).

Faramea has long been classified together with Coussarea, and that relationship was confirmed and those genera were included together in an expanded Tribe Coussareeae by Bremer & Manen (2000; Löfstrand et al., 2019). Faramea was studied with molecular data and a rather broad sampling by Löfstrand et al. (2021), who found the species grouped in two clades that could not be clearly characterized morphologically or geographically. A few incompplete infrageneric classifications had previously been presented based on morphological, usually floristic studies, but their molecular analsyses supported none of these beyond finding  several species from eastern Brazil with unusual enlarged calyx limbs placed on one clade. Löfstrand et al. presented a detailed table of the infrageneric taxa of Faramea that have been recognized, and a more detailed, textual summary of this is presented below on this page as supplemental data for that table. Lófstrand et al. also analyzed and mapped several individual morphological characters of their sampled Faramea species onto their phylogeny, with a focus on characters that were used to diagnose infrageneric taxa; the characters studied were all taken from published literature, and included no new observations as part of that study. Characters they studied included stipule form and arista length, inflorescence position and arrangement, development of bracts, calyx limb morphology, corolla tube size, and fruit and pyrene form, with some of these incompletely documented for the study species because they were not detailed in published description. They found these characters did not diagnose groups found in the molecular analysis. 

Faramea is similar to Coussarea, and these are closely related. In general these differ in their stipules, which are characteristically smooth and acute to rounded in Coussarea, along with their fruits, which are generally spongy or fleshy and yellow to white in Coussarea. Additonally Coussarea has consistently white or yellow flowers; usually (though not exclusively) smooth quadrangular stems; and reproductive stems with a decussate leaf arrangement. Some species of Faramea are also often confused with some genera of Melastomataceae, which have oppposite leaves with similarly well developed submarginal veins, terminal cymose inflorescences, and often also inferior ovaries; but Melastomataceae lack stipules and have free petals . Faramea has also been confused with Palicourea, in particular several Central American species of Palicourea with white nocturnal flowers and thin-walled pyrenes, but Palicourea has bilobed stipules, 5-merous flowers, and bilocular ovaries that produced fruits with 2-5 pyrenes.

Author: C.M. Taylor, in collaboration with Jomar G. Jardim (CEPEC).
The content of this web page was last revised on 20 June 2021.
Taylor web page: http://www.mobot.org/MOBOT/Research/curators/taylor.shtml


Distribution: Moist and humid vegetation from sea level to montane forests, central Mexico and the Antilles to Bolivia and northern Argentina. Faramea has numerous species in most humid continental regions, but only one species throughout the Antilles.
Taxa Included Here:

History of Generic and Infrageneric Classification of Faramea (supplemental discussion to Löfstrand et al., 2021)

Faramea was described from French Guiana with two distinctive species, Faramea corymbosa Aubl. and Faramea sessiliflora Aubl., with the former now designated as the type species. The first infrageneric classification of Faramea was by Candolle (1830), who recognized 3 sections and 17 species based on inflorescence arrangement and bract form. His Faramea sect. Faramea [as "sect. Eufaramea"] was characterized by terminal umbelliform inflorescences borne on 1-3 peduncles and bearing well developed, involucral, caducuous bracts. This section included three distinctive species from the Guianas and one species from Cuba, Faramea sertulifera DC., that was not well known to Candolle; this last species actually lacks the characteristics of the section and is now included within the circumscription of Faramea occidentalis (L.) A. Rich. Candolle's Faramea sect. Tetramerium DC. was characterized by terminal inflorescences that are corymbiform, three-parted, and ebractate, with Faramea occidentalis designated as the type. This section comprised most of the species in the genus and wide range of morphological forms; many of the characters of its constituent species did not actually agree with the diagnosis of the section, however (Candolle, 1830). The third group, Faramea sect. Farameoides DC., was delimited based on its terminal thyrsiform inflorescences and included two species known at that time only from French Guiana; the form of their bracts was not detailed, and these specieds are now both included in Coussarea (Candolle, 1830; Müller, 1881).

Subsequently Hooker (1873) separated some of today's Faramea species into a second genus, Homaloclados Hook.f. His Homaclados included two unusual species from Brazil described in Faramea by Bentham and, provisionally, a third species included in Faramea by Candolle (1830). Hooker did not specifically diagnose his genera but Homaloclados was apparently separated based on its enlarged, petaloid calyx limb with four to eight lobes. Hooker treated his remaining 40 species of Faramea in four infrageneric groups (called "series" in that work, but today sometimes regarded as sections) that were not formally named. Hooker's overview of Faramea here followed but also expanded on Bentham's (1850) treatment of Brazilian Rubiaceae. Hooker's Faramea ser. d was characterized only by its enlarged involucrate bracts; three representative species were cited for this group, and two of them (now considered represent the same species) agree with his diagnosis but the other one actually has a calyx form that matches some species of his Faramea ser. a. Hooker's other three series were separated by combinations of flowers without bracts and various details of stipule and calyx limb form. This first character apparently refers only to a lack of bracteoles subtending an individual flower, because many of the species in these groups do have developed bracts borne elsewhere in their inflorescences. Hooker's Faramea ser. a combined both Sections Faramea and Tetramerium of Candolle (1830), and included most of the species of the genus. He cited five species for his Faramea ser. b, all of them described subsequently to Candolle's (1830) study and most of them now included in a single species, Faramea multiflora A. Rich. (Steyermark, 1967; Taylor & Jardim, 2020). Hooker's Faramea ser. c included two odd species from southeastern Brazil, one of which is now included in Coussarea.

Contemporaneously, Müller (1875, 1881) studied Faramea in Brazil in some detail. He described 67 new species, synonymized Homaloclados with Faramea, presented a formal infrageneric classification with four sections, noted that the inflorescences are variously terminal or axillary, and excluded Faramea Sect. Farameoides with those species transferred to Coussarea. Many of his species are now synonymized (Jardim & Gomes, 2015), but his infrageneric classification was applied by several later authors to the entire genus even though it was based solely on Brazilian species. Müller's Faramea Sect. Homaloclados (Hook.f.) Müll. Arg. was separated by the expanded calyx noted by Hooker for this group, and now comprised 13 distinctive species restricted to southeastern Brazil. Müller recognized two of Candolle's (1830) sections: Faramea sect. Faramea with four species in the Amazon basin, which generally corresponded to Hooker's Faramea ser. d, and Faramea ect. Tetramerium with 64 species distributed throughout the country, which generally corresponded to Hooker's Faramea ser. a. and Faramea ser. c. Müller, however, treated these sections somewhat differently than Candolle and Hooker. He diagnosed both sections by their seeds with a broad rounded excavation on the base (not a pore or opening in the seed coat) and sheathing stipules with well-developed aristae, and then broadened the characterization of each group by additonal features: petaloid, foliaceous bracts and flattened peduncles in Faramea sect. Faramea, and inflorescences branched to two or three orders with small bracts in Faramea sect. Tetramerium. Müller also described a fourth infrageneric group, Faramea ect. Hypochasma Müll. Arg., characterized by seeds with a well-developed, transverse hylar fissure (I.e., pre-formed germination slit), small bracts, and sheathing stipules with short aristas; this group generally corresponds to Hooker's Faramea ser. b.

A decade later, in Schumann's (1891) detailed review world Rubiaceae his tratment of Faramea closely followed that of Müller, and as for Hooker (1870), his view of the genus was heavily biased toward the Brazilian species. Schumann (1891) distinguished and circumscribed Coussarea and Faramea following Müller (1875,1881), citing his diagnosis word for word. Schumann estimated that Faramea included about 100 species, with 87 of them in Brazil. He recognized all four of Müller's sections, and generally characterized them similarly with the addition of some notes about character variation and their distribution outside Brazil.

The next detailed treatment of Faramea reviewed species of northwestern South America was Bremekamp's (1934). In his Rubiaceae treatment for Surinam. Here he expanded the circumscription of Faramea to include Evea Aubl. Its single original species, Evea guianensis (Aubl.) Bremek., is unusual in Faramea in its consistently axillary inflorescences with small cymes that are enclosed in enlarged, involucral bracts. This species has all the other characters of Faramea and clearly belongs here, but its inclusion complicates the infrageneric classification of the genus because its pattern of characters does not agree with any of Müller's (1875, 1881) and Schumann's (1891) sections. Bremekamp did not classify his species in any infrageneric system.

More recently, Steyermark (1967) presented a relatively broad floristic study of Faramea, covering all northwestern and much of central South America. He accepted Bremekamp's inclusion of Evea in Faramea, recognized 30 species with 9 of them new, and synonymized a rather large number (16) of previously recognized species. Steyermark did not group his species in any infrageneric classification, although he did present them in an order that largely corresponds to those of Faramea sect. Faramea followed by those of former genus Evea, then Faramea sect. Tetramerium, and then Faramea sect. Hypochasma. In general throughout his regional study, Steyermark only recognized infrageneric classifications when he studied the significant majority of the species of that genus, which he did not do for Faramea.

Contemporaneously with Steyermark (1967), Dwyer and Hayden (1967a, 1967b) took the opposite approach in their study of general neotropical Rubiaceae floristics. Here, they detailed the Rubiaceae for Panama and also studied selected specimens from western South America, named three new Faramea species (some of which have since been transferred to Coussarea, e.g., Dwyer, 1980), and discovered some apparently new characters for Faramea. They compared their Faramea species with Schumann's infrageneric classification, and when their new species did not agree completely with his groups they separated each into its own monotypic section based on the deviating character. Their Faramea sect. Grandistipulata included only Faramea jefensis Dwyer & M.V. Hayden and was separated based on its unusually long tubular stipules. Several other Faramea species have similar stipules but were not noted by them (e.g., Faramea calophylla Standl.). Their Faramea sect. Integrisepta Dwyer & M.V. Hayden included Faramea persistisepta Dwyer & M.V. Hayden, which was notable in the genus in its ovary and fruit with a well-developed septum; this is indeed an anomalous character for Faramea, and Faramea persistisepta has since been reidentified as Rudgea foveolata (Ruiz & Pav.) Zahlbr. (Taylor et al., 2015). Their third group, Faramea sect. Uniflora Dwyer & M.V. Hayden, included Faramea uniflora Dwyer & M.V. Hayden and was separated based on its solitary flowers. Some other Faramea species also have solitary flowers, and presumably those were intended to be separated from Faramea sect. Uniflora by some other characters but Dwyer & Hayden included no other features in their diagnosis of this section. Their Faramea uniflora agrees in all its other characters with Faramea sect. Hypochasma, and could alternatively be considered an endpoint of inflorescence variation form an this group: from inflorescences with thirty or more flowers in Faramea amazonica Müll. Arg., to often about a dozen flowers in Faramea multiflora A. Rich., to 2-7 flowers in Faramea quinqueflora Poepp. & Endl., to consistently solitary flowers in Faramea uniflora.

Recently, Jardim (2008) presented the study of Faramea with modern phylogenetic methods. He demonstrated the monophyly of the genus with an analysis of morphological and molecular data for 25 accessions (representing 23 species; primarily collected in Brazil). This ground-breaking study is, unfortunately, still not formally published but provides a useful framework. Jardim's phylogenetic analysis placed its three sampled species of Faramea sect. Hypochasma in a well-supported monophyletic group and grouped the remaining species in a weakly supported clade. Within this larger clade the three sampled species of Faramea sect. Homaloclados were grouped in a weakly supported clade but no other groupings were found. Jardim also reviewed the infrageneric taxonomy of Faramea as a whole and Faramea sect. Hypochasma in detail, based on morphological characters. He included Faramea sect. Grandistipulata and Faramea sect. Uniflora within Faramea sect. Hypochasma along with a number of previously unclassified species, and excluded some others from this section (e.g., Faramea pachyantha Müll. Arg.). Jardim’s results supported the synapomorphic status of the seed morphology and fused stipules for Faramea sect. Hypochasma (Müller 1875, 1881), and found other traits considered diagnostic of sections, such as inflorescence structure, to be homoplasious.


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Shrubs, subshrubs, and small trees, unarmed, terrestrial, with raphides in the tissues. Leaves opposite, sessile or petiolate, entire, with higher-order venation not lineolate, occasionally with crypt-type domatia; stipules interpetiolar or fused into a tube or a calyptrate cap, triangular to rounded, acute to usually aristate, apparently imbricated in bud, persistent or caducous. Inflorescences terminal or sometimes axillary, cymose, fasciculate, or reduced, 1- to multiflowered, often blue or white, sessile to pedunculate, bracteate or bracts reduced. Flowers sessile or pedicellate, bisexual, apparently usually distylous, fragrant, diurnal or perhaps sometimes nocturnal, generally medium-sized; hypanthium ellipsoid to turbinate; calyx limb truncate to lobed, lobes 4, without calycophylls; corolla salverform or tubular, blue, purple, or white turning yellow with age, internally glabrous, lobes 4, triangular, valvate in bud, often fleshy and triangular in cross-section, without appendages; stamens 4, inserted at various levels of corolla tube, anthers narrowly oblong, dorsifixed below middle, opening by longitudinal slits, without appendages, included; ovary 1-locular or incompletely 2-locular, with ovules 1--2, basal; stigmas 2, included or exserted. Fruit drupaceous, subglobose to ellipsoid or oblate and laterally or longitudinally flattened, fleshy to leathery or juicy, blue to purple-black, with calyx limb persistent; pyrenes 1(2), 1-locular, subglobose to ellipsoid or oblate, chartaceous to papery, with raphal pore and sometimes with preformed germination slits; seeds 1 per pyrene, ellipsod to subglobose.


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