Corresponding author: Diego F. Morales-Briones (
Academic editor: Alexander Sennikov
Three new species of
Tres nuevas especies de
Morales-Briones DF, Romoleroux K, Tank DC (2019) Three new species of
The genus
The taxonomy of
Based on molecular phylogenetic evidence,
Here, we describe and illustrate three new species of
We used previously published datasets from
Morphological characters were studied using dried herbarium specimens of
The final
Bayesian 50% majority rule consensus tree of the
The final
Bayesian 50% majority rule consensus tree of the
Previous phylogenetic analysis of
Finally,
Ascendent subshrubs; stems erect to slightly decumbent, up to 22–27 cm long, robust, densely sericeous-villous, branched at apex. Basal stipules usually caducous, if present 5–6 mm long, adnate to the petiole at base, free at apex, entirely membranous, brown. Basal leaves usually caducous, if present 3–6-lobed, 4–6 × 3–6 mm; basal petiole 3–4 mm long. Distal leaves reduced, adnate, and connate to the distal stipules forming verticillate, lobed sheaths; sheath lobes 7–8, ascending to spreading at maturity, lanceolate, one lobe 3–5 lobate, lobes (4) 6–8 × 1–2 (4) mm, (2/3 of the entire sheath length), chartaceous, margin revolute, lower surface appressed sericeous-villous to villous, upper surface sericeous-villous to glabrescent. Inflorescences terminal or axilar ± glomerulate cymes; floral bracts lobed, ascending; 8–10 flowers per inflorescence; pedicels 0.3–0.4 mm long, pilose. Flowers 1.5–2.5 mm long; hypanthium turbinate-campanulate, brown-reddish at base, pilose-sericeous outside slightly glabrescent at base, glabrous within; 4 episepals and 4 sepals connivent to ± straight, abaxially sericeous, adaxially glabrous; episepals narrowly ovate, 0.8–1 × 0.2–0.3 mm, apex acute; sepals ovate, 0.5–1 × 0.4–0.5 mm, apex ± acute; stamens 2 adnate to the floral disc; carpels 2 (3), stigma clavate. Two achenes 0.8–0.9 × ca. 0.5 mm, subovoid.
The specific epithet honors Prof. Dr. Werner Rothmaler (1908–1962), a German botanist who studied
Geographic distribution of
Decumbent herbs; branches decumbent up to 12 cm long, sericeous-villous. Basal stipules 5–10 mm long, adnate to the petiole, sparsely villous, membranous, brown at base, free at apex. Basal leaves tri-parted, blade reniform in outline, 6–7 (–10) × 7.5–10 (–1.3) mm, 3 lobes, chartaceous, slightly plicate, lobes unequally obovate-rhomboid, margin dentate-incised, teeth 3–6 on each of the lobes, lower surface sparsely sericeous-villous, upper surface villous; basal petiole 5–12 mm long. Distal stipules and distal leaves reduced adnate and connate, forming verticillate lobed sheaths; sheath lobes 6–10 ascending or slightly spreading; lobes 4–7 × 1.5–3 mm, lanceolate, entire or trilobed, decreasing in size. Inflorescences axilar or terminal glomerulate cymes, flowers aggregate at the distal part. Floral bracts 2–4 mm long, free, incised, and ascending; 3–7 flowers per inflorescence; pedicels 0.5–1 mm long, slightly pilose. Flowers 2–2.5 mm long; hypanthium turbinate to urceolate, green or reddish, pilose-sericeous outside, glabrous within; 4 episepals and 4 sepals green or slightly reddish, straight, abaxially pilose-sericeous, adaxially glabrous; episepals triangular, 0.7–0.8 × 0.4–0. 5 mm, apex acute; sepals triangular-ovate, 0.75–0.85 × 0.5–0.6 mm, apex acute; stamens 2 adnate to the floral disc; carpels 2–3, stigma clavate. Two achenes, ca. 1–1.5 × 0.6–0.8 mm, globose-ovoid.
the following specimens are collections with two different species under the same number, one (a) correspond to
The following specimens have similar habit, leaf shape, and pubescence to
The specific epithet refers to the silvery aspect of the basal leaves.
1 | Subshrubs with erect or decumbent stems; basal leaves caducous or sessile when present; stem leaves reduced, with the adnate stipule forming verticillate sheaths along the entire stem |
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– | Herbs with procumbent or decumbent stems, or subshrubs with suberect or ± decumbent stems; basal leaves petiolate; lower stem leaves subequal to or often longer than the stipules, 3-parted or 3-lobed, uppermost stem leaves reduced, with the adnate stipule forming verticillate sheaths |
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2 | Outer surface of sheath lobes glabrous or puberulent; lobes 4–6 |
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– | Outer surface of sheath lobes conspicuously pubescent; lobes 4–15 |
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3 | Outer surface of sheath lobes glabrous and reticulate, margins strongly revolute |
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– | Outer surface of sheath lobes hispidulous and not reticulate, margins revolute at apex |
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4 | Plants pilose, villous or sericeous; sheath lobes laxly ascending to erect, basal leaves mainly caducous |
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– | Plants hispid, hirsute or villous; sheath lobes widely spreading to abruptly reflexed; basal leaves usually present (except in |
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5 | Stems sparsely villous, pilose to glabrescent, or sericeous; sheath lobes 10–15, linear to linear-lanceolate, all lobes entire |
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– | Stems densely sericeous-villous; sheath lobes 7–8, lanceolate, one lobe 3–5 lobate |
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6 | Sheath lobes 7–8 (10); stems villous; sepals and episepals divergent and equal in length; hypanthium villous |
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– | Sheath lobes 9–13; stems hispid or hirsute; sepals and episepals connivent, episepals usually a little shorter than the sepals |
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7 | Plant copiously hirsute; sheath lobes reflexed; hypanthium glabrous |
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– | Plant sparsely hispid; sheath lobes spreading; hypanthium glabrous or pubescent |
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8 | Caespitose herbs; stems procumbent or slightly decumbent, multiple branched and rooting |
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– | Herbs or subshrubs; stems suberect or ± decumbent, not branched and not rooting |
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9 | Flowers ca. 2–2.5 mm long, hypanthium campanulate, appressed villous, carpels 5–10; stigma clavate |
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– | Flowers ca. 3.5 mm long, hypanthium globose-urceolate, sericeous-villous, carpels 5; stigma subclavate |
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10 | Basal leaves 5-parted, coriaceous; flowers ca. 2.5–3.5 long; carpels 5–6 |
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– | Basal leaves 3-parted, not coriaceous; flowers ca. 2–2.5 long; carpels 2–4 |
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11 | Herbs, sericeous-villous; stems decumbent; sheath lobes 6–10, ascending or slightly spreading |
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– | Subshrubs, villous or hirsute; stems suberect or ascending, sheath lobes 5–8, loosely ascending to reflex |
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12 | Stems suberect; sheath lobes 5–7; hypanthium campanulate-globose; carpels 2–3 |
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– | Stems ascending; sheath lobes 8; hypanthium globose-urceolate; carpels 3–4 |
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Rosette herbs up to 10 cm long, branches decumbent, sparsely hirsute. Basal stipules 10 mm long, adnate to the petiole, sparsely hirsute, membranaceous, brown at base, free at apex. Basal leaves tri-parted, blade reniform in outline, 17–20 × 25–26 mm, 3 lobes, chartaceous, slightly plicate, lobes unequally obovate-rhomboid, lateral lobes divided the length 1/2 of the blade, margin dentate-incised, teeth 8–10 on each of the lobes, lower and upper surface sparsely hirsute-villous; basal petiole 12–20 mm long. Distal stipules 7–8 mm long, connate and adnate to the petiole at base, free and incised at apex, leaf-like in texture. Distal leaves tri-parted, 8–10 × 8–12 mm, lobes obovate-rhomboid, decreasing in size; distal petiole 1–2.5 mm long. Inflorescences axilar or terminal glomerulate cymes, flowers aggregate at the distal part. Floral bracts 1–2 mm long, free, incised, and ascending to slightly spreading; 2–6 flowers per inflorescence; pedicels 0.6–1 mm long, sericeous. Flowers 2–2.5 mm long; hypanthium turbinate-campanulate, green, glabrous outside and within; 4 episepals and sepals green to reddish at apex, straight, abaxially and adaxially glabrous; episepals triangular, 0.7–0.8 × 0.3–0.4 mm, apex acute; sepals triangular-ovate, 0.7–0.8 × 0.5 mm, apex acute; stamens 2 adnate to the floral disc; carpels 2, stigma clavate. Two achenes, ca. 1 × 0.7–0.8 mm, globose-ovoid.
The specific epithet refers to the blue-green color of the leaves.
1 | Herbs with creeping stems, mat-forming; lateral segments of the leaves entire; inflorescences loosely dichotomous; stipules entirely membranaceous |
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– | Herbs with prostrate or loosely decumbent stems, villous or hirsute; lateral segments of the leaves variously lobed; inflorescences glomerulate cymes; stipules herbaceous at least at apex |
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2 | Basal leaves lobes acute; flowers ca. 3 mm long, hirsute-villous to sericeous; stamens 2–4 |
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– | Basal leaves lobes obtuse; flowers ca. 2–2.5 mm long, glabrous or with scattering hairs on the hypanthium lobes; stamens 2 |
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3 | Plants densely sericeous-villous; sepals oblong-ovate |
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– | Plants hirsute or glabrate; sepals ovate |
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4 | Basal leaves subreniform; cauline leaves mainly subsessile; episepals reduced or absent; carpels 1–2 |
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– | Basal leaves orbicular or reniform; cauline leaves mainly petiolate; episepals conspicuous, carpels 2–4 |
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5 | Basal leaf blade reniform; hypanthium turbinate-campanulate; carpels 2 |
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– | Basal leaf blade orbicular; hypanthium oblong-ventricose; carpels 2–4 |
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We are grateful to the curators of ANDES, CAS, COL, F, MEXU, HUT, HAO, ID, JE, MO, NY, QCA, TEX and USM for access or loans of their collections. We also thank Santiago Cordero and Carla Rodríguez for the illustrations, Daniela Cevallos for the Type images, and Simon Uribe-Convers for assistance during fieldwork. Funding for this work was provided by a Secretaría de Educación Superior, Ciencia, Tecnología e Innovación (SENESCYT) doctoral scholarship sponsored by the Instituto Nacional de Biodiversidad (INABIO) to D.F.M-B., Graduate Student Research Grants from the Botanical Society of America (BSA), the American Society of Plant Taxonomists (ASPT), the International Association of Plant Taxonomist (IAPT), and the University of Idaho Stillinger Herbarium Expedition Funds to DFM-B, and a National Science Foundation Doctoral Dissertation Improvement Grant to DCT for DFM-B (DEB-1502049). KR was funded by the Pontificia Universidad Católica del Ecuador and the SENESCYT’s ‘Arca de Noé’ Initiative-Project. Access to genomic and computational resources was granted through the University of Idaho Institute for Bioinformatics and Evolutionary Studies (IBEST) supported by an Institutional Development Award (IDeA) from the National Institute of General Medical Sciences of the National Institutes of Health (P30 GM103324).
These authors contributed equally to this work