Research Article
Research Article
A new species of Raputia (Rutaceae) from the Selva Central of Peru
expand article infoRobin Fernandez-Hilario, Robert Arteaga§
‡ Universidad Nacional Agraria La Molina, Lima, Peru
§ Universidad Nacional Mayor de San Marcos, Lima, Peru
Open Access


Raputia codo-pozuzoensis Rob. Fernandez & Arteaga, sp. nov. (Galipeinae, Rutaceae), a new species endemic to the Huanuco region, in the Selva Central of Peru, is described and illustrated here. The new species differs from other species of Raputia by the combination of 5–7-foliolate leaves (a new characteristic for the genus) and longer petioles. A key to the four Peruvian species of Raputia is presented.


Raputia codo-pozuzoensis Rob. Fernandez & Arteaga, sp. nov. (Galipeinae, Rutaceae), una nueva especie endémica del departamento de Huánuco, en la Selva Central del Perú, es descrita e ilustrada aquí. La nueva especie difiere de las otras especies de Raputia por presentar hojas 5–7-folioladas (una nueva característica para el género) y peciolos largos. Una clave para las cuatro especies peruanas de Raputia es presentada.


Raputia, Galipeinae, premontane forests, Neotropic

Palabras clave

Raputia , Galipeinae , Bosques premontanos, Neotrópico


Rutaceae Juss., in the order Sapindales (APG III, 2009; APG IV, 2016), is a family of mostly tropical and sub-tropical trees, shrubs, and aromatic herbs comprising approximately 2100 species in 154 genera (Kubitzki et al. 2011). In recent years, numerous phylogenetic studies have revealed that the seven sub-families initially proposed by Engler (1931) were paraphyletic (Chase et al. 1999, Scott et al. 2000, Groppo et al. 2008, Groppo et al. 2012, Morton and Telmer 2014), a revelation that has prompted recircumscription of intra-familial groups and new understanding of relationships among them. For example, at the subfamily level, Groppo et al. (2012) reduced the number of subfamilies to two, and Morton and Telmer (2014) to four; and at a more specific level, Bruniera et al. (2015), in the first study of relationships within subtribe Galipeinae (tribe Galipeeae, subfamily Rutoideae), where Raputia belongs, transferred all the species of Almeidea A. St.-Hil. to Conchocarpus J.C. Mikan and determined that the Galipeinae is apparently a monophyletic group.

Raputia and its type species Raputia aromatica were established by Aublet in 1775 based on collections from forest near the Orapu River in French Guiana. In the first comprehensive classifications of the Rutaceae, Engler (1874, 1931) recognized nine species in the genus, forming a group with heterogeneous characteristics.

Emmerich (1978) split this group and placed most of the species in three more homogenous genera (Neoraputia Emmerich, Sigmatanthus Huber ex Emmerich and Raputiarana Emmerich). Subsequently, Kallunki (1990) emended the description of Raputia and making three new combinations. Finally, the last published revision of the genus was made by Kallunki (1994), where she recognized 10 species. Currently, Raputia is found in the subtribe Galipeinae Kallunki [Angostura Alliance sensu Kubitzki et al. (2011)] along with 25 other genera, all restricted to the Neotropics, and characterized by tendencies toward zygomorphic flowers, a more or less tubular corolla, union of the filaments to a corolla tube, reduction in number of fertile stamens from five to two with the transformation of stamens into staminodes, modifications of anthers, loss of endosperm, a curved embryo and conduplicate and plicate cotyledons (Bruniera et al. 2015, Kallunki and Pirani 1997, Kubitzki et al. 2011). In addition, in most taxa with only two fertile stamens, the anthers are variously modified by basal or apical appendages or sterile basal portions above the point of attachment to the filament and, in some, the anthers or appendages are united (Kubitzki et al. 2011).

Brako and Zarucchi (1993), in the “Catalogue of the Flowering Plants and Gymnosperms of Peru”, listed three species known from that country: Raputia ulei (K. Krause) Kallunki, Raputia heptaphylla Pittier and Raputia magnifica Engler. In the revision of the genus, Kallunki (1994) cited no specimens of Raputia ulei from Peru. Instead, she identified the collection (Vásquez et al. 8909) that was the basis for the report of that species from Peru (Brako and Zarucchi, 1993) as Raputia simulans Kallunki. Furthermore, she excluded Raputia heptaphylla and Raputia magnifica from the genus because they showed discordant characteristics and subsequently recognized them as Raputariana heptaphylla (Pittier) Kallunki (Jiménez 2014), and Neoraputia magnifica(Engl.) Emmerich ex Kallunki (Kallunki 2009), respectively. Additionally, Kallunki (1994) described Raputia megalantha Kallunki and transferred Achuaria hirsuta Gereau to Raputia hirsuta (Gereau) Kallunki, which with Raputia simulans Kallunki, are known from Peru (Vasquez and Rodriguez 2002; Ulloa Ulloa et al. 2004).

Raputia comprises a total of 11 Neotropical species, occurring from Venezuela and French Guiana to Amazonian Colombia, Peru and Brazil, principally from lowland areas, with shrubby or tree-like habit, opposite 1–3-foliolate leaves, circinate cauline monochasium, and pentamerous flowers with bilabiate corolla (Kallunki 1994; Kubitzki et al. 2011; Pirani 2005).

During fieldtrips to the premontane forests of Huanuco (Peru) in 2015, we collected an undescribed species of Raputia with 5–7-foliolate leaves, a new characteristic for the genus.

Taxonomic treatment

Raputia codo-pozuzoensis Rob. Fernandez & Arteaga, sp. nov.

Figure 1, 2


Raputiacodo-pozuzoensis differs from others species in this genus by its 5–7-foliolate leaves and longer petioles (8.5–12.5 cm long).


PERU. Huánuco: Prov. Puerto Inca, Dist. Codo de Pozuzo, alrededores de toma de agua cerca al Río Pozuzo, 565 m, 9°40'57.76"S, 75°30'31.35"W, 01 Feb 2015 (fl.), R. Fernandez, R. Arteaga & F. Meza 830 (holotype MOL - 2 sheets).


Monopodial shrub up to 2 m tall; stem cylindrical, 1–1.5 cm in diameter, lenticellate and finely ribbed, dark brown; the terminal buds, young twigs and petioles, and inflorescences pubescent, the hairs short and antrorse. Terminal twigs circular in transverse section, 4–7 mm in diameter, beige-colored when dry, lenticellate; terminal buds ferruginous, stipules absent. Leaves palmately compound, 5–7-foliolate, opposite or verticillate; petiole cylindrical, 8.5–12.5 cm long, 2–3 mm wide; petiolule absent; leaflet blades elliptical, acuminate at apex, decurrent at base, entire at margin, discolorous, sub-chartaceous, the venation brochidodromus, the surface pellucid-punctate, the upper and lower surface glabrous, midrib pubescent beneath, the hairs short and antrorse; central leaflet (21-) 25–36 cm long, 3.5–6 cm wide, the secondary veins (18-) 21–29; lateral leaflet progressively smaller, the basal ones (7.5-) 11–18 cm long, 1.7–3.5 cm wide, the secondary veins 10–16. Inflorescence cauline, of 1–3 monochasia, with 6–14 flowers, 1.8–3 cm long including a peduncle 2–5 mm long. Flowers bisexual, zygomorphic, pentamerous; pedicel 1.5–2 mm long; flower buds slightly curved. Calyx 4–4.5 mm long, 3.5 mm wide at base of lobes, glabrous or pubescent; sepals fused at the base, 5-lobed, the lobes quincuncial, ovate, acute to obtuse at apex, 2 mm long, ciliate, pellucid-punctate. Corolla tubular, unequally 5-lobed, 12–17.5 mm long, bilabiate at anthesis, glabrous in the external surface, sparsely pubescent in the inner base of the tube, woolly in the inner part of the throat, the trichomes ca. 1.2 mm long; the tube white to yellowish, 2–6 mm long to the sinuses of the innermost lobe (inferior lip), 6–9 mm long to the sinuses of the other 4 lobes (superior lip), recurved superior lip; the lobes green, imbricate, oblong, rounded at apex, the inner lobe 10–12 mm long, 3.5–5 mm wide, the other 4 lobes 5–6 mm long, 3.5–4.5 mm wide, pellucid-punctate. Androecium of 2 fertile stamens and 3 staminodia, white-colored; filaments of fertile stamens flanking the inner lobe, adherent from the base to the throat of the corolla tube, the free portion above the throat ca. 2 mm long; staminodia adherent from the base to the throat of the corolla tube, the free portion linear above the throat 9–11 mm long, alternate with the other four corolla lobes; filaments of fertile stamens and staminodia glabrous at the base and apex, only bearded at the throat of the corolla, filaments and back of anthers pellucid-punctate; anthers lanceolate, laterally coherent, basifixed, ca. 5.5 mm long, 1.5 mm wide, glabrous, the appendages flattened, ca. 1.5 mm long, 1 mm wide, glabrous. Gynoecium, ovary of 5 carpels united at the base and by single style, 1.5 mm in diameter, 1 mm high, furrowed, orange-colored; the style 10–11 mm long, slightly curved, glabrous, pellucid-punctate; the stigma 1 mm in diameter, slightly 5-lobed; disc cupular enveloping the ovary, 2.5 mm in diameter, 1.5 mm high, margin 5-lobed, cream-colored, glabrous. Fruit not seen.

Figure 1. 

Raputia codo-pozuzoensis. A Terminal twig B Midrib beneath and lower leaf surface C Inflorescences D Flower E Corolla opened showing two fertile stamens and staminodes F Longitudinal section of calyx, disc cupular, ovary and style. From R. Fernandez et al. 830 (MOL). Drawing by Sara Terreros.

Figure 2. 

Raputia codo-pozuzoensis. A Habit B Inflorescence (R. Fernandez & R. Arteaga 1079; MOL) C, D Flowers (R. Fernandez et al. 830; MOL). Photos by Robin Fernandez.

Distribution and habitat

Raputia codo-pozuzoensis is endemic to humid premontane forest in central Peru, between 565–589 m.a.s.l., growing in zones with shallow to steep slopes in a loamy-silty soil. The only known population of this species occurs in the understory of a forest of tree species, such as: Chrysophyllum sanguinolentum (Pierre) Baehni, Helicostylis scabra (J.F. Macbr.) C.C. Berg, Hevea guianensis Aubl., Iryanthera hostmannii (Benth.) Warb., Mabea speciosa Müll. Arg., Macrolobium gracile Spruce ex Benth., Theobroma subincanum Mart. and Virola pavonis (A. DC.) A.C. Sm.

Figure 3. 

Distribution of Raputia species in Peru.


The specific epithet refers to the Codo de Pozuzo district, the only place where the specimens were found and collected.


Flowering take place from December to February.

Conservation state

We collected individuals of Raputia codo-pozuzoensis in areas of slightly disturbed forest, and we observed individuals sprouting after being cut for the establishment of “trochas” (pathways). We counted 20 individuals in an area of 0.5 ha. Thus, we assume that human activities are not affecting seriously the wild populations of this species. Nonetheless, in our inventories at other sites in Puerto Inca Province and surrounding areas, we and our collaborators have not observed other populations of this new species. Additionally, the extent of occurrence estimated of this species has been decreasing over the last years by deforestation and only remaining less than 100 km2 of the original forest cover. Therefore, under the guidelines of UICN (2012), we assign this species to the category Critically Endangered [CR (B1a+bi)].

Additional specimens examined

PERU. Huánuco: Prov. Puerto Inca, Dist. Codo de Pozuzo, alrededores de toma de agua cerca al Río Pozuzo, 589 m, 9°40'56.72"S, 75°30'30.85"W, 28 Dec 2015 (fl.), R. Fernandez & R. Arteaga 1079 (MOL), R. Fernandez & R. Arteaga 1080 (USM), R. Fernandez & R. Arteaga 1081 (HOXA).


According to the morphologic analyses of Kubitzki et al. (2011), Raputia belongs to a natural group along with the following genera: Apocaulon R.S. Cowan, Decagonocarpus Engl., Ertela Adans., Lubaria Pittier, Ravenia Vell. and Raveniopsis Gleason, characterized by their opposite leaves (alternate in Apocaulon), overlapping sepals, connate petals, basally appendaged (exc. in Ertela) and often laterally coherent anthers, reticulate pollen grains, apocarpous gynoecia, and conduplicate, bilobed cotyledons. Among this group, the two herbaceous genera, Apocaulon and Ertela, stand out and differ noticeably from the others. The former, by its alternate leaves and anthers coherent by their adaxial surfaces, and the later by its strongly unequal sepals, with the two outer much larger and concealing the corolla, and the anthers lacking basal appendages. Previously, Kallunki (1994) recognized that Raputia was related with the following genera: Decagonocarpus, Lubaria, Ravenia and Raveniopsis, forming a group characterized by opposite leaves, a quincuncial calyx (of which the margins are conspicuously overlapping at anthesis), and conduplicate, bilobed cotyledons. Kallunki (1994) differentiated Raputia from these other four genera by the presence of cauline inflorescences and the leathery testa (vs. terminal inflorescences and crustaceous testa). Even though we did not register neither the seeds nor the fruits of Raputia codo-pozuzoensis, the combination of characteristics such as cauline inflorescences, the petals connate, forming a bilabiate corolla with a short tube, and anthers laterally connate, with basal appendages, allow us to locate this new species in the Raputia genus. In Table 1 we display the different characteristics of genera related to Raputia, according to Kubitzki et al. (2011).

Table 1.

Comparison of Raputia with the morphologically most similar genera. Based on Albuquerque (1976), Kallunki (1994, 2005), Pirani (2005) and Kubitzki et al. (2011).

Character Apocaulon Decagonocarpus Ertela Lubaria Raputia Ravenia Raveniopsis
Habit Herbs Shrubs or trees Herbs, sometimes suffruticose Trees Shrubs or trees Shrubs or trees Shrubs or trees
Leaf characters Leaves alternate, congested, often appearing basal, 3-foliolate Leaves opposite, simple Leaves usually opposite on lower part of stem, sometimes alternate or subopposite on upper part, 3-foliolate Leaves opposite, simple Leaves opposite, 1–7-foliolate Leaves opposite or appearing alternate due to reduction of one of two opposite leaves, simple or 3-foliolate Leaves opposite (in Raveniopsis steyermarkii R.S. Cowan some alternate), 1–3-foliolate
Inflorescence position Axillary Terminal Terminal but sometimes appearing axillary Terminal Cauline (rarely axillary) Terminal Axillary or terminal
Inflorescence type Dichasium Monochasium Dichasium Dichasium Monochasium Dichasium, monochasium, or 1 or 2 flowers Dichasium, monochasium, a congested capitulate thyrse, or a solitary flower
Calyx aestivation Unknown Quincuncial Quincuncial Quincuncial Quincuncial Quincuncial Quincuncial
Calyx features Sepals 5, strongly unequal, shortly coherent Cupular, ± equally 5-lobed Sepals 5, free, strongly unequal, the 2 outer much larger and concealing the corolla Sepals 5, free, the 2 outer larger Sepals 5, connate at very base, ± unequal Sepals 5, the 2 outer larger than inner Sepals 5, free or very shortly connate, usually unequal
Corolla aestivation (4)5 imbricate lobes Induplicate-valvate Imbricate Imbricate Imbricate Imbricate Imbricate
Corolla features Petals 5, connate, the tube curved, the 2 lobes opposite the lobe flanked by the 2 stamens joined for a slightly longer distance and forming a bilobed lip Petals 5, connate into a long, slightly curved tube with recurved to spreading lobes, the tube longer than the lobes Petals 5, connate, forming a bilabiate corolla with a short tube, one lip formed by the innermost petal, the other lip 4-lobed Petals 5, the adaxial, innermost one free, the others connate into a 4-lobed lip Petals 5, connate, forming a bilabiate corolla with a short tube, one lip formed by the innermost petal, the other lip forming a 4-lobed, recurved lip Petals 5, connate to the middle or more, forming a bilabiate corolla Petals 5, connate, forming a slightly zygomorphic to markedly bilabiate corolla, the corolla tube cylindric, slightly curved, longer than or equal to the lobes
Anthers Coherent by their adaxial surfaces, with basal appendages Laterally coherent, with basal appendages Laterally coherent in lower half, lacking basal appendages Laterally coherent, with basal appendages Laterally connate, with basal appendages Laterally coherent or not, with or without basal appendages Sometimes laterally coherent, with basal appendages
Testa Crustaceous Crustaceous Crustaceous Crustaceous Leathery Crustaceous Crustaceous
Cotyledons Conduplicate, emarginate at apex Conduplicate, bilobed at apex Conduplicate, bilobed at apex Conduplicate, bilobed at apex Conduplicate, thick, stiff, bilobed at apex Conduplicate, fleshy, bilobed at apex Conduplicate, bilobed at apex (or rarely incumbent and rounded at apex)
Number of species 1 2 2 1 12 11 19

Raputia codo-pozuzoensis is easily distiguished from all other species of the genus by its 5–7-foliolate palmately compound leaves. The other three species found in Peru show unifoliolate (Raputia simulans) or three-foliolate leaves (R. hirsuta and R. megalantha). Raputia codo-pozuzoensis differs further from R. simulans by its much shorter inflorescences 1.8–3 cm long (vs. 19.5–26.5 cm) and from Raputia hirsuta by its short and antrorse hairs (vs. hirsute) on stems, leaves, and inflorescences.

Like Raputia megalantha and Raputia maroana (R.S. Cowan) Kallunki, Raputia codo-pozuzoensis possesses inflorescences shorter than 6 cm and terminal leaflets longer than 20 cm. Raputia codo-pozuzoensis differs, however, from R. megalantha by its petioles 8.5–12.5 cm long (vs. 0.8–3.3 cm) and its corollas 12–17.5 mm long (vs. 30 mm). Although Raputia codo-pozuzoensis shares with R. maroana petioles and corollas of similar lengths, it differs from the latter by its 5–7-foliolate (vs. 3-foliolate) leaves and filaments ca. 2 mm (vs. 11–12 mm) long. In addition, Raputia codo-pozuzoensis is restricted to premontane forest in southwestern Amazonia (Huanuco, Peru), whereas R. megalantha and R. maroana are distributed in lowland forests in northwestern Amazonia (Brazil, Peru, and Venezuela; Kallunki, 1994).

Key to the species of Raputia in Peru

1 5–7-foliolate leaves, petioles longer than 8 cm R. codo-pozuzoensis
1–3-foliolate leaves, petioles shorter than 4 cm 2
2 Leaves 1-foliolate; inflorescences longer than 16 cm; flowers more than 20 R. simulans
Leaves 3-foliolate; inflorescences shorter than 7 cm; flowers fewer than 15 3
3 Central leaflet 8–30 cm long; inflorescences and petioles hirsute R. hirsuta
Central leaflet 50–71 cm long; inflorescences and petioles strigulose R. megalantha

Additional Peruvian specimens of other species

Raputia hirsuta. PERU. Loreto. Prov. Coronel Portillo, Padre Abad, granja del sr. Barrera, NE de la chacra de César Vela (Aguaytia), 17 Oct 1972, V. Schunke 5396 (F, MO, NY); Prov. Loreto, Río Tigre, San Jacinto, Campamento de Occidental Petroleum, 175–205 m, 02°28'S, 75°47'W, 08 Jun 1993, H. Beltrán & R. Foster 435 (F, USM); Prov. Loreto, Dist. Loreto, Pampa Hermosa and vicinity, 3°15'S, 75°50'W, 03–20 Dec 1985, W. Lewis et al. 10328 (F, MO, USM), 04–09 Jun 1986, W. Lewis et al. 10729 (MO, USM), 04–09 Jun 1986, W. Lewis et al. 10782 (MO, USM); Prov. Loreto, Campamento Petrolero San Jacinto, Rio Tigre, 2°15'S, 75°50'W, 16 Sep 1979, C. Díaz & N. Jaramillo 1454 (MO); Prov. Mariscal Ramón Castilla, Margen izquierda del Río Yavari, entre Colonia Angamos y Lago Preto, 4°30'53"S, 71°54'2.77"W, 10 Apr 2003, H. Beltrán et al. 5743 (AMAZ, USM); Prov. Mariscal Ramón Castilla, Alto Río Yaguas, tributario del Río Putumayo, aprox. 80 km NE de Pebas, 140 m, 02°51'53.5"S, 71°24'54.1"W, 07 Aug 2003, M. Ríos et al. 537 (F); Prov. Maynas, Río Blanco, a 3 horas (Jonhson 40 Hp) desde Tamshiyacu, 130 m, 15 Mar 1978, C. Díaz et al. 145 (MO); Prov. Requena, Sapuena, Jenaro Herrera, 170 m, 4°50'S, 73°45'W, 12 Nov 1987, R. Vásquez et al. 9983 (MO, USM).

Raputia megalantha. PERU. Loreto. Prov. Maynas, Mishuyacu, near Iquitos, 100 m, Oct-Nov 1929, G. Klug 544 (F, NY, US); Prov. Maynas, Estación Biológica Rio Blanco, 04°20'S, 72°45'W, 16 Sep 1985, R. Vásquez et al. 6743 (MO, NY); Prov. Maynas, Dist. Allpahuayo, Estación Experimental del IIAP, 04°00'15"S, 73°25'48"W, 30 May 1990, R. Vásquez et al. 13786 (MO, USM); Prov. Maynas, Dist. Iquitos, carretera del caserío de Puerto Almendras, 26 Jun 1984, M. Rimachi 7529 (US); Prov. Maynas, Dist. Iquitos, San Juan, km 31.5 carrera Iquitos-Nauta, 160 m, 3°59'34.7"S, 73°26'42.5"W, 08 Sep 2002, M. Flores et al. 1690 (AMAZ, MO, USM); Prov. Maynas, Dist. Iquitos, Allapahuayo, Estación del IIAP, 150–180 m, 04°10'S, 73°30'W, 18 Jun 1991, R. Vásquez 16810 (AMAZ, MO, NY); Prov. Maynas, Dist. Iquitos, Allpahuayo, Estación Experimental del IIAP, 03°58'16"S, 73°25'08"W, 11 Oct 1990, R. Vásquez & N. Jaramillo 14495 (MO); Prov. Maynas, Dist. San Juan, Reserva Nacional Allpahuayo-Mishana, 128 m, 03°58'02"S, 73°25'08"W, 19 Nov 2008, R. Vásquez et al. 35060 (HOXA). San Martín. Prov. Lamas, Caserio Bonilla, trail to E of road, Km 75 of Tarapoto-Yurimaguas road, 250 m, 6°16'S, 76°17'W, 20 Apr 1986, S. Knapp & J. Mallet 7138 (USM, MO, NY); Prov. Lamas, Santa Rosa de Davidcillo, trail to E of road to Tioyacu, 200 m, 6°16'S, 76°17'W, 22–23 Apr 1986, S. Knapp & J. Mallet 7162 (USM, MO, NY); Prov. Lamas, Santa Rosa de Davidcillo, 220 m, 6°15'S, 76°17'W, 21 Aug 1986, S. Knapp 8109 (MO, USM).

Raputia simulans. PERU. Loreto. Prov. Mariscal Ramón Castilla, cabeceras del Río Apayacu, Noroeste de Pebas, 150 m, 03°07'00"S, 72°42'43"W, 17 Aug 2003, M. Ríos et al. 680 (F, NY); Prov. Maynas, Dist. Iquitos, Quebrada Aucaya, trocha de la cooperativa, 11 Aug 1973, S. McDaniel & M. Rimachi 17701 (MO, NY); Prov. Maynas, Dist. Iquitos, Nina rumi-Rio Nanay, 122 m, 03°48'S, 73°25'W, 05 Mar 1987, R. Vásquez et al. 8909 (MO, NY); Prov. Maynas, Dist. Iquitos, Estación Experimental del IIAP, 04 Nov 1990, R. Vásquez & N. Jaramillo 14560 (MO, NY), 150–180 m, 04°10'S, 73°30'W, Nov 1990, R. Vásquez & N. Jaramillo 14626 (MO), 24 May 1991, R. Vásquez & N. Jaramillo 16426 (MO); Prov. Maynas, Dist. Iquitos, Allpahuayo, 150 m, 04°10'S, 73°30'W, 19 Mar 1992, R. Vásquez 17727 (MO); Prov. Maynas, Dist. Iquitos, carretera Iquitos-Nauta, km 28, trocha del Fundo Pichiri, 150 m, 23 Jul 1997, M. Rimachi 12005 (USM); Prov. Maynas, Dist. Putumayo, NE de Iquitos y Pebas, en la esquina del trapezoide de Colombia, 80 m, 02°43'15.9"S, 70°34'30.6"W, 28 Oct 2010, I. Huamantupa et al. 14752 (F).


This research has been developed as part of the biological monitoring program of the 2D seismic project of Petrolifera Petroleum of Peru, company of the Gran Tierra group. We express our sincere gratitude to Pacific PIR for the logistical facilities in the field and Jacquelyn A. Kallunki for her support and comments during the drafting of this manuscript. The curators from the AMAZ, HOXA and USM herbaria are gratefully acknowledged for their logistical support during the review of botanical material. We also thank Sara Terreros for the excellent illustration and Andrea Bravo for the distribution map, as well as Kenneth J. Wurdack and Carla P. Bruniera for their valuable feedback on the manuscript.


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