Nordic Journal of Botany 33: 421–431, 2015

doi: 10.1111/njb.00714, ISSN 1756-1051

© 2015 The Authors. Nordic Journal of Botany © 2015 Nordic Society Oikos
Subject Editor: Bertil Ståhl. Editor-in-Chief: Torbjörn Tyler. Accepted 19 October 2014

The genus Bipinnula (Orchidaceae: Chloraeinae) in Argentina

Agustín Sanguinetti, Cristiano Roberto Buzatto and Rodrigo B. Singer

A. Sanguinetti (sango@bg.fcen.uba.ar), Laboratorio de Biología del Desarrollo de las Plantas, Depto de Biodiversidad y Biología Experimental,
Facultad de Ciencias Exactas y Naturales, Univ. de Buenos Aires. Ciudad Univ., C1428EGA, Ciudad de Buenos Aires, Argentina. AS also at:
Inst. de Biodiversidad y Biología Experimental y Aplicada. CONICET-UBA, Argentina. – C. R. Buzatto and R. B. Singer, Programa de
Pós-graduação em Botânica, Depto de Botânica, Inst. de Biociências, Univ. Federal do Rio Grande do Sul. Bento Gonçalves 9500, 91501-970,
Porto Alegre, Rio Grande do Sul, Brazil. CRB also at: Univ. Estadual de Feira de Santana, Depto de Ciências Biológicas, Av. Transnordestina s.n.,
44036–900, Feira de Santana, Bahia, Brazil.

A taxonomic synopsis of Bipinnula Comm. ex Juss. (Orchidaceae: Chloraeinae) in Argentina is presented. Three species
are recognized: B. biplumata Rchb.f., B. penicillata (Rchb.f.) Cisternas & Salazar and B. polysyka Kraenzl. A key to separate
these species is presented and their distribution in Argentina is presented. All species are described in detail and photo-
graphs of diagnostic vegetative and floral features, as well as distributional, ecological and nomenclatural comments are
included. Lectotypes for the basionyms Arethusa biplumata L.f., Chloraea arechavaletae Kraenzl. and B. polysyka Kraenzl. are
designated. All studied species are terrestrial orchids that grow in grasslands of the heavily transformed Pampas biome; two
of them are restricted to a few counties. Therefore, an assessment on their conservation status is urgently needed.

Orchidaceae Juss., with about 24 000 species, ranks together et al. 2008). Some species extend north to southern Brazil,
with Poaceae and Asteraceae among the more species-rich reaching Paraná State (Buzatto et al. 2014), while some
angiosperm families (WCSP 2014). In Argentina, this fam- species extend south, reaching the southern tip of the
ily is represented by 239 species in 74 genera (Correa 1996). Buenos Aires Province (this study).
As currently circumscribed, Chloraeinae Pfitzer comprises So far, three species have been recorded in Argentina:
about 70 species in the three genera Bipinnula Comm. B. biplumata Rchb.f., B. penicillata (Rchb.f.) Cisternas
ex Juss., Chloraea Lindl. and Gavilea Poepp. This orchid & Salazar [formerly, Geoblasta penicillata (Rchb.f.) M. N.
subtribe dwells principally in the South-American Andes, Correa] and B. polysyka Kraenzl. (Correa 1996). Vegetative
from Tierra del Fuego and insular territories of the south- features in B. biplumata and B. polysyka are inconspicuous
ern Atlantic to Peru (Correa and Sánchez 2003). However, and, in addition, both species present a short flowering
nine species differ from this general distribution pattern period (Results). As a consequence of this, these species are
and occur outside the Andes, in southern Brazil and nearby rarely collected. Consequently, information regarding their
parts of Uruguay and Argentina (Buzatto et al. 2014). In this distribution and habitat in Argentina is scarce and frag-
contribution we follow the circumscription of Chloraeinae mentary (Results). In addition, some floral and vegetative
adopted by Chemisquy and Morrone (2012) and Cisternas features are poorly known and have never been illustrated in
et al. (2012a), which is strongly supported by phylogenetic detail. Therefore, the main aim of the present contribution
analyses based on molecular characters. The various earlier is to present an updated (in nomenclatural and taxonomic
interpretations of the delimitation of Chloraeinae have been terms) synopsis of the Bipinnula species occurring in Argen-
summarized by Buzatto et al. (2014) and Chemisquy and tina together with a complete set of illustrations (photos)
Morrone (2010). of vegetative and floral features and updated data regarding
As currently accepted, Bipinnula comprises 10 their distribution, phenology and ecology.
species (Table 1; updated from Schinini et al. 2008). Five of
these species (hereafter, the occidental species) are endemic
to Chile and the remaining species (hereafter, the oriental Methods
species) dwell outside the Andes, at the eastern edge of the
continent (Fig. 109.2 of Correa and Sánchez 2003). All The following Argentinean and foreign herbaria were
occidental species present multi-florous inflorescences. Con- consulted in 2011–2014: AMES, B, BA, BAA, BAF, BBB,
versely, the oriental species are uni-florous (exceptionally, BM, CEN, F, G, GH, GOET, HAS, HB, HEPH, IBGE,
bi-florous). The distributional center of the oriental species ICN, K-L, K, LINN, LP, LPS, M, MBM, MVFA, MVM,
is Uruguay, where all species occur (Izaguirre 1973, Schinini NY, PACA, PEL, R, RB, S, SI, SP, UB, UEC, VIC and W.

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Table 1. Currently accepted Bipinnula species, sorted according to than the sterile ones, partially or totally covering the ovar-
their geographical distribution. ium. Flowers showy, sessile, with sepals and lateral petals
Occidental spp. (Andean) Oriental spp. (non-Andean) light-green coloured, with a darker green reticulated pattern
B. apinnula Gosewijn B. biplumata (L.f.) Rchb.f. and labellum differently shaped, coloured and sized than
B. fimbriata (Poepp.) I. M. Johnst. B. gibertii Rchb.f. the other perianth parts. Dorsal sepal concave, triangu-
B. plumosa Lindl. B. montana Arechav. lar to lanceolate, with entire margin and acuminate apex,
B. taltalensis I. M. Johnst. B. penicillata (Rchb.f.) sometimes crenate. Lateral sepals usually longer and nar-
Cisternas & Salazar rower than the dorsal one, pectinate-fimbriate in its apical
B. volkmanii Kraenzl. B. polysyka Kraenzl.
third (although entire in B. apinnula and B. penicillata),
with simple to trifid laciniae. Petals asymmetric, ovate,
with entire to crenate margin. Labellum sessile to unguicu-
In addition, high-definition photos of specimens deposited late and articulated at the base of the column, ecalcarate,
at BR, CORD, HBG, MPU, P, SGO, UPS, Z⫹ ZT were entire to tri-lobed, fleshy and darker than the other perianth
checked online. As a whole, 72 Bipinnula exsiccates from parts, usually with dorsal and/or lateral parts ornamented
Argentina were consulted. with warts, pubescence and/or projections. Column erect,
Data concerning distribution, habitat, phenology and slightly curved, flat in its adaxial surface, without noticeable
complementary information were obtained from the exam- column wings; in occidental species two fossae may occur
ined exsiccates as well as from personal observations during at the column base. Anther dorsal and terminal, bi-locular,
fieldwork. Fieldwork in Argentina extended from 2011 to triangular to circular in shape. Pollinarium made up by two
late 2013. Vegetative and floral features were recorded from oblong, subdivided, granular, yellow pollinia but devoid of
living specimens with the help of a digital camera. Data from viscidium. Ovarium obconic to clavate. Capsule ellipsoid to
pressed specimens and from their respective protologues and pyriform, dehiscent through two longitudinal slits along the
relevant literature (Correa 1968a, 1968b, Izaguirre 1973, dorsal carpelar suture.
Correa and Sánchez 2003, Buzatto et al. 2014) was also
taken into account. Most of our own collections consist of Historical treatments
inflorescences and a few leaves such that collection impact The name Bipinnula was coined by Commerson and
was minimized and orchid individuals were preserved. A published by Jussieau (1789, p. 65), in order to set apart
minimum of whole specimens were removed in order to species formerly placed in Arethusa L. (1753, p. 950) whose
illustrate subterranean vegetative characters or vegetative lateral sepals have pectinated-ciliated apices, among other less
features absent during anthesis. Vouchers were deposited at important features. Phylogenetic studies based on molecular
SI and BA. Only one voucher is cited per administrative unit characters have clearly pointed out that the species currently
(county, department, etc.). To ensure their conservation, the placed in Bipinnula (Table 1) form a well-supported clade
precise collection data are not given here. This information which, in turn, is the sister-group of Gavilea. Both clades are
is, however, available for bona fide researchers, on request. inserted within the species-groups of Chloraea, rendering the
Specific orchid morphological terms follow Dressler (1993). latter genus paraphyletic (Chemisquy and Morrone 2012,
More specific Chloraeinae terminology follow Sanguinetti Cisternas et al. 2012a).
et al. (2012). Kraenzlin (1903, p. 18) first recognized two well-
differentiated species assemblages within Bipinnula by
Bipinnula Comm. ex Jussieu (1789, p. 65) placing the oriental species in the “Uniflorae” group, and
the occidental species in the “Spicatae”, accompanying
Type: Bipinnula commersonii Lindl. (1827, p. 52). nom. this segregation with the short phrase “Species spicatae terrae
illeg. – Bipinnula biplumata (L.f.) Rchb.f. (1883, p. 62); Chilensis, uniflorae regionis Bonariensis et Brasiliae meridionali
lectotypified by Kraenzlin (1903, p. 18). sincolae”. Nieuwenhuizen (1993, p. 13) published the most
recently described species in the genus and then proposed
Taxonomic synonyms: Geoblasta Barb. Rodr. (1891, dividing it into three sections: B. sect. Bipinnula (contain-
p. 132). Type: Geoblasta teixeirana Barb. Rodr. (1891, ing the type species B. biplumata and all oriental species)
p. 133). – Jouyella Szlach., in Szlachetko and Margonska (Table 1), B. sect. Trilobatae Gosewijn and B. sect. Multiflo-
(2001, p. 124). Type: Jouyella fimbriata (Poepp.) Szlach. & rae Gosewijn. Species in B. sect. Bipinnula are non-andean
Marg. (2001, p. 125). – Chloraea fimbriata Poepp. (1833, and easily diagnosed on the basis of their uni-florous inflo-
p. 15). rescences, entire to obscurely tri-lobed, sometimes fleshy
lip, covered by papillae or pubescence and fimbriate lateral
Description sepals, often with bifid laciniae. Species in B. sect.
Terrestrial herbs; geophytes. Roots fascicled, cylindrical Trilobatae and Multiflorae (two and three species, respec-
and tuberose. Leaves basal, fascicled to rosulate, elliptic to tively) are exclusively Andean (Chilean). Species within B.
linear-lanceolate, with entire margin and obtuse to acute sect. Trilobatae have pauciflorous spikes (2–10 flowers),
apex; generally withering or absent during anthesis in clearly 3-lobed labellum with erect lateral lobes and mid-
the oriental species. Inflorescence a multi-florous spike in lobe covered by warty or papilose projections, apices of
the occidental species and uni-florous (exceptionally bi- lateral sepals without laciniae or shortly pectinate and base
florous) in the oriental ones. Scape erect and covered by of the column with two shallow fossae. Species within
some acute, sterile bracts. Floral bracts acuminate and larger B. sect. Multiflorae are diagnosed by their multi-florous

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(⬎ 10 flowers), wide entire labellum which often has fleshy to this species based on morphological standards. Using the
projections, fimbriate lateral sepals with long laciniae and same criterion that prompted the nomenclatural transfer of
column base with two fossae projecting inside the ovarium. G. penicillata to Bipinnula, the name B. apinnula should
Barbosa Rodrigues (1891, p. 133) proposed the genus be kept as the correct one in order to avoid paraphyly of
Geoblasta, when publishing G. teixeirana, based on speci- Bipinnula. Therefore, the following treatment is proposed:
mens collected in Curitiba, Paraná, Brazil. Much later,
Hoehne (1940) correctly synonymized G. teixerana with Bipinnula apinnula Gosewijn, in Nieuwenhuizen
Chloraea penicillata Rchb.f. (a name with nomenclatural (1993, p. 11)
priority). However, Hoehne (1940) was emphatic in that Based on the same type: Chloraea apinnula (Gosewijn)
this taxon could be separated from Chloraea owing to a Szlach., in Szlachetko and Margonska (2001, p. 20). –
distinct set of morphological features, therefore suggesting Ulantha apinnula (Gosewijn) Szlach., in Szlachetko and
the revalidation of Geoblasta (Hoehne 1940). Considering Tukallo (2008, p. 115).
all these precedents, Correa (1968a, p. 71) proposed the
rehabilitation of Geoblasta, with G. penicillata (Barb. Rodr.) Type: Chile. Talca, Cerro Peine, 1800 m a.s.l., 6 Dec 1989,
Hoehne ex M. N. Correa as its only species. However, it Gosewijn s.n. (holotype: SGO-1352 [image!], isotype SGO-
is important to highlight that Correa (1968a, p. 71) was 1353 [image!]; paratypes CONC-117825, SGO-110620, K).
the first to notice the morphological resemblances between
Geoblasta and some uni-florous species of Bipinnula.
Key to the Argentinian species of Bipinnula
Cisternas et al. (2012b, p. 10), on account phylogenetic
analyses based on molecular characters (Chemisquy and 1. Leaves linear, erect; labellum apex velutinous, entire ........
Morrone 2012, Cisternas et al. 2012a) and morphological ................................................................ B. biplumata
features (Correa 1968a) transferred G. penicillata to Bipin- – Leaves circular-ovate, decumbent or prostrate; labellum
nula, thus creating the new combination Bipinnula penicillata apex ornamented with non-velutinous warts or projec-
(Rchb.f.) Cisternas & Salazar. By making this, they obtained a tions, not entire …………………………………. 2
consistent delimitation of Bipinnula (Cisternas et al. 2012b). 2. Apex of lateral sepals entire; labellum sessile, with long,
Keeping Geoblasta as an accepted genus would have rendered claviform, dark and yellow projections …….. B. penicillata
Bipinnula paraphyletic (Cisternas et al. 2012b). – Apex of lateral sepals fimbriate; labellum articulated,
Szlachetko and Margonska (2001), based on observa- unguiculate, with short, warty, dark projections ……
tion of morphological features on herbarium specimens, …………………….................................. B. polysyka
suggested that Bipinnula was not monophyletic and pro-
posed a new genus (Jouyella Szlach.) for the species in the Bipinnula biplumata (L.f.) Rchb.f. (1883, p. 62)
B. sect. Multiflorae. However, ongoing phylogenetic analyses (Fig. 1)
(Cisternas et al. 2012a, 2012b) strongly suggest that
Bipinnula species of the sections Bipinnula and Trilobatae are Basionym: Arethusa biplumata L.f., in Linnaeus (1782,
nested among the species of B. sect. Multiflorae ( ⫽ Jouyella). p. 405).
Based on this evidence, Buzatto et al. (2014) synonymized
Jouyella with Bipinnula. Based on the same type: Bipinnula bonariensis Spreng.
(1826, p. 745), nom. illeg. – Bipinnula commersonii Lindl.
Notes on extra-Argentinean Bipinnula (1827, p. 52), nom. illeg.
In the following, some issues regarding extra-Argentinean
species are commented in order to clarify the generic cir- Type: “in Freto Magellanico”, 1767, Commerson s.n.
cumscription (Table 1). This, however, does not pretend to (lectotype designated here: LINN-HS-1394.5 [image!],
replace a thorough generic revision. Bipinnula ctenopetala isolectotypes: MPU-017561 [image!], MPU-017562
Schltr. has recently been considered a dubious name and a [image!], UPS-Thunb-21419 [image!], P-372103 [image!],
probable synonym of B. montana Arechav. (Buzatto et al. P-372104 [image!], P-372105 [image!], P-372108
2014). Based on morphological (mostly floral) characters, [image!]).
Szlachetko transferred B. apinnula to the genus Chloraea
[as Chloraea apinnula (Gosewijn) Szlach; Szlachetko 2001] Description
and later to the genus Ulantha [as U. apinnula (Gosewijn) Herb, 15–50 cm high (Fig. 1A). Roots numerous, fascicled,
Szlach; Szlachetko and Tukallo 2008]. Therefore, Bipinnula tuberous, up to 8 cm long, with conical apex and annu-
apinnula was placed as a synonym of Ulantha apinnula lar thickenings at the base. Leaves fascicled, with sheating
in the “Catálogo de Plantas Vasculares del Cono Sur” base, numerous, erect, linear, 14–20 cm long and 1.0–
(Schinini et al. 2008). Furthermore, in the ‘World checklist 1.5 cm wide, with apiculate apex, persistent or wither-
of selected plant families’ (WCSP 2014) Chloraea apinnula ing during the anthesis (Fig. 1C). Scape erect, uni-florous
is the currently accepted name, this being reflected in several (exceptionally bi-florous), with short, invaginant bracts
major sources of taxonomic information such as Tropicos slightly longer than the inernods. Flower violaceous–
(2014) and ‘The plant list’ (2013). However, phylogenetic green, held by an invaginant bract with acute apex
studies have inequivocally shown that this taxon is well- (Fig. 1B). Dorsal sepal triangular to lanceolate, 25–
nested within Bipinnula (Cisternas et al. 2012a), thus sup- 30 mm long and 8–11 mm wide, concave, with entire
porting the position that Nieuwenhuizen (1993) assigned margin and acuminate apex. Lateral sepals linear, recurved

423
Figure 1. Bipinnula biplumata. (A) view of whole plant, inset: detail of bi-florous individual, (B) flower, (C) vegetative features, (D) flower
in lateral view, lateral petal and lateral sepal removed, (E) pollinarium, (F) adaxial view of labellum, (G) ovarium and column, (H) detail of
the apex of lateral sepal, (I) fruit, Scales: (A)–(C) and (I) ⫽ 1 cm, (D) ⫽ 5 mm, (E)–(H) ⫽ 1 mm. (A) and (I) from Sanguinetti 63 (BA), (B)
and (D)–(H) from Sanguinetti 119 (SI).

and involute, slightly curved, 30–45 mm long and 3–5 mm green (Fig. 1D–F). Column erect, slightly curved, 17 mm
wide in basal part, narrowing towards the apex and with long and 4 mm wide. Stigmatic surface ovate, distal, 4 mm
pectinate apical part, with cylindric, somewhat curved, long. Rostellum light green, thick and glandular, transver-
sometimes bifid, laciniae that are 2–4 mm long (Fig. 1H). sal (Fig. 1G). Anther incumbent, triangular. Pollinarium
Petals ovate, asymmetric, 20–25 mm long and 14–18 mm 7 mm long and 4 mm wide (Fig. 1E). Ovarium obconic,
wide, with entire margin and acute apex. Labellum shortly 10 mm long (Fig. 1D). Fruit an obovoid capsule, 20 mm
unguiculated, articulated at the base of the column, long and 13 mm wide (Fig. 1I).
obscurely 3-lobed, 14–17 mm long and 10–14 mm wide;
lateral lobes involute, white, with transversal, contrasting Phenology
and regularly-spaced black warts; the median lobe pulvi- This species flowers from November to December and fruc-
nate, with dark, muricate surface; apex obtuse, recurved tifies from December to January after which they become
and revolute, with velvety surface coloured in dark olive quiescent. Plants sprout again in February–March.

424
Distribution Additional material examined
Bipinnula biplumata extends to the north, with discontinu- Argentina. Buenos Aires: Olavarría, 20 Nov 1901, Spegazzini
ities, to the Brazilian State of Paraná (Buzatto et al. 2014), s.n. (LPS-1989!). Azul, 22 Nov 2013, A. Sanguinetti 119
Brazil and Uruguay. To the south, this species reaches the (SI!). Santa Fé: Iriondo, 1886/1887, Berndt s.n. apud Kurtz
central hills of the Buenos Aires Province. The few known 5156 (CORD [image!]).
Argentinean specimens come from the sierras of Olavarría
and Azul, with a single known specimen from low eleva- Iconography
tions in Santa Fé Province (Fig. 4). Lamarck (1797, Pl. 729, Fig. 4); Smith (1789, XXII; based
on the lectotype); Cogniaux (1893, Pl. 21, Fig. 1; reproduced
Habitat and ecology by Correa 1968b, p. 590, Correa et al. 2009, p. 351, Herter
Bipinnula biplumata is associated with rocky outcrops in hilly 1939, p. 250); Kraenzlin (1903, Pl. I, Fig. D); Izaguirre
landscapes, just like B. penicillata. However, B. biplumata (1973, p. 265, 1984, p. 414); Buzatto et al. (2014, p. 5).
grows in deeper soils and may be surrounded by denser and
taller vegetation than the latter species. In Buenos Aires it is Bipinnula penicillata (Rchb.f.) Cisternas & Salazar in
found either in hilly grasslands dominated by Piptochaetium Cisternas et al. (2012a, p. 10). (Fig. 2)
spp. and Nassella spp., or in shrubby communities with pre-
dominance of Baccharis tandilensis Speg. and Eupatorium Basionym: Chloraea penicillata Rchb.f. (1878, p. 51).
buniifolium Hook. ex Arn. During anthesis, B. biplumata
may or not present fresh leaves, depending on light condi- Based on the same type: Geoblasta penicillata (Rchb.f.)
tions and water availability. Hoehne ex M. N. Correa (1968a, p. 71).

Notes Type: “Orange Harbour. Tierra del Fuego”, 1838, Wilkes

Bipinnula biplumata is the type species of the genus. This Expedition s.n. (holotype: AMES-106737!).
species was cited for Argentina more than a hundred years
ago (Hicken 1910, p. 77). However, B. biplumata is repre- Taxonomic synonyms: Geoblasta teixeirana Barb. Rodr.
sented by very few herbarium specimens in local and foreign (1891, p. 133); based on the same type: Chloraea teixeirana
herbaria. For instance, the four most important herbaria (Barb. Rodr.) Cogn. (1893, p. 107). Type: Brazil. Paraná:
from Buenos Aires Province (SI, LP, BAA and BA) had – to Curitiba, Barbosa Rodrigues s.n. (destroyed, fide Sprunger
date – a single collection made by Spegazzini, in 1901, in et al. 1996, Buzatto et al. 2011, 2013). Lectotype designated
Buenos Aires Province. by Buzatto et al. 2014: Barbosa Rodrigues's original illustra-
Spegazzini (1916, p. 140) was the first researcher that tion deposited at the Biblioteca de Barbosa Rodrigues, Jar-
gave a precise location for this species in Argentina. Unfor- din Botanico de Rio de Janeiro, Iconographie des Orchidées
tunately, he referred to this species as “B. biplumosa Reich.”, du Brésil. Vol. 1, Pl. 34, holotype cited as Pl. 865 [inedit],
thus causing confusion with the Chilean, multi-florous, Barbosa Rodrigues (1891), reproduced by Sprunger et al.
B. biplumosa Lindl. This error was later corrected by (1996. Vol. 1, p. 87).
Hauman and Vanderveken (1917, p. 309). Correa (1968b,
p. 589) refers to the Spegazzini specimens in her ‘Flora – Chloraea arechavaletae Kraenzl. (1888, p. 316); based on
de Buenos Aires’, defining the habitat of the species as the same type: Asarca arechavaletae (Kraenzl.) Kuntze (1898,
follows: “vive en campos abiertos” (living in open grass- p. 298) and Geoblasta arechavaletae (Kraenzl.) Szlach. &
lands). After almost a hundred years, Orfila (2000, p. 5) Marg. (2001, p. 125). Type: Uruguay. Montevideo: “entre
again collected this species at the sierras of Azul (Buenos las piedras de Independencia”, Nov 1874, Arechavaleta 2615
Aires Province). His publication (Orfila 2000) was scarcely (lectotype designated here: HBG-500263 top left specimen
divulgated. However, Orfila (2000) provided photographic [image!], isolectotypes rest of HBG-500263 specimens,
illustrations and more precise habitat references. The MVFA!, probably HBG-500925 and HBG-500926 [not
known distribution of this taxon in Argentina is restricted seen]).
and noteworthy. So far, B. biplumata has been collected
at the northeastern extreme of the Tandilia mountain – Chloraea bergii Hieron. (1879, p. 380); based on the same
system. This species has never been found in other parts type: Geoblasta bergii (Hieron.) Szlach. & Marg. (2001,
of this mountain system that, as a whole, is very well- p. 125). Type: Argentina. Provincia de Buenos Aires:
explored from the floristic point of view since Spegazzini Carmen de Patagones, 1874, Berg s.n. (holotype: CORD-
(1916) to the present (Frangi 1975). More remarkable is 2208 [image!]).
the only known specimen (dated 1886–1887) from Santa
Fé Province. This pressed voucher comes from the north- Description
ern limit of the Rolling Pampas phytogeographic district Herb, 10–20 cm high (Fig. 2A). Roots fascicled and dimor-
(sensu Soriano 1991), with deep loessic soils and devoid phic; 3–5 tuberose and thick, 4 to 7 cm long and 2.5 cm
of mountains or rocky outcrops. The natural grasslands wide, with annular thickenings at the base; 3–7 slender and
of this region have mostly become crop lands (especially cylindric, up to 10 cm long and 0.3 cm wide, with conic apex
soybean plantations) (Baldi and Paruelo 2008). Therefore, and devoid of basal annular thickenings. Leaves scarce, rosu-
is quite unlikely that there are relictual populations of late, prostrate, elliptic to ovate, 3–7 cm long and 1.5–3.0 cm
B. biplumata in that region. wide, with obtuse to rounded apex, sometimes mucronulate,

425
Figure 2. Bipinnula penicillata. (A) view of whole plant, (B) vegetative features, (C) column, (D) fully opened flower, (E) from left to right,
labellum in adaxial, lateral and abaxial views, respectively, (F) closed flower, (G) fruit, (H) pollinarium glued to a pin by means of rostellar
secretion. Scales: (A)–(G) ⫽ 1 cm, (H) ⫽ 1 mm. (A) from Sanguinetti 103 (SI); (C)–(F) and (H) from Sanguinetti 102 (SI).

normally withering during anthesis (Fig. 2B). Scape erect, Phenology

uni-florous (exceptionally bi-florous, P-372174), covered by Flowers from October to early December. Fructifies from
4–6 apiculate, invaginant bracts up to 4 cm long. Floral bract November to December after which they become quiescent.
bigger than the scape bracts and totally covering the ovarium. Sprouts again in March.
Flower whitish–green, with darker, fleshy, shiny and insecti-
form labellum. The remaining perianth parts membranous, Distribution
semi-transparent and with a contrasting dark–green reticu- Among the oriental species, this is the one with the widest
lated pattern (Fig. 2A, 2D). Dorsal sepal lanceolate, 25–50 distribution. The known northern distributional limit is the
mm long and 10–15 mm wide, entire, acute to acuminate. Brazilian state of Paraná and its southernmost record is that
Lateral sepals lanceolate, 25–45 mm long and 12 mm wide, of the holotype of Chloraea bergii, which was collected in
entire, acuminate (Fig. 2A, 2F). Petals ovate to lanceolate, Carmen de Patagones, Buenos Aires Province, the northern
slightly asymmetric, 25–45 mm long and 13–15 mm wide, limit of the Patagonia (Buzatto et al. 2014, this study). In
with entire margin, sometimes slightly serrated at the api- Argentina, this species has been collected in the eastern part of
cal part, and acute apex. Labellum sessile, elliptic to slightly the Entre Ríos Province and, in the Buenos Aires Province, from
rhombic or obtrullate, in its basal and median parts green, the central Sierras towards the south (Fig. 4). Correa (1968a,
fleshy and shiny, 20–22 mm long and 12–14 mm wide. The p. 71) explains that the Tierra del Fuego citation appear-
labellar basal part is covered by villose whitish green pubes- ing at the label of the type specimen of Chloraea penicillata
cence. Two longitudinal, shiny, fleshy and smooth carenae is an error and that this material may actually have been col-
stand out from the rest of the smooth labellar surface. Label- lected at the estuary of Río Negro (near Carmen de Patagones)
lar apex covered by short clavate to capitate brownish projec- where the Wilkes Expedition disembarked. Vervoorst (1967,
tions; margin of basal part entire and margin of distal part p. 104) states that Chloraea bergii ( ⫽ B. penicillata) is present
revolute and provided with several simple, cylindrical, yel- in the same plant communities as B. polysyka (below), but this
low projections that are ca 3–6 mm long (Fig. 2E). Column may also be an error. Whereas Chloraea membranacea Lindl.
incurved, 13 mm long and 4 mm wide. Stigmatic surface (Chloraeinae) is indeed quite common in these plant commu-
roundish, wider than the column, ca 6 mm long. Rostel- nities, B. penicillata has never been collected in these localities.
lum glandular, prominent and transverse (Fig. 2C). Anther
circular. Pollinarium 5 mm long and 4 mm wide (Fig. 2H). Habitat and ecology
Ovarium obconic, 10 mm long. Fruit an obovoid capsule Usually common in prairies on hills (Spegazzini 1901,
20–30 mm long and 14 mm wide (Fig. 2G). p. 47), B. penicillata is generally associated with shallow

426
soils in rocky environments where survival to dry periods C. bergii and C. teixerana in the synonymy of C. penicillata.
is granted by means of its tuberous roots. There, it can be The history of the rehabilitation of Geoblasta and its further
found together with other drought-tolerant plants such as transference to Bipinnula is described above (see comments
Wigginsia tephracantha (Link & Otto) D. M. Porter, Oxalis after the generic description).
articulata Savigny, Gomphrena perennis L., Vernonia flexuosa When Correa (1968a) transferred Chloraea penicil-
Sims, Gamochaeta filaginea (DC.) Cabrera, Dichondra repens lata Rchb.f., creating the new combination Geoblasta
var. sericea (Sw.) Choisy and Aristida spegazzini Arechav. penicillata, the spelling of the basionym was changed to
(Frangi 1975, pers. obs.). In the Entre Ríos Province this “pennicillata” without further justifications. Ever since
plant has been recorded in grasslands and Baccharis spp. this spelling was kept in numerous important taxonomic
maquis as well, in relatively high, well-drained areas (Ciotek works (Correa 1968b, 1996, Correa and Sánchez 2003,
pers. comm.). This species is pollinated by males of Schinini et al. 2008, Correa et al. 2009, Cisternas et al.
Campsomeris bistrimacula (Lepeletier) that attempt copu- 2012b). In one of the latest works where Correa partici-
lation with the insectiform labellum (Ciotek et al. 2006). pated (Correa et al. 2009) the etymology of “pennicillata”
During the sunnier and hottest hours, sepals and lateral is explained as derived from “penna” (feather) and the
petals of fresh flowers incurve, totally exposing the column suffixes “-cillus” and “-atus” (indicating diminutness and
and labellum Remarkably, the same floral parts have the abil- likeness, respectively). This may be a hint that the change
ity to turn back and hide/protect the column and labellum was made on purpose believing there was an orthographic
after late afternoon and/or under adverse weather conditions error, thus attending the Art. 73/Note 4 of the Edinburgh
(Fig. 2F). Code (Lanjouw et al. 1966), which corresponds to Art.
60.3 in the present Code (McNeill et al. 2012). In any
Notes case, whether this change was intentional or was lapsus
Bipinnula penicillata is the species with the most numer- calami is irrelevant and B. penicillata should be retained.
ous herbarium records from Argentina. In Buenos Aires Reichenbach filius published the name in that manner
Province, most records come from the Partidos (counties) in its protologue and in subsequent works (Reichenbach
of Tornquinst (23%), Tandil (21%), Balcarce (17%) and 1878–1883: 28, 61, pl. 229), not describing the etymology
Saavedra (17%). All these localities are placed within the though; where penicillata is a feminine and singular com-
Tandilia and Ventania mountain systems. When in bloom, pound adjective formed by the noun “penicillus” (brush)
this species is quickly and easily found, because it is relatively and the adjectival suffix “-atus” (above). Moreover, this
frequent in some specific localities. When devoid of flow- author published a drawing (Reichenbach 1878–1883:
ers, however, the plant is remarkably inconspicuous for the pl. 229) based on the holotype where a resemblance to a
non-trained eye. Yet, this species tends to occur in similar paintbrush is clearly noticed.
microsites at the outcrops, mostly in places with sparse veg-
etation. These mountain systems are very well explored from Additional material examined
the floristic point of view, a fact that may have contributed Argentina. Buenos Aires: Azul, 7 Nov 2012, A. Sanguinetti
to the relative richness of records of this species in Argentin- 62 (BA). Balcarce, 31 Oct 1959, E. Grondona 7056 (BAA).
ean herbaria. Conversely, adjacent regions (such as Coronel Coronel Dorrego, 4 Nov 2008, F. Biganzoli and C. Larsen
Dorrego, General Alvarado, Tres Arroyos) have not been 1987 (SI). Coronel Pringles, 28 Nov 1932, Stegmann s.n.
explored so thoroughly and, consequently, few specimens (SI-25685). General Alvarado, 3 Dec 1946, A. L. Cabrera
are known from there. The same factors may apply for the 10378 (SI). General Pueyrredon, 11 Nov 1962, O. Boelcke
Carmen de Patagones region (southern Buenos Aires Prov- et al. 9340 (BAA). Olavarria, 1909, C. Spegazzini s.n. (LPS-
ince) and for the Entre Rios Province as well. Because this 2085). Patagones, 1874, C. Berg s.n. (CORD [image]).
species is inconspicuous and seems to have a low abundance Saavedra, 10 Nov 1932, L. R. Parodi 10310 (BAA). Tandil,
outside rocky environments, it is possible that its distribu- 3 Nov 1928, A. Burkart 2773 (BAA). Tornquist, Oct 1907,
tion in non-montane environments is actually underrepre- C. M. Hicken s.n. (SI-40232).
sented.
This species was originally referred to the genus Chloraea References of non-examined material
(see synonymy). Kraenzlin (1903, p. 44) published an arti- Argentina. Buenos Aires: Bahía Blanca, 1884, M. G.
ficial key to set apart four species that are today considered Mansel s.n. (BM-95739, cited in Correa 1968a). Tres
heterotypic synonyms (C. arechavaletae, C. bergii, C. pen- Arroyos, 15 Nov 1987, C. B. Villamil and M. G. Canizzaga
icillata, C. teixerana), based on geographical distribution 5424 (BBB, C.B. Villamil, pers. comm.). Entre Ríos: Colón,
and labellar characters. Remarkably, each of these different cited in Ciotek et al. (2006, p. 366). San Salvador, R. Báez
names agrees with different collection localities (Montevi- s.n. in Hauman (1920, p. 99).
deo, Buenos Aires, Tierra del Fuego and Paraná). Hauman
(1920, p. 44) was the first to synonymize Chloraea bergii Iconography
under C. arechavaletae, recognizing that labellar features Reichenbach (1883, p. 229, Fig. 1; based on the holotype of
are variable, even within the same region. Indeed, Hauman C. penicillata); Cogniaux (1893, Table 20, Fig. 2 reproduced
(1920) emphasized the already well-known floristic affinities in Correa et al. 2009, p. 14); Kraenzlin (1903, Table IV,
between Uruguay, Entre Rios and Buenos Aires argue against Fig. A–C); Herter (1939, p. 250); Correa [1968a, p. 72,
Kraenzlin's (1903) geographic arguments. Later, using simi- 1968b, p. 592, reproduced in Cabrera and Zardini (1993,
lar criteria, Hoehne (1940, p. 196) placed C. arechavaletae, p. 216)]; Izaguirre (1984, p. 411); Buzatto et al. (2014, p. 13).

427
Bipinnula polysyka Kraenzl. (1888, p. 317) (Fig. 3) at low altitudes, near water sources (Izaguirre 1973, p. 262).
In Argentina, this species has been found in plain grass-
Type: Uruguay. Montevideo: Cerro de Montevideo, ladera lands and in low altitudes, in association with Nassella char-
oeste, Nov 1875. Arechavaleta 2627 (lectotype designated ruana (Arechav.) M. E. Barkworth and other grasses such as
here: ZT-14858 third specimen from left to right [image!], Aristida murina Cav., Briza brizoides (Lam.) Kuntze, Briza
isolectotypes rest of ZT-14858 specimens, MVFA!). subaristata Lam., Bothriochloa laguroides (DC.) Herter,
Danthonia montevidensis Hack. & Arechav., Lolium
Description multiflorum Lam. snd Piptochaetium montevidense (Spreng.)
Herb 12–25 cm high (Fig. 3A). Roots fascicled and dimor- Parodi (Vervoorst 1967; pers. obs.). These areas are normally
phic; 2–5 tuberose-globose 2–4 cm long and 1.0–1.5 cm used to graze livestock and the soil is very argilose, hard
wide, with annular thickenings at the base; 2–5 cylindrical (when dry) and poorly drained.
to 5 cm long and 2 mm wide, with conical apex and devoid
of annular thickenings. Leaves rosulate, numerous, decum- Notes
bent, ovate, 2–5 cm long and 1.0–2.5 cm wide, obtuse to Bipinnula polysyka was the last species in the genus to be
acute at apex , withering during anthesis (Fig. 3B). Scape found in Argentina (Correa 1959, p. 180). Remarkably, the
erect, uni-florous (exceptionally bi-florous), covered by api- first Argentinian voucher (LP 54527) was collected in 1944
culate invaginant bracts about 3–5 cm long. Floral bract at a very urban, accessible place, as it was Barrio Elizalde,
slightly bigger than the scape bracts, sometimes covering in the outskirts of La Plata City. This fact highlights the
the ovarium, sometimes not. Flower terminal, with black, inconspicuosness of this species. Veervoorst (1967), as part
shiny labellum contrasting with the other, mostly greenish of his phytogeographic studies of the Rio Salado basin, gave
floral parts (Fig. 3C). Dorsal sepal ovate 19–22 mm long the first accurate information regarding the habitat of this
and 9–12 mm wide, concave and recurved, with entire mar- orchid. In fact, the association of B. polysyka with commu-
gin and acute and crenate to condilomatose apex. Lateral nities of N. charruana pointed out by Veervoorst (1967)
sepals linear, recurved and involute, 23–30 mm long and made it possible to find the plants recorded and illustrated
5–7 mm wide in the basal part, narrowing towards the apex, here. In the early 1990's, a remnant population was found
with margins of apical part fimbriate, bearing irregular, tor- at the locality of Esteban Echeverría (Alejandro Taborda,
tuous, simple to trifid laciniae that are 2–4 mm long (Fig. pers. comm.; voucher BAA-24936). This population was
3E). Petals ovate, slightly asymmetric 16–20 mm long and lost as a consequence of urban development and no further
9–12 mm wide, margin entire and apex obtuse, crenate to records are known from this locality. Yet, it is possible that
condilomatose, with marginal protuberances extending B. polysyka also grows in Nassella charruana grasslands at the
towards the medial region. Labellum shortly unguiculated, Entre Rios Province (e.g. Gualeguaychú), considering the
articulated at the base of the column, 16 mm long and 7 proximity with localities in Uruguay where this species has
mm wide. Labellar disc arched, somewhat panduriform, at already been collected (Izaguirre 1973, SNAP 2014).
base yellowish–green, umbonate and puberulent, darkening Kraenzlin comments in the protologue as well as on the
towards the apex and aquiring a foveolate surface and a shiny lectotype label that B. polysyka is very close to B. giberti, a
black colour; lateral margins reduced to a short condiloma- species that – to date – has only been collected in Uruguay
tose wing bearing clavate, flat to geniculate projections. The and southernmost Brazil (Izaguirre 1973, Buzatto et al.
labellar apex crowned by a dense fascicle of short black pro- 2014). Yet, Kraenzlin emphasizes that these species can be
jections (Fig. 3D). Column arched, 8 mm long and 4 mm separated on the basis of the crenate apex of the dorsal sepal
wide (Fig. 3D, 3F). Stigmatic surface eliptic, ca 4 mm long. in B. polysyka, vs the non-crenate condition in B. giberti.
Rostellum glandular and transversal (Fig. 3F). Anther ovate. These characters are well illustrated by Cogniaux (1893, Pl.
Pollinarium ca 4 mm long and 3 mm wide (Fig. 3G). Ovar- 21) which, in turn, based his illustrations in the specimens
ium obconic, 6–8 mm long. Fruit as an obovoid capsule, 13 cited in the respective protologues. Williams (1938, p. 138)
mm long and 5 mm wide (Fig. 3H). studied pressed specimens from Uruguay and reaffirms that
these species can be separated on the basis of the dorsal sepal
Phenology
features, but establishes that some specimens of B. polysyka
Flowers from late October to late November and fructifies
present a non-crenate dorsal sepal. Finally, Williams (1938)
in November and early December after which it becomes
suggests that the study of more specimens may ultimately
quiescent. Plants sprout again in March.
lead to the conclusion that B. polysyka is just a variety of B.
giberti. After having studied living specimens of both spe-
Distribution
cies (Buzatto et al. 2014, this study) we can affirm that B.
This species has been collected in Uruguay, from the county
polysyka can be separated from B. giberti on the basis of its
of Soriano to the margins of the Río de la Plata (Izaguirre
larger perianth parts, its whitish base of petals and sepals and
1973, SNAP 2014). In Argentina, this species has a nar-
its shiny black labellum (vs the opaque, greenish–grey label-
row distribution in the Buenos Aires Province, from Ezeiza
lum of B. giberti).
to Pipinas, and from Río Samborombón to Río de la Plata
Bipinnula polysyka is easily recognizable among
(Vervoorst 1967; this study) (Fig. 4).
Argentinean Bipinnula species because of its massive, shiny
Habitat and ecology black labellum and remaining whitish floral parts. The
In Uruguay this species has been found growing in exposed, leaves of B. polysyka are easily confused with those of
dry places such as mountain summits as well as in locations B. penicillata. However, the leaves of B. polysyka can be

428
Figure 3. Bipinnula polysyka. (A) view of whole plant, inset: detail of bi-florous individual, (B) vegetative features, (C) flower, (D) lateral
view of flower with lateral petal and sepals (dorsal and lateral) removed and adaxial view of labellum, (E) detail of the apex of lateral sepal,
(F) frontal view of column, (G) pollinarium, (H) fruit. Scales: (A)–(D) and (H) ⫽ 1 cm, (E)–(G) ⫽ 1 mm. (A) from Sanguinetti 60/64 (SI);
(C)–(G) from Sanguinetti 60 (SI).

longer (Description) and slightly less fleshy. Just like Philibert Commerson and Jeanne Baret. The main collection
B. penicillata, this species may lose its leaves when flowering. of Commerson is held at P where many duplicates were distrib-
uted by Jussieau to different European herbaria – particularly
Additional material examined P, G, LINN, UPS – (Stafleau and Cowan 1976). According to
Argentina. Buenos Aires: Ezeiza, 21 Nov 1993, A. Castillo Art. 9.3 (McNeill et al. 2012) all these Commerson’s dupli-
et al. s.n. (BAA-24936). La Plata, 9 Nov 1944, A. L. Cabrera cates of Arethusa biplumata are treated as original material. We
8424 (LP). Magdalena, 15 Nov 2013, A. Sanguinetti 110 designate the specimen at LINN-HS as the lectotype because
(SI). Punta Indio, 17 Nov 2013, A. Sanguinetti 112 (SI). this belonged to the collection which Linnaeus filius mostly
worked with (Stafleau and Cowan 1976).
Iconography
Kraenzlin described both Bipinnula polysyka and Chlo-
Cogniaux (1893, Pl. 21, Fig. 3; based on original material,
raea arechavaletae ( ⫽ B. penicillata) based on specimens sent
modified in Herter 1939, p. 251, Fig. 1001, and Correa
by the Uruguayan botanist Arechavaleta (Kraenzlin 1888),
1968b, p. 590, Fig. 124A–C); Kraenzlin (1903, Pl. I, Fig.
without mentioning the herbarium of reposition (Christen-
E); Correa (1959, p. 181); Izaguirre (1973, p. 273, 1984, p.
son 1994). The primary set of Kraenzlin's types was sold to
414); Szlachetko and Margonska (2001, p. 124, Fig. 1B, 2).
Berlin (B) in 1907 and most likely lost during Allied air raids
Typifications in World War II (Stafleau and Cowan 1976, Poppendieck
2001). In agreement with this scenario, two of the authors
Linnaeus filius described Arethusa biplumata based on (CRB and RBS) thoroughly revised the orchid collection
material collected in 1767 on the Rio de la Plata basin by at B in late 2012, without finding any of these types. For-

429
 cessfully located type specimens of Kraenzlin, Schlechter and
Mansfeld which were also thought to be lost at B during the
WWII. They speculate that these specimens reached Z ⫹
ZT due to the active collaboration between these botanists
and Hans Shinz, based in Zürich. As for C. arechavaletae,
parts of the original material of B. polysyka corresponding to
isolectotypes were kept by Arechavaleta in MVFA.

Conclusions

The vast majority of the extra-Argentinean examined her-

barium exsiccates not collected by the present authors
belong to B. penicillata (87%), followed by B. polysyka (8%)
and B. biplumata (5%). This numerical predominance of
B. penicillata over the other two species may be – to a great
extent – explained by its wider distribution (Fig. 4). Other
factors that may have contributed to the rareness of her-
barium collections of B. biplumata and B. polysyka may be
their vegetative inconspicuousness and their short flower-
ing period. In Argentina, all these species dwell in Pampean
grasslands (B. penicillata has also been collected in north-
eastern Patagonia) and therefore suffer from the pressure of
land conversion to agriculture, urban sprawl, mining and
extractivism by hobbyists. In the case of B. biplumata and
B. polysyka all these factors are worsened by their restricted
distribution. It is urgent to evaluate the conservation status
of these species (preferably, under the IUCN criteria) as well
as to detect the eventual risks they are exposed to. Addi-
tional studies addressing these issues are on the way.

Acknowledgements – All the personnel of the consulted herbaria are

greatly thanked for their valuable assistance on our work. We thank
The Rufford Foundation and Neotropical Grassland Conservancy
for fieldwork funding. AS acknowledges Consejo Nacional de
Investigaciones Científicas y Técnicas (CONICET) for his doctoral
scholarship and thanks Ideawild for donating field equipment.
Figure 4. Map of the distribution of Bipinnula species recorded CRB and RBS thank Conselho Nacional de Desenvolvimento
from Argentina. Gray: B. biplumata. Black: B. polysyka. Horizontal Científico e Tecnológico (CNPq, projects Reflora 563556/2010–2
lines: B. penicillata. and Universal 470353/2011–2) for financial support; CRB is grate-
ful for Postdoctoral Fellowship (CNPq, project 150075/2014–6).
tunately, Kraenzlin kept fragments for his study and this
private collection was acquired by HBG in 1935 and thus References
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