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An unusual new record of Baccharis (Asteraceae) from the Peruvian Andes and its relation with the northern limit of the dry puna


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Background and Aims: The knowledge of the richness and distribution of the flora in the Peruvian Andes is scarce; about 70 taxonomic novelties have been reported for this area in the last decade, a large part belonging to the Asteraceae. The objective of this paper is to contribute to the knowledge of this flora through a new record of Baccharis acaulis. In addition, the distribution pattern of this species is examined. Methods: As part of floristic inventories in the high Andes of Peru, various field trips were carried out, several plant specimens were collected, among them a previously unreported species, which was identified by consulting some specialists and confirmed using taxonomic keys. Furthermore, predicting species distribution based on ecological niche modeling was made use the machine learning algorithm Maxent with the bioclimatic variables of the WorldClim database. Key results: A species of the genus Baccharis was registered for the first time for the flora of Peru. This species is recognized for its unique habit; it is a rhizomatous herb in which the aboveground part consists of small leaf rosettes surrounding a solitary capitulum. The species was collected in the south of the Peruvian Andes. This record extends the known distribution of the species considerably to the north. Conclusions: This record suggests the need to continue studying the Andean flora, as well as the links of their distribution patterns with the phytogeographical regions.
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Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 1
Artículo de investigación
An unusual new record of Baccharis (Asteraceae) from the
Peruvian Andes and its relation with the northern limit of
the dry puna
Un inusual nuevo registro de Baccharis (Asteraceae) de
los Andes peruanos y su relación con el límite norte de la
puna seca
Paúl Gonzáles1,4 , Asunción Cano1, 2 , Jochen Müller3
Background and Aims: The knowledge of the richness and distribuon of the ora in the Peruvian Andes is scarce; about 70 taxonomic noveles have
been reported for this area in the last decade, a large part belonging to the Asteraceae. The objecve of this paper is to contribute to the knowledge
of this ora through a new record of Baccharis acaulis. In addion, the distribuon paern of this species is examined.
Methods: As part of orisc inventories in the high Andes of Peru, various eld trips were carried out, several plant specimens were collected, among
them a previously unreported species, which was idened by consulng some specialists and conrmed using taxonomic keys. Furthermore, pre-
dicng species distribuon based on ecological niche modeling was made use the machine learning algorithm Maxent with the bioclimac variables
of the WorldClim database.
Key results: A species of the genus Baccharis was registered for the rst me for the ora of Peru. This species is recognized for its unique habit; it is
a rhizomatous herb in which the aboveground part consists of small leaf rosees surrounding a solitary capitulum. The species was collected in the
south of the Peruvian Andes. This record extends the known distribuon of the species considerably to the north.
Conclusions: This record suggests the need to connue studying the Andean ora, as well as the links of their distribuon paerns with the phyto-
geographical regions.
Key words: Compositae, ora, Heterothalamus, new record, phytogeography.
Antecedentes y Objevos: El conocimiento de la riqueza y distribución de la ora en los Andes peruanos es escaso; en la úlma década han habido
alrededor de 70 novedades taxonómicas para esta área, siendo la familia Asteraceae la de mayor aporte. El objevo de este trabajo es contribuir al
conocimiento de esta ora, a través de un nuevo registro de Baccharis acaulis. Además, se examina su patrón de distribución.
Métodos: Como parte de los inventarios oríscos en los altos Andes del Perú, se realizaron numerosos viajes de campo, se recolectaron varios es-
pecímenes de plantas, entre ellos una especie no reportada previamente, que se idencó mediante la consulta a algunos especialistas y se conrmó
ulizando claves taxonómicas. Además, se predijo la distribución de la especie basada en modelos de nichos ecológicos, ulizando el algoritmo de
aprendizaje automáco Maxent con las variables bioclimácas de la base de datos WorldClim.
Resultados clave: Se registró por primera vez para la ora del Perú una especie del género Baccharis. Esta especie se disngue por su singular hábito,
es una hierba rizomatoza en la que la parte supercial consiste en pequeñas rosetas de hojas que rodean a un capítulo solitario. La especie fue reco-
lectada al sur de los Andes peruanos. El registro exende la distribución conocida de la especie a una zona más norteña.
Conclusiones: Este registro sugiere la necesidad de connuar estudiando la ora andina, así como los vínculos de sus patrones de distribución con
las regiones togeográcas.
Palabras clave: Compositae, togeograa, ora, Heterothalamus, nuevo registro.
1 Universidad Nacional Mayor de San Marcos, De-
partamento de Dicoledóneas, Museo de Historia
Natural, Laboratorio de Florísca, Av. Arenales 1256,
Jesús María, Lima, Peru.
2 Universidad Nacional Mayor de San Marcos, Facultad
de Ciencias Biológicas, Instuto de Invesgación de
Ciencias Biológicas Antonio Raimondi (ICBAR), Lima,
3 Friedrich-Schiller-Universität, Herbarium Haussk-
necht, Fürstengraben 1, 07737 Jena, Germany.
4 Author for correspondence: pgonzalesarce@hot-
Received: May 17, 2018.
Reviewed: June 25, 2018.
Accepted by Marie-Stéphanie Samain: August 6, 2018.
Online rst: October 10, 2018.
Published: Acta Botanica Mexicana 126 (2019).
To cite as:
Gonzáles, P., A. Cano and J. Müller. 2018(2019). An
unusual record of Baccharis (Asteraceae) from the Pe-
ruvian Andes and its relaon with the northern limit
of the dry puna. Acta Botanica Mexicana 126: e1393.
DOI: 10.21829/abm126.2019.1393
This is an open access arcle under the Cre-
ave Commons 4.0 Atribuon-Non commer-
cial License (CC BY-NC 4.0 Internaonal). e-ISSN: 2448-7589
Gonzáles et al.: A new record of Baccharis (Asteraceae) from Peruvian Andes
Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 2
The knowledge of the richness and distribuon of the ora
in the Peruvian Andes is sll incomplete, which is reected
by the constant reports of new species and new records for
the country (Gonzáles et al., 2016; León et al., 2018), where
in the last decade these noveles have reached 70 species
and of which 20% corresponds to the Asteraceae (Beltrán,
2009; Montesinos-Tubée, 2014; Montesinos-Tubée et al.,
2015, 2017, 2018; Gonzáles et al. 2016; Beltrán and Gran-
da, 2017).
In the present work, Baccharis acaulis (Wedd. ex
R.E. Fr.) Cabrera, a new record for the high Andean ora of
Peru, is documented. This species had not previously been
registered by Brako and Zarucchi (1993), Ulloa Ulloa et al.
(2004; 2017), Beltrán et al. (2006), and much less for other
works that have listed new records or nomenclatural nov-
eles for the Peruvian ora (Dillon and Sagástegui, 2002;
Vásquez et al., 2002; Rodríguez et al., 2006; Linares et al.,
2010; Gonzáles et al., 2016). This nding is the result of a
study whose purpose is to develop the checklist of vascular
plants of the Peruvian Andes.
Baccharis L. is a large genus of the tribe Astereae
with 300 to more than 500 species (Malagarriga, 1976; Ne-
som, 1990; Bremer, 1994; Hellwig, 1996) widely distributed
in America, but with a mayor diversity throughout the An-
des (Müller, 2006). The genus contains about 77 species in
Peru (Brako and Zarucchi, 1993; Ulloa Ulloa et al., 2004), 17
of them being endemic to Peru (Beltrán et al., 2006).
Baccharis acaulis was inially described and valid-
ly published as Psila caespitosa Phil. (Philippi, 1891), and
subsequently as Heterothalamus acaulis Wedd. ex R.E. Fr.
(Fries, 1905); however, due to its paleate female capitula,
it has later been subordinated to Pseudobaccharis Cabre-
ra (Cabrera, 1944). Finally, it was located inside Baccharis
(Cabrera, 1975).
This taxon had only been previously menoned as
growing in Peru (Müller, 2013), but no herbarium material
was known to support its presence, which is why we make
this formal as a new record. Therefore, the objecve of this
paper is to contribute to the knowledge of the Peruvian o-
ra through a new record of Baccharis acaulis. In addion,
the distribuon paern of this species is examined in func-
on of its habitats of occurrence.
Material and Methods
Field work and herbarium study
In recent exploraons realized in the department of Cusco
(South Peru), we collected a specimen of Asteraceae that
at rst glance did not t within the known taxa for this area
of Peru. A review of available literature and consultaons
with colleagues was necessary. The collecons were depos-
ited in the herbarium of the Museum of Natural History of
the Naonal University of San Marcos (USM). Addionally,
we consulted the principal Peruvian herbaria (CPUN, CUZ,
HUSA, HUT, MOL, and USM) to nd more records. These
herbarium specimens were studied under a stereomicro-
scope (Leica-EZ4 1X–4.5X, Leica Microsystems, Wetzlar,
Potential distribution model
The locaons in which the species has been registered
were obtained by consulng the collecons of the herbaria
US and USM.
The machine learning algorithm Maxent version
3.3.3e (Phillips et al., 2006) was used for developing the
habitat suitability maps, for being one of the most reliable
methods when working with a small number of occurrence
records (Elith et al., 2006; Hernandez et al., 2006; Benito de
Pando and Peñas de Giles, 2007; Pearson et al., 2007; Wisz
et al., 2008; Särkinen et al., 2013). We ran Maxent based
on the 25 records from Peru, Bolivia, Chile and Argenna to
idenfy potenal suitable habitat areas for Baccharis acau-
lis. The model was run with default sengs (see details in
Särkinen et al., 2013). To evaluate model performance, we
ran the model with cross-validaon because it uses all of
the data for validaon. For the interpretaon of the mod-
el we used the receiver operang characterisc (ROC) and
the area under the curve (AUC) (Hanley and McNeil, 1982;
Fielding and Bell, 1997; Segurado and Araújo, 2004; Phillips
et al., 2006; Phillips and Dudík, 2008). Values of AUC close
to 1 indicate an opmal funconing of the model, values
close to 0.5 indicate performance equal to random (Phillips
et al., 2006).
The model was run with 19 bioclimac variables,
based on the interpolaon of temperature and precipita-
on data of the WorldClim database (Hijmans et al., 2005)
Gonzáles et al.: A new record of Baccharis (Asteraceae) from Peruvian Andes
Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 3
with a 30 arc second spaal resoluon (ca. 1 km2) (Hijmans
et al., 2005). Aer the jackknife analysis, 10 variables were
selected to run the nal model. The remaining 10 variables
included layers describing the seasonality of the habitat
(bio1, bio2, bio3, bio5, bio6, bio9, bio10, bio11, bio13,
bio14). A nal distribuon map for B. acaulis was produced
based on the logisc output format, where all areas with
relave suitability greater than 50% were considered as po-
tenal areas of occurrence for the species.
Taxonomical details
Baccharis acaulis (Wedd. ex R.E. Fr.) Cabrera, Bol. Soc. Ar-
gent. Bot. 16(3): 255. 1975. (Fig. 1).
= Psila caespitosa Phil. Anales Mus. Nac. Sanago de Chile
Botánica 1891(8): 36, t. 1, f. 4. 1891. SYNTYPES: Chile. Re-
gion Antofagasta, province San Pedro de Atacama, planta
femenina de Machuca (small Andean village), 3200 m, plan-
ta hermaphrodita de Guanaqueros allatae sunt, without
date, Philippi s.n. (SGO-43827, SGO-64394, LP-002306!).
Heterothalamus acaulis Wedd. ex R.E. Fr., Nova Acta Regi-
ae Soc. Sci. Upsal., ser. 4, 1: 79. 1905.
Pseudobaccharis acaulis (Wedd. ex R.E. Fr.) Cabrera, Notas
Mus. La Plata 9: 248. 1944. TYPE: Argenna. Province Jujuy,
department Tumbaya, locality Moreno, 3500 m, 23.X.1901,
masculine owers, Fries 701 (lectotype: (designated by
Müller, 2006) S 04-474!, isolectotype: US 00129267!). Illus-
traons: Philippi (1891: Tab. I, Figs. 4A-H), Cabrera (1978:
Figs. 90A-H), Müller (2006: Figs. 113A-O).
Perennial herb, rhizomatous, creeping, low-growing,
forming mats 0.5-2 cm tall; plants with long pale subterra-
neous rhizomes; leaves sessile, rosee-forming, linear, 5-20
× 0.1-1 mm, sheathing at the base, apex obtuse, eshy,
margin enre, 3-veined at base (1-veined at lamina); syn-
orescences of solitary sessile terminal capitula, solitary at
the apex of dense leaf rosees; male capitula hemispheri-
cal, 4-7 mm long, receptacle without palea, owers 20-35,
involucres 4-6 × 3-5 mm, phyllaries 15-25 in 3-4 series, the
outermost phyllaries narrowly ovate and acute, median
phyllaries ellipc, broadly lanceolate or ovate, innermost
phyllaries longer than outermost ones, linear-oblong, ob-
tuse or subobtuse, all phyllaries whish green, apically
nged with purple, phyllaries margin rather broadly scari-
ous, short to long-dentate in the distal poron; corolla 3-4
mm long, tube 1.2-1.8 mm long, throat poron 1-1.5 mm
long, campanulate, lobes 0.8-1.1 mm long, curved at ma-
turity, pubescent in distal half, anther length double that
of laments; style not or slightly exceeding the corolla,
sterile ovary glabrous, pappus with inconspicuously broad-
ened bristles at apex, 2.4 mm long, bristles 25-50; female
capitula hemispherical, 4-9 mm long, receptacle with linear
palea, owers 30-50, involucre 3-7 × 3-4 mm, cup shaped,
phyllaries 25-40 in 2-4 series, such as the male; corollas
2.5-4.5 mm long, apex inconspicuously denculate, style
branches short but separate, achenes 1-2 mm long, 5-an-
gled, glabrous, pappus 2-seriate, accrescent.
Material examined: ARGENTINA. Province Catamar-
ca, department Belén, Reserva Laguna Blanca, 3260 m,
01.VI.2002, M. Borgnia 6 (LP). Province Jujuy, department
Humahuaca, 3529 m, B. Ruthsatz 157 (LP). Department
Tumbaya, locality Moreno, 3500 m, 23.X.2001, R . E. Fries 701
(CORD, S, US). Province Salta, department Los Andes, 3800
m, A. L. Cabrera 10608 (LP); Los Andes, Mina Sijes, 3800
m, 02.XII.1988, A. Charpin 20789 (SI, US 3107570); Mina
Sijes, 3800 m, 3.XII.1986, L. Novara et al. 5742 (MCNS); San
Antonio de Los Cobres, 3800 m, 21.XII.1948, M. H. Humbert
21077 (P-P00218234); San Antonio de los Cobres, puna de
Jujuy, 21.X.1948, R. E. Fries s.n. (S); 3800 m, 03.XII.1934, E. J.
Ringuelet 116 (LP). Department Rosario de Lerma, puna de
Jujuy, Diego de Almagro, 21.X.1948, R. E. Fries s.n. (S); loc.
cit., A. L. Cabrera 10612 (LP). BOLIVIA. Department Cocha-
bamba, province Carrasco, Siberia, 17°49'47"S-64°44'07"W,
2940 m, 20.II.2005, S. Altamirano Azurduy 3077 (BOLV); Ca-
rrasco, Siberia, 17°49'47"S-64°44'07"W, 2940 m, 20.II.2005,
E. Fernández Terrazas 3216 (BOLV). Province Tapacarí, Jacha
Rancho, Comunidad de Japo, km 125 Cochabamba - Oruro,
4000 m, A. Pestalozzi 866 (LPB); Campanani, Comunidad de
Japo, km 125 Cochabamba - Oruro, 4180 m, 19.XI.1995, H.
U. Pestalozzi 735 (BOLV); Comunidad de Japo, km 125 Co-
chabamba - Oruro, Jacha Rancho, 4200 m, 18.IX.1996; H. U.
Pestalozzi 867 (LPB). Department La Paz, province Aroma,
Umala, 3810 m, K. Graf 172 (NY). Province Larecaja, Viciniis
Gonzáles et al.: A new record of Baccharis (Asteraceae) from Peruvian Andes
Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 4
Figure 1: Baccharis acaulis (Wedd. ex R.E. Fr.) Cabrera; A. habit; B. details of rhizomatous plant; C. details of male flowering branches; D. details of
female flowering branches; E. capitulum showing the rows of phyllaries; F. capitulum of male flowers; G. details of male flowers; H. fruits; I. leaf
(upper side), J. details of phyllaries.
Gonzáles et al.: A new record of Baccharis (Asteraceae) from Peruvian Andes
Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 5
Sorata, prope Apacheta de Logena, in graminosis, 4000 m,
VIII.1859, G. Mandon 109 (G, NY, P-P00218233, S). Province
Pedro Domingo Murillo, 5.5 km E of the juncon with road
to Collana, on road between Cota Cota and Palca, open,
heavily grazed slopes with some boggy areas along streams
and seeps, 16°32'S, 67°58'W, 3900 m, 25.IX.1982, J. C. So-
lomon 8248 (MO 3681776, NY, US); “Plain above La Paz, in
damp places near borders of lakes, 14500 ., R. Pearce s.n.
(BM); La Paz, 3900 m, Williams 2342 (NY). Department Oru-
ro, province Avaroa, Pampa de Huancané, 3800 m, C. Troll
3024 (B). Province Sabaya, sur de Sabaya, Salar de Coipasa,
3670 m, S. G. Beck 21566 (JE, LPB). Province Sajama, near
the village of Sajama along the banks of the Rio Sajama,
4260 m, 18.X.1967, B. Vuilleumier 317 (GH, MO 3230979,
NY, US). Province Sur Carangas, S de Andamarca, Laguna Ta-
cata, 3700 m, Stab B388 (LPB). Department Potosí, province
Antonio Quijarro, cerca de Tica Tica, camino Uyuni - Potosí,
3600 m, E. Jordan et al. 57 (LPB). Province Chayanta, 1.6 km
SW de Ocuri, 4000 m, M. Massy 65 (LPB). Province Enrique
Baldivieso, 3 km de Alota, 4000 m, M. Liberman 222 (LPB).
Department Tarija, province José María Avilés, Avilés, cerca
Passajes, borde de Laguna (sal), 3255 m, casi plano, suelo
limoso, 29.I.1986, Basan 635 (LPB); Iscayachi 23.7 km ha-
cia Villazón, Hoyada chica, 3800 m, S. G. Beck 11025 (LPB).
CHILE. Region Antofagasta, province Antofagasta, 4180 m,
23.II.2009, F. O. Zuloaga 11153 (SI). Province El Loa, Salar de
Ascotan, 21.6°S, 68.3°W, 3800 m, 01.II.1997, S. Teillier 4225
(CONC). Region Arica y Parinacota, Province Parinacota,
quebrada profunda antes de Colpitas, 4086 m, 20.X.2012,
A. Moreira 2023 (SGO); Putre, Laguna Chungará, 4590 m,
B. Ruthsatz 7990 (JE). PERÚ. Department Cusco, province
Chumbivilcas, Chamaca, arriba de Uchucarco, Laguna Yana-
cocha, 4364 m, 4.IX.2017, P. Gonzáles 4192 (USM); Chum-
bivilcas, Velille, arriba de Chilloroya, Cochapampa, 4305 m,
4.IX.2017, P. Gonzáles 4195 (USM); Chumbivilcas, Chamaca,
Puente Arizona, camino a Uchucarco, 3917 m, 4.IX.2017, P.
Gonzáles 4196 (USM), 4197 (USM); Chumbivilcas, Chama-
ca, debajo de Mina Catanga, 3973 m, 4.IX.2017, P. Gonzáles
4198 (USM), 4199 (USM); Chumbivilcas, Chamaca, debajo
de Uchucarco, puente hacia Velille, 3845 m, 4.IX.2017, P.
Gonzáles 4200 (USM), 4201 (USM). Province Espinar, Es-
pinar, Alto Huancané, 3961 m, 01.X.2015, P. Gonzáles 3850
(USM); Espinar, intersección de quebrada Tintaya con río
Salado, 3917 m, 8-18.IX.2016, P. Gonzáles 3958 (USM);
4 km de Espinar, debajo del puente en Río Salado, 3900
m, 30.VIII.2017, P. Gonzáles 4190 (USM); Espinar, 8-9 km
de Espinar, desvío a Kanamarca, 3950 m, 30.VIII.2017, P.
Gonzáles 4191 (USM). Department Puno, province Lampa,
Santa Lucia, moist environs on the side of the Río Verde,
14000 ., 2.II.1939, J. E. Sharpe 101 (K); province Puno,
San Antonio de Esquilache, bogs, 14600 ., 30.VIII.1934, D.
Staord 1121 (K).
Real and potential distribution
The distribuon of Baccharis acaulis covers the Alplano
and adjacent ranges in Peru (Cusco, Puno), Bolivia (Cocha-
bamba, La Paz, Oruro, Potosí, Tarija), northern Chile (Ari-
ca y Parinacota, Tarapacá, Antofagasta), and northwestern
Argenna (Catamarca, Jujuy, Salta). It can be found in the
high Andean wetlands, mainly at the margin of streams and
brackish lagoons at 3200-4600 m (Giuliano, 2001; Müller,
2006; Freire et al., 2011; Moreira-Muñoz et al., 2016).
The nal model yielded AUC values >0.98 indicang
good model performance. The two most important cli-
mac variables were minimal temperature of the coldest
month (bio6) and mean temperature of the coldest quarter
(bio11). Other important variables included annual mean
temperature (bio1), mean temperature of driest quarter
(bio9), precipitaon of weest month (bio13) and precip-
itaon of driest month (bio14).
Along the Andes, the potenal distribuon mod-
el indicates the existence of very suitable areas for the
presence of B. acaulis between 13° and 26°S, whose most
extensive areas are between Bolivia and Chile (17°-19°S),
as well as between Bolivia and Argenna (21°-23°S) (Fig.
2). The northern limit of the potenal area (more than
70%) of distribuon predicted along the Andes is only 1.5°
of the record found in the department of Cusco, which
shows the usefulness and eecveness of the species dis-
tribuon models.
In Peru, the potenal distribuon model showed
very suitable climac condions for B. acaulis in the de-
partments of Cusco, Puno, Apurímac, Arequipa, Ayacucho,
Moquegua, and Tacna (Fig. 2). The last ve departments
remain without conrming the presence of this species.
In addion, it idened areas that are reasonably suitable
Gonzáles et al.: A new record of Baccharis (Asteraceae) from Peruvian Andes
Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 6
Figure 2: Real and potential distribution map of Baccharis acaulis (Wedd. ex R.E. Fr.) Cabrera showing the known collections (white square). The
map shows the areas identified with very suitable habitats (>50%). The collection of Baccharis davidsonii Cuatrec. are also shown (black asterisk).
Gonzáles et al.: A new record of Baccharis (Asteraceae) from Peruvian Andes
Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 7
towards central Peru, where in a recent expedion we re-
corded a populaon of Baccharis davidsonii Cuatrec. (Lima,
Oyón, Raura, 4770 m, 17.X.2017, A. Cano et al. 22585
(USM), 22587 (USM)), a closely related species in Baccha-
ris sect. Psila (Phil.) Cuatrec. (Heiden and Pirani, 2016). The
model also predicted the presence of B. acaulis in Bolivia,
Chile and Argenna, but in these countries, there are re-
cords of specimens for the regions (Chile), departments
(Bolivia) or provinces (Argenna) where their potenal dis-
tribuon is predicted.
In recent evaluaons throughout the Peruvian Andes
and search eorts in herbaria, ve localies of B. acaulis
were recorded in the Andean zone of southern Peru. These
ve localies together with the localies of Bolivia, Chile
and Argenna suggest a connuity of the distribuon of B.
acaulis throughout the dry puna (13°-26°S).
Baccharis acaulis grows always associated with humid ar-
eas named wetlands or locally named “bofedales”; how-
ever, it is not a main component of this vegetaon type
(Gonzáles, 2015), as it is only associated with the edges of
small rivers or channels that run through the “bofedales”
without moving more than four meters from the edge. This
suggests a predilecon for permanently saturated water
soils. Baccharis acaulis also grows in brackish areas, and its
adapve mechanisms to those soil are sll unknown. On
the one hand, in the dry puna, the “bofedales” and “sala-
res” are the only environments with high water availabili-
ty, being the salt ats those that have a permanent water
regime in comparison to the “bofedales” whose regime is
very variable and temporary. On the other hand, the re-
cent record of B. davidsonii in the central Andes of Peru
corresponds to the humid puna, which presents vegetaon
types with higher humidity than the dry puna (Josse et al.,
2011). Despite the greater humidity of the “bofedales” in
the central Andes of Peru, B. davidsonii as well as B. acau-
lis connue to grow at the edges of this vegetaon type
and are always associated with water courses. In addion
to the “bofedales” in the humid puna, there are other areas
with high levels of humidity such as rocky and cryoturbated
soils, where some cushions of B. davidsonii have also been
recorded (own observaon).
Discussion and conclusion
Distinctive morphology
Baccharis acaulis diers from all other species of Baccharis
in its unique habit; it is a rhizomatous herb in which the
aboveground part consists of small leaf rosees surround-
ing a solitary capitulum (Müller, 2006).
Most species of Baccharis are woody shrubs, erect
and larger than 10 cm tall; but there are also some small
and procumbent shrubs such as Baccharis caespitosa
(Ruiz & Pav.) Pers., Baccharis alpina Kunth, Baccharis
humifusa Kunth and Baccharis pumila Joch. Müll., which
form small cushions that grows at ground level. In this
context, B. acaulis as well as B. davidsonii are unique
species, because they are the only two high Andean spe-
cies with herbaceous habit, growing like a rhizomatous
The capitulescence type in Baccharis has several
forms (panicles, racemes, spikes); however, the most pecu-
liar forms are the solitary and terminal capitula as registered
for Baccharis acaulis, B. alpina, B. davidsonii, B. pumila and
B. tola Phil. This form is very common in diverse Andean
genera of Asteraceae such as Hypochaeris L., Paranephelius
Poepp., Senecio L., Werneria Kunth, and Xenophyllum V.A.
Funk, amongst others.
The unusual characteriscs of B. acaulis have led to
its classicaon in dierent genera (Cabrera, 1944, 1955;
Cuatrecasas, 1967), and even in a dierent infrageneric
posion within Baccharis. Müller (2006) placed it in the
Baccharis juncea (Lehm.) Desf. group due to its glabrous,
eshy and narrow leaves, the pappus of female owers
which is strongly elongated at achene maturity, compris-
ing several bristle series. Giuliano (2001) placed it in the
secon Angusfoliae Baker due to its linear, one-veined
leaves, solitary capitula, corollas of female owers with
truncated apex, pappus of female owers which is bise-
riate and accrescent, and the achenes with ve sides.
Recently, it has been placed in Baccharis subgen. Hetero-
thalamus (Lessing) G. Heiden within Baccharis sect. Psi-
la, which consists of halophyllous, shrubby cryptophytes
with long and mulbranched subterraneous rhizomes
bearing deciduous scales and presenng ephemeral aeri-
al branches with solitary capitula subtended by leafy ro-
sees (Heiden and Pirani, 2016).
Gonzáles et al.: A new record of Baccharis (Asteraceae) from Peruvian Andes
Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 8
In the dry regions on the Alplano and in the Andean
southern Peru, owering periods of Asteraceae are largely
restricted to the wet season (Müller, 2006), but they may
also occur in the dry season (August to December, rarely
extended to February) as registered for B. acaulis.
Müller (2006) menons that some species of Baccharis are
restricted to the Alplano zone centred in Bolivia, southern
Peru, northern Chile and northwestern Argenna; among
these species is B. acaulis, although no collecons from
Peru were cited.
The habitat exclusively in high Andean zones of B. acau-
lis rearms the orisc anity between wetlands “bofedales”
of Bolivia, northern Chile, northern Argenna and southern
Peru, due to the homogeneity and environmental connu-
ity of the high Andean wetlands, whose bioc limits do not
necessarily respond to internaonal borders (García, 2007;
Gonzáles et al., 2016). This distribuon paern of B. acaulis
agrees with the distribuon of other typical species of bofeda-
les” such as Patosia clandesna (Phil.) Buchenau and Oxychloe
andina Phil. which have registered their northernmost distri-
buon in the south of Peru (Balslev, 1996; Ruthsatz, 2012).
Baccharis acaulis grows in humid, oen saline, soil for
which it needs to present some mechanisms to counter-
act stress condions (Lieth and Mochtchenko, 2003; Teillier
and Becerra, 2003; Ruthsatz, 2012). Frequently, salt toler-
ance is associated with tolerance to water stress; howev-
er, if salt toxicity is the main cause of the eect of salinity,
salt tolerance is not necessarily linked with water stress
tolerance (Villagra and Cavagnaro, 2006); this could be the
mechanism of B. acaulis, since it would be the most import-
ant for the ecological success of this species. Furthermore,
this salt stress tolerance could be an adapve factor in the
niche dierenaon of B. acaulis and its capacity to grow in
salt soil, since the habitats where B. acaulis grows are the
unique place with highest level of water in the dry puna.
Apparently, B. acaulis is a facultave halophyte which
can tolerate salt stress but not drought, as has been report-
ed for other species such as Suaeda salsa (L.) Pall. and Ka-
lanchoe daigremonana Raym.-Hamet & H. Perrier (Zhao
and Harris, 1992; Zhao et al., 2003). This idea is reinforced
by the lower tolerance of B. acaulis to water stress (eld
observaons), which supports the armaon that physi-
ological and ecological responses are dierent between
halophytes and xerophyc plants (Zhao and Harris, 1992;
Lieth and Mochtchenko, 2003; Zhao et al., 2003).
Baccharis acaulis should then have mechanisms al-
lowing it to survive under salt stress condions, which are
not related to the presence of salt eliminaon organs; rath-
er the higher salt tolerance of B. acaulis could be related
to its ability to counteract the toxic eect rather than the
osmoc eect of salt, as has been proposed for other An-
dean species of Prosopis L. (Villagra and Cavagnaro, 2005).
This report is relevant for contribung to the knowl-
edge of an addional species for the Peruvian ora, besides
registering the northernmost area (15°S) for the species and
showing the biogeographical importance of the dry puna
for housing species typical of this zone. It also supports the
hypothesis that this area in southern Peru would coincide
with the northern limit of the renowned dry puna (Josse
et al., 2009, 2011). However, the record of B. davidsonii in
the central Andes of Peru is more than 700 km away from
the populaons of B. acaulis in southern Peru, being the
northernmost distribuon limit of Baccharis secon Psila.
Author contributions
JM and PG performed the study of herbarium materi-
al, PG performed sampling, manuscript preparaon, and
data analysis, PG, AC and JM performed interpretaon and
preparaon of the nal version.
We thank the Vicerrectorado de Invesgación y Postgra-
do, Universidad Nacional Mayor de San Marcos, FLORAM
Research Group, by the nancial support (Grant Number
We acknowledge Luis Ariza, Daniel Giuliano, and Gustavo
Heiden for their taxonomic comments; Virginia Palche
for his comments on halophyc species; Blanca León for
her valuable observaons on the manuscript; Susy Casllo,
Gonzáles et al.: A new record of Baccharis (Asteraceae) from Peruvian Andes
Acta Botanica Mexicana 126: e1393 | 2019 | 10.21829/abm126.2019.1393 9
Huber Trinidad, Elluz Huamán, Sebasan Riva and Estefany
Ccollana for eld work support.
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... The first cluster analysis with 373 nr of relevés (153 this survey and 220 nr from literature) revealed an outgroup (61 relevés by [4,20,22,28,31,32,35,[58][59][60][61] which was removed prior to the second run with the remaining 312 relevés including the relevés from the present survey as well as relevés from literature ( Table 3, see heading of Table 3 for detailed description of the communities from other studies included in the analysis). Table 2. Table with the type relevés of associations and subassociations described as new (including one community and a pro-visional association). ...
... A synoptic-syntaxon table (Table 2) was constructed by reordering the species into syntaxonomic species groups after comparison with publications from [21,22,24,27,58,59]. ...
... New syntaxa were described following the International Code of Phytosociological Nomenclature [62] and new syntaxonomic units were defined after reviewing [21,22,24,27,58,59]. ...
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The present work is a phytosociological synthesis and syntaxonomic overview of the vegetation of the highest subnival parts (superpuna) of the open alpine vegetation of the high plateaus (puna) of the Andes of Moquegua, South West Peru, as related to the main environmental gradients. Using TWINSPAN and DCA ordination analysis, 153 phytosociological relevés were analyzed. For each association, subassociation and community, the syntaxonomy, floristic diversity and relation with environmental variables are described. The syntaxonomy and synecology of superpuna vegetation was studied in 19 localities at an altitude of 4450–4800 m. The study area has a pluviseasonal climate with yearly rainfall (December-April). Four main highland vegetation types were distinguished: 1. slope and scree chasmophyte vegetation composed of shrubs, cushions, ground rosettes and grasses, 2. grasslands (grazed and ungrazed) characterized by great species richness in shrubs, cushions, ground rosettes, grasses and herbs, 3. vegetation of plateaus with cushions, shrubs, ground rosettes, herbs and grasses and 4. nitrophilous vegetation with high cover and low species richness. Within the vegetation of the orotropical and cryorotropical bioclimatic belts three phytosociological classes can be distinguished: Argyrochosmetea niveae (chasmophytic vegetation), Calamagrostietea vicunarum (grasslands with cushions), Anthochloo lepidulae-Dielsiochloetea floribundae (highland slopes and plateaus) and a nitrophylous community. One new association from rock and scree slopes was described within the Saxifragion magellanicae (Argyrochosmetea niveae). Within the Calamagrostion minimae, which comprises grasslands with cushions and mat-forming plants, one new association with two subassociations could be distinguished. Within the grassland and cushion associations of the Azorello-Festucion (Calamagrostietea vicunarum), three new associations were described, comprising nine subassociations. In the Anthochloo-Dielsiochloetalia one new and one previously described association and one community are distinguished. In addition, the nitrophilous community of Tarasa nototrichoides and Urtica flabellata has been described. In total the vegetation comprised 172 vascular species belonging to 32 families. Our study provides the first syntaxonomic revision of chasmophytes, cushion associations and high-altitude grasslands in the Andes of North Moquegua. The proposed syntaxonomic scheme contains the associations distributed under similar habitat conditions throughout the Southern Andes of Peru, but also the associations reflecting the local floristic and environmental patterns. The subnival vegetation of Moquegua hosts some rare endangered and/or protected plant species.
The genus Baccharis is composed of ca. 440 species, distributed primarily in South and Central America, many of which are of great ecological, economic, and cultural importance. Baccharis species are mostly dioecious and highly diverse in chemistry, ecology, architecture, and phenology, occupying many different niches and habitats across several gradients of light, temperature, humidity, altitude, and succession. Its species are found in natural, urban, and highly polluted environments. Many species host a large number of associated organisms, including the largest fauna of gall-inducing insects in the Neotropics, and play crucial roles in biodiversity maintenance as foundation species or ecosystem engineers, while others are invasive species with economic implications around the world. Many species are geographically restricted or endemic. Baccharis is also well known for being the source of innumerable chemical compounds widely used in folk medicine and in the cosmetics and pharmaceutical industries. It is one of the most studied genera in the world, owing to these multiple factors that have captured the attention of the scientific community.
Baccharis is a monophyletic genus characterized by functionally unisexual florets, generally distributed in distinct individuals (dioecy), but also including monoecious, gynodioecious, and polygamous species. The genus has not been revised taxonomically as a whole for nearly two centuries. Recent country- or dependent territory-level checklists are hardly comparable and mostly outdated. A comprehensive checklist on the diversity and distributions of Baccharis at generic, infrageneric, and specific levels, including putative hybrids, and adventitious occurrences is provided. Baccharis comprises 442 species classified into 47 sections and 7 subgenera. The genus is native in the Americas, from southeastern Canada and northwestern USA to Tierra del Fuego, with species native to the Falkland/Malvinas Islands, across most of the Caribbean islands and the Galápagos archipelago. Complete lists of species per country and territory are provided. Brazil (185 species, 114 endemics), Argentina (110 species, 25 endemics), and Bolivia (76 species, 22 endemic) are the richest countries for the genus. Four species are highlighted for occurring in more than ten countries or territories within their native range (B. dioica, B. pedunculata, B. trinervis, B. salicifolia), while at least 218 species are endemic to a single country or territory. The role of hybridization in the genus diversity, ecology, and evolution is still a neglected subject, and 38 putative hybrid taxa were described so far. Some species were spread outside the American continent by anthropogenic dispersals, and at least two have established naturalized alien populations: B. halimifolia in Europe and Oceania and B. spicata in Europe.
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The cataloging of the vascular plants of the Americas has a centuries-long history, but it is only in recent decades that an overview of the entire flora has become possible. We present an integrated assessment of all known native species of vascular plants in the Americas. Twelve regional and national checklists, prepared over the past 25 years and including two large ongoing flora projects, were merged into a single list. Our publicly searchable checklist includes 124,993 species, 6227 genera, and 355 families, which correspond to 33% of the 383,671 vascular plant species known worldwide. In the past 25 years, the rate at which new species descriptions are added has averaged 744 annually for the Americas, and we can expect the total to reach about 150,000.
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Senecio sagasteguii y Senecio qosqoensis (Asteraceae, Senecioneae), dos nuevas especies procedentes de la región Cusco, en el sureste del Perú, son descritas e ilustradas. Los nuevos taxones son asignados a Senecio sección Senecio serie Chilenses subserie Caespitosi en virtud de su hábito subarbustivo y sus capítulos discoides solitarios o escasos. Se incluyen algunos comentarios sobre sus posibles afinidades morfológicas, su distribución geográfica y sus preferencias ecológicas.
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The family of the Asteraceae is the most diverse in the Chilean flora; nevertheless, only some tribes have undergone a recent systematic treatment, while the knowledge about the geographical distribution of the taxa is still very incomplete. From the review of herbarium specimens and fieldwork, a survey of the Compositae flora of the Arica y Parinacota Region was performed. The study shows the presence of 144 taxa of the Asteraceae family in the region of which 82.6% is native (119) and 9% is endemic to Chile (13), while 8.3% are considered as exotic species (12). The genus Senecio is the one showing the highest diversity in the region, with the presence of 27 species, including six endemic ones. The revision of the botanical material reveals the presence of five new species for the Chilean flora: Achyrocline ramosissima Britton, Gamochaeta humilis Wedd., Mniodes kunthiana (DC.) S.E. Freire et al. (=Lucilia violacea Wedd.), Pseudogynoxys cordifolia (Cass.) Cabrera, and Senecio evacoides Sch. Bip. A new species has been recently described based on our recent field surveys: Pseudognaphalium munoziae N. Bayón, C. Monti & S.E. Freire. Additionally, the correct denomination of the endemic species Stevia philippiana has been recently undertaken. The greatest diversity of species is found in the province of Parinacota, above 3,000 m asl. Compositae stand out in the region not only for its diversity but also as a crucial element of the vegetation, being a fundamental component of the precordillera and altiplano vegetation belts, known as "tolar", in which species pertaining to genera like Parastrephia, Lophopappus, Baccharis, or Senecio tend to dominate. Indeed, most of the vegetation belts described in the Parinacota province show members of the Asteraceae as dominant species. In spite of a big amount of the regional surface under formal protection, several species occur outside the protection units.
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Names of new taxa, new combinations and names at new rank are proposed for subgenera and sections in Baccharis to move towards a phylogenetic infrageneric classification of this New World genus. Two earlier segregated genera and two previously recognised sections are moved to the subgeneric rank (as B. subgen. Coridifoliae, B. subgen. Heterothalamus, B. subgen. Heterothalamulopsis, and B. subgen. Oblongifoliae). Three new combinations and/or names at new rank are proposed for the following sections: B. sect. Axillares (assigned to B. subgen. Baccharis), B. sect. Heterothalamulopsis (assigned to B. subgen. Heterothalamulopsis), and B. sect. Pluricephalae (assigned to B. subgen Coridifoliae). Four new sections are described to accommodate taxa not corresponding to any previously described section: B. sect. Andina and B. sect. Illinitae (assigned to B. subgen. Baccharis), B. sect. Bradeanae (assigned to B. subgen. Heterothalamus), and B. sect. Polifoliae assigned to B. subgen. Molina). All taxa here recognized correspond to monophyletic groups based on highly supported clades in a recent molecular phylogenetic analysis.
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Como resultado de recientes estudios florísticos en las zonas altoandinas del Perú, se dan a conocer doce nuevos registros para la flora peruana: Jalcophila boliviensis Anderb. & S. E. Freire, J. ecuadorensis M. O. Dillon & Sagást., Senecio aquilaris Cabrera, Werneria spathulata Wedd., Xenophyllum pseudodigitatum (Rockh.) V. A. Funk (Asteraceae) Draba alyssoides Humb. & Bonpl., D. loayzana Al–Shehbaz, Petroravenia friesii (O. E. Schulz) Al–Shehbaz, P. werdermannii (O. E. Schulz) Al–Shehbaz, Rorippa beckii Al–Shehbaz, Weberbauera cymosa Al–Shehbaz (Brassicaceae) y Stellaria weddellii Pedersen (Caryophyllaceae). Para cada taxón se menciona la información del tipo, su distribución geográfica actualizada y datos del hábitat. Se interpretan, además, algunos aspectos generales sobre la fitogeografía de los géneros con dichas novedades florísticas para el Perú. Palabras Clave. Andes, fitogeografía, nuevos registros, taxonomía.
Three new species of Senecio (Compositae: Senecioneae) from Central Peru are described and illustrated. Morphological characters are used in order to distinguish them from the closest species. Senecio crassiandinus Montesinos & Pino has succulent leaves with cuticle composed of translucid white glands, bright yellow disc florets with blotches of pale reddish brown in the corolla throat and linear-oblong achenes covered with trichomes, which distinguish it from the other South American species of the genus. Senecio inghamii Montesinos has oblanceolate-ovate to rhombic-quadrate leaves, short sized, with serrate-quadrate dentation, upper surface lustrous and glossy, lower surface densely covered with whitish lanuginose pubescence, capitules bear trilobulated ray florets, achenes copiously covered by fimbriate hairs which differentiates it from its closest relative, S. pensilis Greenm. Senecio roseoandinus Montesinos & R. Zárate is a unique pubescent species characterized for being a perennial rhizomatous herb, with lanuginose and arachnoid trichomes, leaves elliptic-lanceolate, thick, densely arachnoid to glabrous (lustrous, glossy, dark green), margin entire to crenate, synflorescence in racemes, capitules discoid, corolla pale yellow but turning fuchsia-pink at maturity, achenes oblong, striate, and glabrous. All three new species are distributed in the Alto Marañon river mountain slopes, at altitudes ranging from 2000 up to 4200 m a.s.l. The species are, so far, only known from their type localities in the Huánuco region.
The aquatic fern Pilularia americana A. Braun is known from several countries in South and North America. Here we provide a first report of this species for Peru, from 2 localities in the Ancash and Ayacucho regions (central Peru), which confirm its presence in the national flora.
A new species of the genus Senecio (Asteraceae, Senecioneae) belonging to S. ser. Suffruticosi subser. Caespitosi is described from the pumices of the Ticsani Volcano, central-eastern part of the Moquegua Department in southern Peru. The new species differs from S. moqueguensis by its longer indumentum, different leaf shape and size, wider involucres, narrower calycular bracts, a greater number of phyllaries per capitula, and shorter pappus. A table with the morphological comparison with other species of Senecio is provided and the conservation status of the new species is assessed.