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Boletín de la Sociedad Argentina de Botánica

versión On-line ISSN 1851-2372

Bol. Soc. Argent. Bot. vol.45 no.3-4 Córdoba jul./dic. 2010



Inkaliabum, a new Andean genus of Liabeae (Asteraceae) from Peru

Diego G. Gutiérrez1

1 Assistant researcher, CONICET. División Plantas Vasculares, Museo de La Plata (UNLP), Paseo del Bosque s.n., B1900FWA La Plata, Buenos Aires, Argentina. Email:


Summary: Inkaliabum, a new monotypic genus endemic to valleys and slopes of the Andes of Cuzco (Peru), is described. The species of genus Inkaliabum is a slender scandent subshrub with pinnately veined leaves, lacking latex, with few capitula in terminal and/or axillary capitulescences, widely campanulate involucres, many phyllaries, many florets per capitulum, and disc florets with corollas slightly differentiated into tube and throat, long style branches, and a short pubescent part of the style shaft. The new combination Inkaliabum diehlii, based on Liabum diehllii is established, and the species is illustrated. In addition, a key to the genera of Liabeae from Peru plus Oligactis is provided.

Key words: Asteraceae; Endemism; Inkaliabum; Inkaliabum diehlii; Liabeae; Liabum; Peru.

Resumen: Inkaliabum, un nuevo género andino de Liabeae (Asteraceae) de Perú. Se describe Inkaliabum, un nuevo género monotípico endémico de los valles y pendientes de los Andes de Cuzco (Perú). La especie del género Inkaliabum es un subarbusto grácil escandente con hojas pinnatinervadas, ausencia de latex, pocos capítulos dispuestos en capitulescencias terminals y/o axilares, involucros anchamente acampanados, filarias numerosas, flores numerosas por capítulo y flores del disco con corolas levemente diferenciadas en tubo y limbo, ramas de estilo largas y la porción pubescente del estilo por debajo del punto de bifurcación corta. Se establece la nueva combinación, Inkaliabum diehlii, basada en Liabum diehllii, especie que es ilustrada. Además, se provee una clave de los géneros del Liabeae de Perú más Oligactis.

Palabras clave: Asteraceae; Endemismo; Inkaliabum; Inkaliabum diehlii; Liabeae; Liabum; Peru.



The small Neotropical tribe Liabeae includes more than 150 species distributed in 18 genera (Dillon et al., 2009). Despite being a tribe well defined, news in several taxonomic levels were presented recently based on morphological and molecular data.Liabeae was divided in three subtribes: Munnoziinae, Paranepheliinae, and Liabinae, based on morphological and palynological characters (Robinson, 1983a; Robinson & Marticorena, 1986). Results from morphological cladistic studies by Bremer (1994), Funk (1985) and Funk et al. (1996) demonstrated the monophyly of the first two subtribes but not the Liabinae. Later, however, the monophyly of Munnoziinae was questioned in a further molecular study (Kim et al., 2003). Recently, the results of molecular investigations on Liabeae, have recovered consistent placements for several taxa (Gutiérrez et al., 2007; Dillon et al., 2009). The consensus tree of last phylogeny of Liabeae shows four main clades named as Liabinae, Sinclariinae, Paranepheliinae, and Munoziinae (Dillon et al., 2009).
This better knowledge based on morphological and molecular information showed some needed changes at the genera and specific levels: the monotypic genus Bishopanthus H. Rob. and Dillandia Funk & H. Rob. were described (Robinson, 1983b; Funk & Robinson, 2001), the genus Austroliabum H. Rob. & Brettell was reduced to synonymy under Microliabum Cabrera (Robinson, 1990), the subgenus Andromachiopsis of Oligactis (Kunth) Cass. was elevated to the generic level as Sampera Funk & H. Rob. (Dillon et al., 2009), the monotypic genus Sinclariopsis Rydb. was resurrected (Dillon et al., 2009), and some taxonomical rearrangements have been proposed in Liabum Adans. (Gutiérrez & Katinas, 2006) and Oligactis (Gutiérrez, 2008; Dillon et al., 2009).

Liabum, one of the largest and more widedistributed genera of Liabeae, contains 26 species which grow in a wide variety of habitats throughout southern Mexico, Central America, the West Indies (Cuba, Jamaica, and Hispaniola), and western South America from northern Venezuela to northwestern Argentina (Gutiérrez, 2003; Gutiérrez & Katinas, 2006; Gutiérrez & Katinas, in prep.). The greatest diversity of this genus is found in Ecuador and northern Peru where not less than 15 species occur. Liabum is characterized by a combination of morphological characters: perennial herbs to shrubs (rarely small trees), latex absent, slightly or strongly triplinerved (acrodromous) leaves, leaf abaxial surface albotomentose, capitulescences of usually many capitula, long style branches, disc florets style with its pubescent portion below the bifurcation point equal or shorter than the style branches, yellow pale anthers, eglandular cypselae and subquadrate crystals of cypsela wall.

However, three endemic species from Peru have several morphological features that deviate from Liabum: L. ferreyri H. Rob. and L. sandemanii H. Rob. with densely glandular cypselae and L. diehlii H. Rob. with scandent habit and pinnately veined leaves (Gutiérrez, 2004; Gutiérrez & Katinas, in prep.). L. ferreyri and L. sandemanii are being studied in order to determine their taxonomic ubication (Gutiérrez & Katinas, in prep.).

The species Liabum diehlii was described by Robinson (1980) who included it under Liabum because of its general aspect and type of cypsela trichomes. However, he recognized that the narrowly elliptical leaves with pinnate venation of this species are commonly found in the genus Oligactis s.l. Recently, during the preparation of a revision of Liabum (Gutiérrez, 2004; Gutiérrez & Katinas, in prep.), a detailed morphological study of the specimens of L. diehlii showed that many systematic characters such as the habit, leaf venation, type and position of capitulescence, size of involucre, number of phyllaries, number of florets, and type of disc florets do not correspond to the genus Liabum. According to the findings above L. diehlii is proposed to be excluded from Liabum.


This study as a part of the doctoral thesis is based on more than 1500 specimens or digital photographs from the following herbaria (Holmgren et al., 1990): B, BAB, BM, CTES, F, FI, G, GH, IJ, K, LIL, LINN, LP, MO, MY, NY, P, PMA, S, SI, US, USM, VEN, W. The data were supplemented by field observations and information from the literature. For microscopic examination, vegetative and reproductive parts were rehydrated, treated with a clearing process, stained with 2% safranin, and mounted on microscope slides. Drawings were made using a stereomicroscope Wild M5 and Olympus CH2 microscope with a camera-lucida attachment.


As a result of a morphological study, Liabum diehlii is excluded from the genus Liabum and the new genus Inkaliabum is proposed. Inkaliabum is morphologically closely related to Liabum, Oligactis, and Sampera but can be easily differentiated from these genera by a combination of traits, including its slender scandent habit, pinnately veined leaves, capitulescence with few capitula, ca. 200 linear phyllaries, long style branches, and eglandular cypselae. More characters comparing Inkaliabum with the genera Liabum, Oligactis, Sampera, Dillandia, and Ferreyranthus H. Rob. & Brettell, which belong to a monophyletic clade in the more recent phylogeny of Liabeae (Dillon et al., 2009), are shown in Table 1.

Table 1. Characters comparing Inkaliabum with Dillandia, Ferreyranthus, Liabum, Oligactis, and Sampera.

The genus Inkaliabum is endemic to Peru, where the greatest diversity in Liabeae is found.

Key to Peruvian genera of Liabeae plus Oligactis

1. Abaxial surfaces of leaves hispide or strigose; leaves and stems with stiff hairs with enlarged bases

     2. Leaves 5-9 palmately veined; pappus of bristles or scales; cypselae usually with 4 ribs


     2'. Leaves 3-veined; pappus absent; cypselae with 2 ribs


1'. Abaxial surfaces of leaves with white or brown tomentum; leaves and stems without stiff hairs with enlarged bases

     2. Anther thecae dark brown or black

          3. Herbs perennial; leaves in a rosette or grouped close together on a short stem; pappus white


          3'. Usually shrubs or subshrubs, sometimes annual or perennial herbs; leaves cauline; pappus yellow or orange


     2'. Anther thecae pale yellow

          3. Leaves usually 3-veined, sometimes slightly 5 palmately veined, rarely pinnately veined

               4. Base of petioles not fused into a sheath; pappus of capillary bristles or sometimes somewhat paleate bristles


              4'. Base of petioles fused into a sheath; pappus of capillary bristles or absent

                   5. Capitulescence with many capitula densely grouped; approximately 5 ray florets per capitulum;
                    cylindrical involucre; pappus absent


                   5'. Solitary capitula on leafy peduncles or few (2 or 3) capitula laxly grouped; 20-40 ray florets per capitulum;
                   campanulate involucre; pappus of denticulate or plumose capillary bristles

                        6. Disc corollas yellow; pappus of denticulate capillary bristles


                        6'. Disc corollas reddish; pappus of plumose capillary bristles


          3'. Leaves conspicuously pinnately veined

              4. Herbs perennial, acaulescent or caulescent

                   5. Leaves in a rosettte; capitula sessile or subsessile


                   5'. Leaves cauline or grouped close together on the stem; capitula pedunculate

                        6. Ray florets of corolla red


                        6'. Ray florets of corolla yellow


              4'. Shrubs, subshrubs, vines or small trees

                   5. Shrubs or small trees; bases of petiole pair fused into a sheath;
                   crystals of cypsela walls elongate; cypselae glandular hairs with head abruptly distinct from the stalk


                   5'. Vines or scandent subshrubs; bases of petiole pair not fused into a sheath
                   (sometimes with perfoliate leaf bases); crystals of cypsela walls quadrate;
                   cypselae glandular hairs absent, or when present with head gradually distinct from the stalk

                        6. Capitulescence with few capitula (2 or 3); phyllaries ca. 200 linear or narrowly long-ovate;
                         cypselae without glandular hairs


                        6'. Capitulescence with several capitula (usually more than 50);
                         phyllaries 25-55 narrowly ovate or ovate; cypselae with glandular hairs

                             7. Slender vines; axillary and/or terminal, racemose, spiciform or glomerulose cymes;
                             capitula 6-15-flowered; anther apical appendages papillose


                             7'. Robust vines; terminal, corymbiform cymes; capitula 16-42-flowered;
                              anther apical appendages smooth


Taxonomic treatment

Inkaliabum D. G. Gut., gen. nov. Type: Liabum diehlii H. Rob., Phytologia 46: 99. 1980. Inkaliabum diehlii (H. Rob.) D. G. Gut.

Lianae vel frutices scandentes. Folia opposita sparsa subsessilia anguste elliptica, supra glabra, subtus albotomentosa, pinnatinervia, falsis stipulis instructa. Capitula in cymis umbelliformibus paucicephalis vel geminata. Capitula heterogama, involucro late campanulato multiseriato, bracteis anguste ovatis vel linearibus apice attenuato. Flores radii ca. 50 et disci ca. 200. Antherae base digitata; rami stigmatici longi involuti pubescentes. Achaenia pilis geminis vestita. Pappus setis scabrosis inaequalibus.

Slender scandent subshrubs or vines. Stems with dense and persistent pubescence. Pseudostipules forming nodal discs. Leaves opposite, sparsely arranged, sessile, elliptical or narrowly ovate, bases clasping, apex attenuate, margins sparcely mucronatedentate, revolute, adaxial surface glabrous, abaxial surface albotomentous, pinnately veined (camptodromous). Capitulescence with few capitula (2 or 3) grouped in umbelliform or geminate cymes. Capitula heterogamous, radiate, long or shortly pedunculate. Involucre widely campanulate; phyllaries imbricate in 5-7 series, linear or narrowly long-ovate, apex attenuate. Receptacle fimbriate or setiferous with laciniate setae. Ray florets ca. 50, pistillate, fertile, with corollas with limb 3-dentate, yellow; style 2-branched with style branches elongate, coiled, externally glabrous, internally with the stigmatic surface continuous. Disc florets ca. 200, hermaphroditic, fertile, with corollas tubularfunnelform slightly differentiated into tube and limb, deeply 5-lobed, yellow; stamens with anthers pale yellow, linear, shortly calcarate and shortly caudate, digitate at the base, apical appendages smooth, planate, ovate, apically rounded; style 2-branched with style branches coiled, longer than the pubescent part of the style shaft; externally, branches hispidulous, with sweeping hairs formed by 1-celled papillae, triangular in shape, apically acute, and internally with the stigmatic surface continuous. Cypselae cylindrical or obconical, brown, with conspicuous carpopodium, costate, pubescent with twin hairs, eglandular, with quadrate crystals. Pappus 2-seriate, with capillary scabrous bristles, pale yellow, outer series short, few or absent, inner series long, numerous.

Distribution. Inkaliabum is endemic to the Department of Cuzco, Peru (Fig. 1) and inhabits in valleys and slopes of the Andes.

Fig. 1.
Geographical distribution of Inkaliabum diehlii.

Etimology. The generic name refers to Liabum, the genus from where the species of Inkaliabum was originally described, and "Inka", the name of the original people where Inkaliabum occurs.

Inkaliabum diehlii (H. Rob.) D. G. Gut., comb. nov. Basionym: Liabum diehlii H. Rob., Phytologia 46: 99. 1980. Type: PERÚ. Cuzco. Prov. La Convención, Quellouno, 750 m, 22-V-1930, C. Bues s.n. (F!, holotype) (Figs. 2, 3).

Fig. 2.
Holotype of Inkaliabum diehlii conserved in F.

Fig. 3.
Inkaliabum diehlii. A. flowering branch with a terminal umbelliform cyme; B. involucre; C. inner (left) and outer (right) phyllaries; D. disc floret without cypsela; E. ray floret without cypsela; F. stamen; G. disc floret style, showing upper shaft and style branches; H. cypsela and pappus (Bues s.n., F); I. flowering brach with an axillary and terminal cyme with geminated capitula (Diehl 2450a, F) (Scale bars: A-I: 1 mm; scale bars are the same for D-E and F-G).

Slender scandent subshrubs or vines. Stems terete or slightly hexagonal, densely and persistently albotomentose. Pseudostipules forming nodal discs up to 0.9 cm long on terminal branches, without auricles, adaxial surface glabrescent and abaxial surface densely and persistently albotomentose. Leaves distributed evenly along terminal branches, sessile, 7-15 x 1.5-2 cm, blades narrowly elliptical, narrowly ovate, or linear, bases decurrent, apices attenuate, margins mucronulate, slightly revolute, adaxial surface glabrous, sometimes with a persistent arachnoid pubescence, abaxial surface densely and persistently albotomentose, chartaceous; pinnately veined (camptodromous) with 7-10 secondary veins on each side of the midrib. Capitulescence of 2 or 3 capitula in axillary or terminal cymes (mostly in an umbelliform cyme, rarely in a pair with one capitulum subsessil and the other long pedunculate), sometimes with a pair of leafshaped bracts on the peduncles near the involucre; peduncles, when present, 1.5-3 cm long, persistently and densely albotomentose. Involucre 11-14 x 10-15.5 mm, widely campanulate, phyllaries ca. 200, 3.5-13 x ca. 0.5 mm, slightly wavy, imbricate in 5-7 series, outer series subulate to linear with attenuate apices, arachnoid pubesent, and inner series narrowly long-ovate or linear with attenuate apices, glabrous, pale green. Receptacle fimbriate or setiferous with laciniate setae, 1-1.2 mm long. Ray florets ca. 50, female, fertile, corolla 9-12.5 mm long, true ray, glabrous, yellow, tube 5-6 x 0.25-0.3 mm, limb 4.5-6.5 x 1-1.5 mm, obovate, 4-veined, apex 3-dentate with three reduced equal lobes; style ca. 10 mm long, style branches ca. 2.8 mm long, glabrous. Disc florets ca. 200, hermaphroditic, fertile, corolla 8.5-9.5 mm long, tubulose-funnelform, deeply 5-lobed, slightly and gradually diferentiate into tube and limb, glabrous, yellow, tube ca. 5.5 x ca. 0.25 mm, limb 3.5-4 x 0.5-0.7 mm; lobes 1.5-2 x ca. 0.2 mm, apex pubescent; stames with thecae ca. 3.5 x 0.25 mm, pale yellow, apical appendages ca. 0.47 x 0.15 mm, ovate and smooth; style ca. 10 mm long, styles branches ca. 2.6 mm long, pubescent. Cypselae 0.9-1.2 x 0.4-0.6 mm, cylindrical or obconical, densely setulose with twin hairs. Pappus 0-7,5 mm long, outer series with short bristles up to 1.5 mm long or absent, inner series 7-7.5 mm long, pale yellow.

An interesting morphological variation in the specimens of Inkaliabum diehlii was found in its capitulescence. It is a terminal umbelliform cyme with two or three subsessile capitula or with short peduncles of equal length, each one with two leafy bracts near the involucre or without them. However, in the specimen Diehl 2450a the capitulescence is an axillary and terminal cyme with paired capitula, one of them subsessile and the other one long-pedunculate, without leafy bracts (Fig. 3. I).

Aditional material examined. PERU. Cuzco: Prov. Calca, Hacienda Pavayoc, Valle Lares, IX-1925, A. Diehl 2406 (F); prov. La Convención, Quebrada Versalles, IX-1925, A. Diehl 2450a (F); unknown locality (probably Pillpinto in prov. Paruro), 700 m, 13-V-1930, C. Bues s.n. (F).


Thanks are given to Daniel Potter, Liliana Katinas, Gisela Sancho, Mariana A. Grossi, and the reviewers for helpful comments on this manuscript. Also, I thank Michael O. Dillon and Anna Balla for assistance with the specimens of F, M. Alejandra Migoya for the illustration, Daniel A. Giuliano for the Latin diagnosis, and curators. This work was supported by the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina.


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Recibido el 29 de abril de 2010
Aceptado el 28 de junio de 2010.

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