Phytotaxa 521 (3): 177–192 ISSN 1179-3155 (print edition)

https://www.mapress.com/j/pt/
Article PHYTOTAXA
Copyright © 2021 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.521.3.3

Prosopanche cocuccii (Hydnoraceae): a new species from Southern Brazil

JORDANO D. TAVARES DE CARVALHO1,5*, CATARINE MARKUS2,6, ALDO MEROTTO JUNIOR2,7, RENATO
A. ZÁCHIA3,8, MARILIA SCHUCH4,9, ESTÉFANI SULZBACH2,10 & JORGE E. A. MARIATH1,11
1
Laboratório de Anatomia Vegetal (LAVeg), Programa de Pós-Graduação em Botânica, Departamento de Botânica, Instituto de Bio-
ciências, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves, 9500, Porto Alegre, RS, Brazil.
2
Departamento de Plantas de Lavoura, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves, 7712, Porto Alegre, RS,
Brazil.
3
Departamento de Biologia, Universidade Federal de Santa Maria, Av. Roraima, 1000, Santa Maria, RS, Brazil.
4
JTI Processadora de Tabaco do Brasil LTDA, Santa Cruz do Sul, RS, Brazil.
5

jordano.tavares@ufrgs.br, https://orcid.org/0000-0002-2423-2186
6

catarine.markus@ufrgs.br, https://orcid.org/0000-0002-5330-3502
7

merotto@ufrgs.br, https://orcid.org/0000-0002-1581-0669
8

renato.zachia@ufsm.br, https://orcid.org/0000-0003-2842-6129
9

Marilia.Schuch@jti.com, https://orcid.org/0000-0002-3254-0242
10

estefanisulzbach@gmail.com, https://orcid.org/0000-0002-8699-6640
11

jorge.mariath@ufrgs.br, https://orcid.org/0000-0002-2776-6781
*
Author for correspondence: Jordano D. Tavares de Carvalho,
jordano.tavares@ufrgs.br

Abstract

Prosopanche cocuccii is described as a new species from Southern Brazil. It resembles P. bonacinae due to the trigonous
rhizome and by the general size of the flower organs. Nevertheless, P. cocuccii is distinguished by the main floral morpho-
logical characteristics used to differentiate species. In addition, the new species presents some rare characters for the genus,
such as a highly branched rhizome and flowers usually grouped in fascicles. We provide a description, illustrations, field
photographs, morphological details under the stereomicroscope, and comments on habitat, ecology, distribution, and con-
servation of the new species, as well as an updated key for the genus Prosopanche. In addition, we identified that P. cocuc-
cii produces a large number of rhizomes and fruits for each single host specimen of tobacco (Nicotiana tabacum), which
resulted in a decrease in height and growth of its only known host plant.

Keywords: Atlantic Forest, haustorial roots, Hydnora, Nicotiana tabacum, Prosopanche bonacinae, Piperales, root hol-
oparasite.

Resumo

Prosopanche cocuccii é proposta como uma espécie nova do sul do Brasil. Assemelha-se a P. bonacinae pelo rizoma trígono
e tamanho geral das peças florais. Contudo, P. cocuccii se distingue pelas principais características florais utilizadas para
diferenciar espécies. Além disso, a nova espécie apresenta algumas características raras para o gênero, como o rizoma alta-
mente ramificado e flores geralmente agrupadas em fascículos. Apresentamos uma descrição, uma ilustração, fotos a campo,
detalhes morfológicos sob estereomicroscópio e comentários sobre o habitat, ecologia, distribuição e conservação da nova
espécie, como também uma chave atualizada para o gênero Prosopanche. Além disso, identificamos que P. cocuccii produz
um grande número de rizomas e frutos para cada espécime de tabaco (Nicotiana tabacum) hospedeiro, o que resulta em um
decréscimo na altura e crescimento de sua única planta hospedeira conhecida.

Palavras-chave: Floresta Atlântica, holoparasita de raízes, Hydnora, Nicotiana tabacum, Piperales, Prosopanche bonaci-
nae, raízes haustoriais.

Introduction

Hydnoraceae C. Agardh comprises root holoparasitic plants with a remarkable specialization of vegetative organs and
very distinctive flower morphology. After many divergences (Nickrent et al. 2002; Barkman et al. 2007; Naumann
Accepted by Marie-Stéphanie Samain: 10 Sept.2021; published: 30 Sept. 2021 177
et al. 2013; Massoni et al. 2014, APG IV 2016), Jost et al. (2021) recognize a series of morphological and molecular
evidences that place the family among the perianth-bearing Piperales, together with other three monophyletic families:
Aristolochiaceae Jussieu, Asaraceae Ventenat and Lactoridaceae Engler. Hydnoraceae thus includes two genera:
Hydnora Thunberg (1775: 69), with eight species distributed in Africa and Arabic Peninsula (Martel et al. 2018), and
Prosopanche de Bary (1868: 267), with six species occurring in South America and Costa Rica (Funez et al. 2019).
Prosopanche differs from Hydnora mainly by the fusion of the polythecous anthers and by the presence of
interstaminal appendages, often interpreted as staminodes (Cocucci 1965). Given the modified flower morphology,
with few taxonomically informative characters, most of the first species of Prosopanche were described using artificial
characters, often based on few fertile materials. In that context, Cocucci (1965) proposed a complete morphological/
nomenclatural and taxonomic revision for the genus, recognizing only two species, Prosopanche americana (Brown
1845: 245) Baillon (1886: 27) and P. bonacinae Spegazzini (1898: 19–22) (the correct spelling for the epithet “bonacinai”;
see below in Nomenclatural note), distributed throughout the Chaco domain in Argentina, Bolivia, Paraguay, and
possibly Southern Brazil. In this study, Cocucci (1965) delimited the taxonomically informative characters used later
in the description of new species, such as the number of rhizome angles, measurements of floral organs, number of
folds in the stigmatic lamellae, morphology and measurements of staminodes, and seed morphology. Gómez & Gómez
(1981: 53–54) later described P. costaricensis Gómez & Gómez from the rainforest of Costa Rica. Three decades
later, Machado & Queiroz (2012) described P. caatingicola R.F.Machado & L.P.Queiroz (2012: 59) from the Caatinga
domain, a Brazilian nucleus of dry forest. More recently, Martel et al. (2018) described P. panguanensis C.Martel
& Rob Fernandez (2018: 242) from the rainforest of central Peru, also expanding the morphological description of
P. costaricensis. Finally, Funez et al. (2019) described P. demogorgoni Funez (2019: 94) from the Atlantic Forest of
Southern Brazil.
The first known mention for Prosopanche in Brazil is Harms (1935), based on a specimen collected by Jürgens
parasitizing tobacco in an uncertain location in the state of Rio Grande do Sul. Later, some collections identified as
P. bonacinae from the same State were also observed, unfortunately lacking well-preserved botanical material (e.g.,
SMDB 998). At the end of 2020, infestations of a species of Prosopanche were found in cultivated tobacco fields near
Atlantic Forest remnants on the southern slope of the Southern Brazilian plateau, in the state of Rio Grande do Sul.
Previous analyzes indicated an affinity of this species with P. bonacinae, but detailed morphological observations
revealed that it represents, indeed, a new species.

Material and Methods

Plant specimens (ca. 8) were collected in January 2021 and February 2021 with ca. 30 flowers at pre- and post-anthesis,
rhizomes and some fruits. The fresh material was measured and morphologically examined. Part of the material was
dried and part was fixed in FAA 50% and stored at the spirit collection of the Laboratório de Anatomia Vegetal (LAVeg)
at Universidade Federal do Rio Grande do Sul (UFRGS). Vouchers from the type material were deposited at ICN
herbarium (UFRGS), and isotypes were distributed to the herbaria RB and SMDB. Herbarium vouchers from SMDB
and virtual vouchers from BM, F, HUEFS, MBM, MO, NY, and SI herbaria were also examined, as well as characters
illustrated in the protologues of most species. We also carefully examined the taxonomic and morphological revisions
proposed by Cocucci (1965) and Cocucci & Cocucci (1996) for P. bonacinae. The distribution map was elaborated
using the QGIS 3.10 ‘A Coruña’ software (2019), with topological data from the Topodata project (Geomorphometric
Database of Brazil-INPE) that offers the Digital Elevation Model (MDE), data from IBGE for the administrative
limits, and Google Earth images. The conservation assessment complies with the IUCN criteria (2017).

Taxonomic treatment

Prosopanche cocuccii Tavares de Carvalho, Záchia & Mariath sp. nov. (Fig. 1–8)
Prosopanche bonacinae affinis, qua imprimis differt numero thecarum cuiusque antherae (9–13 vs. 20–30), numero plicarum in lamellis
stigmatis (10–30 vs. 4–10), forma staminodiorum (sessilia, reflexa vs. pedunculata) et dimensionibus ovari (25–70 × 23–34 mm vs.
15–20 × 10–20 mm). Etiam differt rhizoma ramoso (vs. rhizoma raro diviso).
TYPE:—BRAZIL. Rio Grande do Sul: Agudo, parasitando tabaco, 29°31’07.7”S, 53°13’18.8”W, 12 January 2021, J.D.T. de Carvalho
112 (holotype ICN!; isotypes RB!, SMDB!).

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Herb, root holoparasite, achlorophyllous, hypogeal except for the upper portion of the flower. Rhizomes trigonous, rarely
tetragonous, fleshy, with numerous branches; primary rhizome 6–13 mm diam., branches 2–5.5 mm diam.; haustorial
rudiments numerous on the angles, 0.5–1.5 mm apart, 0.5–2 mm long. Leaves absent. Bracts absent. Flowers in 2–6-
flowered fascicles or isolated on the fascicle’s periphery, monoclinous, semi-hypogeal (emergent at the perigonial
tube’s height); pedicel 17–128 × 17–35 mm; perigonial tube 10–60 × 12–20 mm; tepals (2–)3, valvate, ovate to
oblong-ovate, fleshy, apex acuminate, externally rugose, ochre to light brown, at maturity internally smooth, dark
brown; synandrium 12–20 × 7–16 mm, ovoid, pale yellow before dehiscence, becoming dark brown after dehiscence,
stamens 3, filaments 1–3 × 2–2.5 mm, anthers 9–13-thecous, extrorse, with longitudinal dehiscence; staminodes 3,
alternate to the filaments, ca. 5–6 mm wide, sessile, reflexed, bilobed, lobes fleshy, rugose, appendiculate; ovary
inferior 25–70 ×23–34 mm, ovoid to obovoid, hypogeal; stigmatic surface triradiate, lamellate, 9–12 lamellae per side,
smallest ones with 10–12 folds, the largest with 25–30 folds; ovules many, reduced, embedded in a laminar, parietal
placenta. Fruit ca. 70 mm × 50 mm, globose to ovoid, hypogeal, fleshy, rugose, dehiscing by irregular longitudinal
fissures. Seeds numerous, 5–7 mm wide, ovoid, foveolate, dark brown.
Additional specimens studied:—BRAZIL. Rio Grande do Sul: Agudo, parasitando tabaco, 29°28’27.7”S,
53°10’30.7”W, 12 January 2021, J.D.T. de Carvalho 113 (ICN); Gramado Xavier, parasitando tabaco, 29°22’48.7”S,
52°33’12.0”W, 24 February 2021, J.D.T. de Carvalho 114 (ICN).
Etymology:—The specific epithet honors Alfredo Elio Cocucci (1926-2015), a prolific professor, taxonomist and
plant embryologist from Argentina, for his taxonomic, morphological and embryological contributions to Prosopanche
and many other angiosperm groups.
Phenology:—Flowers of P. cocuccii were collected between November and March and fruits between January
and April.
Distribution, habitat and ecology:—Prosopanche cocuccii is known from three localities in the central region
of Rio Grande do Sul, Southern Brazil, in the municipalities of Agudo and Gramado Xavier (Fig. 2). This region’s
irregular topography results from the erosion of the edge of the Southern Brazilian plateau, with a mosaic of forest
elements (Seasonal Forest, Atlantic Forest) and agricultural properties, mainly tobacco crops (Fig. 3). A probable
occurrence of the species in the municipality of Arroio do Tigre is considered dubious, since we were only able to
analyze photos.
The only known host plant is tobacco, Nicotiana tabacum Linnaeus (1832: 462), in all localities. Field search
in the native surrounding vegetation did not lead to the collection of other individuals of P. cocuccii, and no other
hosts were found in the tobacco field. However, there is a great possibility that its origin is from the vegetation
near the tobacco fields, requiring a more active search for the original host. Another hypothesis to consider is that P.
cocuccii has changed its original host, although it is unknown how this plant would maintain its life cycle in this case,
since tobacco is an annual crop. Nevertheless, the morphological characteristics are identical in all populations. The
individuals apparently develop normally, with rhizomes and flowers observed in different developmental stages and
fruits producing abundant seeds (Fig. 4–8).
When comparing the host plants of Prosopanche species (Tab. 1), there is a preference for Fabaceae Lindley
species, with exception of P. demogorgoni, which parasitizes Baccharis uncinella De Candolle (1836) (Asteraceae
Berchtold & J.Presl) and, notably, P. bonacinae, which apparently has no host preference, parasitizing at least 15 species
from seven families (Amaranthaceae Jussieu, Anacardiaceae R.Brown, Apiaceae Lindley, Aquifoliaceae Berchtold
& J.Presl, Asteraceae, Malvaceae Jussieu, and Solanaceae Jussieu) (Table 1). This species has been occasionally
associated with at least three crops of agronomic or commercial importance (Cocucci 1965), including Gossypium
hirsutum Linnaeus (1763: 975) (cotton), Ilex paraguariensis A. Saint-Hilaire (1822: 351) (mate), and Foeniculum
vulgare Miller (1768) (fennel). However, there are few data on the parasitism characteristics in these species. Although
the damage in tobacco was not directly quantified, we identified that tobacco plants infested with P. cocuccii had
lower growth in comparison with unattached plants. This effect occurred before the parasite emerges from the soil,
which makes it difficult to diagnose infestations in a large area because the removal of all plants is not possible. A
large number of rhizomes with fruits parasitizing a unique tobacco plant was identified (Fig. 3C). This results in large
demand for photoassimilates obtained by the haustorial roots that limit the growth of the host plant (Fig. 3C).
The flowers at anthesis exude a pleasant minty odor, resembling the scent of the roots of Polygala Linnaeus
(1753: 701). After anthesis, Coleoptera and Diptera were found in abundance, inside the perigonial tube and sometimes
in galleries into the ovary. The mature fruit has a slightly sweet flavor and develops a putrid odor, with larvae and
adult beetles and ants feeding on the fleshy placenta. The fruit opens through irregular fissures and does not seem to
naturally reach the surface, being probably dug up by larger animals (Fig. 8).

PROSOPANCHE COCUCCII (HYDNORACEAE) Phytotaxa 521 (3) © 2021 Magnolia Press • 179
FIGURE 1. Line drawings of Prosopanche cocuccii. A. Detail of the tepals and synandrium. B. Stigma surface. C. Detail of a staminode
in frontal view. D. Upper view of the staminodes E. Synandrium in bottom view showing the staminodes. F. Seed. G. Habit (the hatched
line indicates the height of the soil surface). H. Portion of the rhizome with a cluster of flower buds. I. Host root and rhizome with two
flower buds on the left. (Illustrated by Rafaella Migliavacca Marchioretto).

180 • Phytotaxa 521 (3) © 2021 Magnolia Press DE CARVALHO ET AL.

FIGURE 2. Known distribution for Prosopanche cocuccii in Rio Grande do Sul, Brazil.

PROSOPANCHE COCUCCII (HYDNORACEAE) Phytotaxa 521 (3) © 2021 Magnolia Press • 181
FIGURE 3. A. Habitat of Prosopanche cocuccii, showing the property near the type locality with Atlantic Forest remnants in the
background. B. Tobacco field in the type locality. Arrow indicates a location with parasitized plants. C. Detail of a plant with severe
damage due to parasitism (arrow). To show the parasite, part of the roots was exposed. Arrowheads indicate some of the many flowers of
the parasitic plant.

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FIGURE 4. Photographs of Prosopanche cocuccii. A. Two flowers at anthesis in upper view. B. Fascicle of flowers of the same individual,
after partial removal of the soil (arrows: flower buds or fruits. C–E. Emergent portion of flowers. C. Flower at early anthesis. D. Flower
during anther dehiscence. E. Flower at post-anthesis. Bars: 1 cm.

PROSOPANCHE COCUCCII (HYDNORACEAE) Phytotaxa 521 (3) © 2021 Magnolia Press • 183
FIGURE 5. Underground structures of Prosopanche cocuccii. A. Fascicle of flower buds at a portion of the rhizome, showing a fascicle
on the left and solitary flowers on the right. B. Flower at anthesis next to a developing flower bud. C. Rhizome fragment and host root
(arrow), showing some of its branches (detail: rhizome in cross-section). Bars: A–C. 1 cm. Detail in C: 5 mm.

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FIGURE 6. Flower morphology of Prosopanche cocuccii. A–D. Synandrium in frontal view of one of the anthers. A. Synandrium at pre-
anthesis. B. Synandrium at very early anthesis (arrows: insertion point of the filament). C. Synandrium at anthesis, with dehiscent thecae.
D–F. Stigma in upper view. D. Fresh stigma at pre-anthesis, with a mucilaginous substance in the surface (sl: stigmatic lamella). E. Dried
stigma at pre-anthesis (sl: stigmatic lamella). F. Detail of the stigmatic lamellae (arrow: fold in a stigmatic lamella). G. Ovarian region in
cross-section, showing the laminar, parietal placenta in the center, and channels with a mucilaginous substance in the periphery. H. Detail
of a lamella in the placenta showing many ovules (arrow: ovule). Bars: A. 1.6 mm. B–F: 1 mm.

PROSOPANCHE COCUCCII (HYDNORACEAE) Phytotaxa 521 (3) © 2021 Magnolia Press • 185
FIGURE 7. Staminode morphology of Prosopanche cocuccii. A–D. Sections of fresh flowers at pre-anthesis. A. Longitudinal section of
the synandrium and the perigone tube with a frontal view of a staminode. B. Cross section in perigonial tube, showing a bottom view of
the synandrium and staminodes. C. Staminodes in upper view. D. Detail of a staminode in frontal view. E–F. Cross-sections of perigonial
tube of dried flowers at pre-anthesis. E. Bottom view. F. Upper view. (pt: perigonial tube; sy: synandrium. Arrow indicates the appendage
in a staminode lobe. Bars: A–C. 1 mm. D–E. 2 mm. F. 0.6 mm. G. 5 mm. H. 0.5 mm.

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FIGURE 8. Fruits and seeds of Prosopanche cocuccii. A. Immature fruits. On the right, a fruit just after the emergent portion of the flower
falls. The soil was partially removed. B. Mature fruit, showing the irregularly longitudinal dehiscence. C. Upper view of a dehiscent fruit
being visited by ants, showing the fleshy pulp and numerous seeds (arrow: ant). D. Dried seed. Bar: 0.1 mm.

TABLE 1. Comparison between P. cocuccii and other species of Prosopanche.

P. cocuccii P. americana P. bonacinae P. caatingicola P. costaricensis P. demogorgoni P. panguanensis
Rhizome 3(–4)-gonous (4–)5(–6)- 3(–4)-gonous 3(–4)-gonous (4)–5-gonous 3(–4)-gonous 3–5-gonous
cross section gonous
Rhizome Numerous Rare Rare Not reported Not reported Not reported Not reported
branches
Flowers In fascicles/ Solitary Solitary Solitary In fascicles Solitary Solitary
solitary
Tepal (mm) 35–45(–60) × 60–70 × 35–55 × 9–20 18–22 × 4–14 32–56 × 24–36 15–20 × 5–8 14.2–26 ×
13–25 30–50 11.4–21
Synandrium 12–20 × 7–16 30–40 × 15–20 × 6–8 9–9.5 × 4–5 19–26 × 5–6 × 3–4 13.6–22 × 11–14
(mm) 23–25 11–15,7
Staminodes Sessile, Sessile, Pedunculate, bilobed at Sessile, bilobed. Sessile, Sessile, bilobed Sessile, unilobed,
bilobed. bilobed. apex. Rugose lobes. Oblongoid bilobed. at apex. cucculate and
Fleshy- Globose, fleshy lobes. Smooth fleshy Papillose lobes. fleshy.
appendiculate smooth, lobes.
lobes. fleshy lobes.
Thecae 9–13 20–30(?) 20–30 5–8 7–17 3–4 8–11
Ovary (mm) 25–70 × 50–80 × ? 15–20 × 10–20 15–20 × 12–17 15–20 × ? 15–23 × 12–15 17.1–27.1 ×
23–34 19.6–22.4
......continued on the next page

PROSOPANCHE COCUCCII (HYDNORACEAE) Phytotaxa 521 (3) © 2021 Magnolia Press • 187
 TABLE 1. (Continued)
P. cocuccii P. americana P. bonacinae P. caatingicola P. costaricensis P. demogorgoni P. panguanensis
Folds in the 10–12 14 4 (smallest)–10 ? 12 ? 4–20
stigmatic (smallest)– (smallest)– (biggest)
lamellae 25–30 30 (biggest)
(biggest)

Seed Oval, Oval to Spheroidal, foveolate, ? Elliptc, ? Rugulose

foveolate, elliptic, 5 mm wide rugulose-
0,5–0,8 mm foveolate, verrucose,
wide 1 mm wide 1–1.2 mm
wide
Host plant Solanaceae: Fabaceae: Amaranthaceae: Fabaceae: Fabaceae: Asteraceae: Fabaceae:
Nicotiana Prosopis Salicornia L. Parapiptadenia Inga Baccharis Inga Miller
tabacum Linnaeus (1753); oerstediana uncinella (1754), Abarema
zehntneri
(1767) Benth. ex Pittier (1927),
Anacardiaceae: (Harms 1924)
Schinus dependens Lima & Lima Seemann Zygia P. Browne
Ortega (1798); (1984), Mimosa (1853), (1756)
ophthalmocentra Gliricidia
Apiaceae: Foeniculum
Martius ex sepium (Jacq.)
vulgare
Bentham (1875), Kunth ex
Aquifoliaceae: Ilex
Calliandra Walpers (1842)
paraguariensis
depauperata
Asteraceae: Ambrosia
Bentham (1875)
tenuifolia Spreng.
(1826), Baccharis
salicifolia (Ruiz
& Pav.) Persoon
(1807), Gaillardia
megapotamica
(Spreng.) Baker (1884),
Hyalis argentea D.
Don ex Hooker &
Arnot (1835), Proustia
ilicifolia Hooker &
Arnot (1830)
Malvaceae: Sida
arborea Linnaeus f.
(1782), Gossypium
hirsutum; G.
barbadense Linnaeus
(1753);
Solanaceae: Solanum
auriculatum Aiton
(1789), Salpichroa
origanifolia (Lam.)
Thellung (1912),
Fabiana Ruiz & Pavón
(1794)

Conservation status:—Prosopanche cocuccii is known from three localities where it seems to be abundant. The
number of individuals cannot be estimated, added to the fact that it has not been found in native vegetation. Thus, we
suggest it be assessed as DD (Deficient Data), according to the IUCN categories and criteria (IUCN 2017).
Nomenclatural note on P. bonacinae:—We adopt the spelling “bonacinae” instead of “bonacinai” since
it is grammatically correct, as recommended by Cocucci (1965). Machado & Queiroz (2012) justified the use of
“bonacinai” for this species since “Spegazzini (1898) named this plant after the priest Pedro Bonacina (Giacomelli
1906), and therefore the specific epithet should be in the genitive case retaining its gender in accordance with Article
23.5 (ICBN; McNeill et al. 2006)”. Since then, this spelling has been used in official species lists (e.g., Flora do Brasil
2020; tropicos.org) and descriptions of new species (Martel et al. 2018; Funez et al. 2019). However, it is not possible

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to maintain the termination -i, if the name in question ends with the vowel -a, even if the honored person is male. In this
case, the genitive must be indicated with the ending -e, in the singular, or -erum in the plural (ICN, Art. 60.8; Turland
et al. 2018).
Taxonomic comments:—This species resembles P. bonacinae, by the stems usually trigonous in cross-section
and by the flower measurements, except by the ovary dimensions (Table 1). It differs, however, by the main flower
characters proposed by Cocucci (1964) in the morphological concept of P. bonacinae. Prosopanche cocuccii has fewer
thecae in each anther (9-13 vs. 15-30) and more folds in the stigmatic lamellae (10-12 in the smallest and 25-30 in the
largest vs. ca. 4 in the smallest and 10 in the largest) (Fig. 6). Despite being bilobed, the staminodes are sessile and
reflexed in P. cocuccii (Fig. 7) (vs. short pedunculate in P. bonacinae). It also differs, albeit subtly, by seed morphology
(Fig. 8 D) (ovoid in P. cocuccii vs. spheroidal in P. bonacinae).
The large number of second-order branches of the primary rhizome, usually with a smaller diameter, is a remarkable
vegetative feature of P. cocuccii (Figs. 1 I; 5 C). These branches can give rise to flowers or to third-order branches,
which are even thinner. Regardless of the branching order, the rhizome maintains its trigonous shape in cross-section
(Fig. 5 C, detail). According to Cocucci & Cocucci (1996), the rhizomes of P. americana and P. bonacinae rarely
present branches, with only a few dichotomies. Likewise, for the remaining species of this genus, there are no reports
in protologues or type materials that indicate ramifications in the primary rhizome.
The flower insertion in the rhizome is another crucial diagnostic character of P. cocuccii. Prosopanche species
have solitary flowers, with P. costaricensis being the only species with flowers grouped in fascicles. In P. cocuccii,
the individuals exhibited clusters of flowers, with at least one flower group in a fascicle, although solitary flowers
on the periphery of the rhizome also occur (Figs. 1 H; 4 A–B; 5 A–B). Flowers in fascicles seem to be one of the
only characters shared with the Costa Rica endemic P. costaricensis, which shows morphological affinities with P.
americana and P. panguanensis.
Floral organ measurements are important in delimiting most species of Prosopanche. In P. cocuccii, however, some
measurements of synandrium, tepals and perigonial tube individually overlap when compared to most of Prosopanche
species, except by the dimensions of the ovary (Table 1). Even at pre-anthesis flowers, the ovary is larger than most
species of Prosopanche with developed fruits. Only P. americana, with the largest flower of the genus, presents a
larger ovary. Similar to other species of the genus, pedicels and ovaries of P. cocuccii have channels that store a yellow
mucilaginous substance. This substance can be observed when the tissue of that region (mucilage channels, sensu
Cocucci & Cocucci 1996) is ruptured (Fig. 6 G).
Additional specimens of Prosopanche examined:
Prosopanche americana:—ARGENTINA. San Juan: Quebrada del Carrizal, A. Lutz 1119 (MBM 148039); Pampas Reise (=Pampas
trip): January 1892, C. E. O. Kuntze s.n. (NY 887985); San Luis: Quines, Estancia La Cortadera, 2 January 1944, R. Báez, s. n.
(SI071061).
Prosopanche bonacinae:—BRAZIL. Rio Grande do Sul: Faxinal do Soturno, Santos Anjos, 30 January 1967, I. Cervo s.n. (SMDB
998!).
Prosopanche caatingicola:—BRAZIL. Bahia: São Gabriel, Alto da Jurema, 11º11’42”S, 41º50’29”W, 5 April 2009, R.F. Machado 652
(HUEFS 187671, holotype); São Gabriel, Alto da Jurema, 11º11’42”S, 41º50’29”W, 10 August 2010, R.F. Machado 653 (HUEFS
187672, paratype).
Prosopanche costaricensis:—COSTA RICA. Cartago: Callejón section 18, finca La Lola, 11 June 1981, L.D. Gómez 7335 (BM000947287,
isotype).
Prosopanche panguanensis:—PERU. Río Tambo, Comunidad Nativa Parijaro, camino al Parque Nacional Otishi, 11º58’10”S,
73º40’19”W, 15 November 2013, L. Valenzuela et al. 26751 (MO 6717522).

Key to species of Prosopanche

1. Staminodes cucullate, perigonial tube extending above the synandrium insertion .......P. panguanensis C.Martel & Rob.Fernandez
- Staminodes bilobed, perigonial tube not extending above the synandrium insertion ........................................................................2
2. Rhizome (4–)5(–6)-gonous ................................................................................................................................................................3
- Rhizome 3(–4)-gonous .......................................................................................................................................................................4
3. Flowers solitary; rhizome (4–)5(–6)-gonous; synandrium 30–40 × 23–25 mm; tepals 60–70 × 30–50 mm; South America ............
................................................................................................................................................................P. americana (R.Br.) Baillon
- Flowers in fascicles; rhizome (4–)5-gonous; synandrium 19–26 × 11–16 mm; tepals 32–56 × 24–36 mm; Costa Rica ....................
....................................................................................................................... Prosopanche costaricensis L.D. Gómez & Gómez-L.
4. Anthers with 3–8 thecae; synandrium 5–9.5 × 3–5 mm; tepals 15–22 × 5–14 mm ...........................................................................5
- Anthers with 9–30 thecae; synandrium 12–20 × 6–16 mm; tepals 35–60 × 9–25 mm ......................................................................6

PROSOPANCHE COCUCCII (HYDNORACEAE) Phytotaxa 521 (3) © 2021 Magnolia Press • 189
5. Anthers with 5–8 thecae; synandrium 9–9.5 × 4–5 mm; perigonial tube 5–10 mm long ....................................................................
....................................................................................................................................... P. caatingicola R.F.Machado & L.P.Queiroz
- Anthers with 3–4 thecae, synandrium 5–6 × 3–4 mm; perigonial tube 36–85 mm long ................................ P. demogorgoni Funez
6. Flowers solitary; rhizome often unbranched; ovary 15–20 × 10–20 mm; anthers with 15–30 thecae, staminodes short-pedunculate;
4–10 folds per stigmatic lamella; seeds spheroidal, ca. 0.5 mm wide...........................................................P. bonacinae Spegazzini
- Flower often in fascicles; rhizome often many-branched; ovary 25–70 × 23–34 mm; anthers with 9–13 thecae; staminodes sessile,
reflexed; 10–30 folds per stigmatic lamella; seed ovoid, 0.5–0.8 mm wide ........................................................................................
............................................................................................................................ P. cocuccii Tavares de Carvalho, Záchia & Mariath

Acknowledgments

This research was supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), grants
numbers 141695/2019-6 (J.D.T.C.) and 303840/2019-6 (J.E.A.M.).
We thank Laboratório de Anatomia Vegetal (LAVeg) from Universidade Federal do Rio Grande do Sul for technical
support.
We thank the tobacco growers that provided access to the sampling areas and information about the region and
crop management (Silvio Leanel Ceconet, Solange Bindani Ceconet, Dionatan Festinalli and Tania Hoffmann) and
JTI Processadora de Tabaco do Brasil LTDA team for their field assistance: Erick Rizzetti, Fabiano Borth and Jesum
Felipe Lemos (Agronomy Technicians); Catiusca Peretto and Tiago Schuck (Agronomy Development Technicians)
and Roberto José Huff (Plant Breeder Assistant).
We thank the three anonymous reviewers for their contribution to the final version of the article.
We also thank Rafaella Migliavacca Marchioretto for the illustration and Henrique Mallmann Büneker for some
photographs in Fig. 5.

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